BULLETIN ieee
OF THE
VOL. 49
FouNDED BY WILLIAM HENRY LEGGETT 1870
EDITOR
ALEXANDER WILLIAM EVANS
ASSOCIATE EDITORS
JEAN BRoapHuRST MICHAEL LEVINE
JAMEs ARTHUR Harris GEORGE EDWARD NICHOLS
GEORGE T. HASTINGS ARLOW BuRDETTE STOUT
MARSHALL Avery HowE NORMAN TAYLOR eo
NN
@
NEW YORK
1922
PouBLISHED FOR THE CLUB
Tue INTELLIGENCER PRINTING COMPANY
Lancaster, PreNNSYLVANIA
CONTENTS
—— — }. Epiphyllous plants of certain regions in Jamaica
So Pe pb ae Pe a ey ee era ete ss Cea I
Bunn. ‘Bas ys he penne Carmenere 20 ele ae te ee 5 15
Muwnz, Parie A., and JoHNston, IvAN M. Miscellaneous notes on
plants of Sotitheri-Calioraia foo oe ve oe es 31
Farr, CirpporD H. Quadripartition by furrowing in Sisyrinchium
(PRC) i er eo els Pee ose vee 51
ROUND, Epa: Ma Avmodern plant fossil see eo Sees ee ae 63
Hazen, Tracy E. The phylogeny of the genus Brachiomonas (plates
Boy a oe ee Pe ea Be 75
STEVENS, O. A. New records and other notes on North Dakota plants. 93
YuncKER, T. G. Three new species of Cuscuta from Mexico......... 107
Hazen, Tracy E, New British and American species of Lobomonas: a
study in morphogenesis of motile algae (plates 5, 6).............. 123
ANDERSON, FLora. The development of the flower and embryogeny of
: Marttyaia louisiana (plates 7,8) c6e0 ss a ed es 141
OvERHOLTs,*-L. O. Mycological notes for 1920 (plate 9).............- 163
GrarF, Pau W. shake maha plants from Glacier National Park...... 175
OsTERHOUT, GEO. E. Two new plants from Colorado................ 183
ARTHUR, JOSEPH CHARLES. peta species of Uredineae—XIV......... 189
SMITH, CHARLES Piper. Studies in the genus Lupinus—VII. L. suc-
CUIGHEe a BVetas oo ae i kk ee oe 197
Howe, Marswatt A., and Hoxiicx, ArtHuR. A new American fossil
aU See ee ey evs bee Guill es oo arc ee aoe 207
CocKkERELL, T. D. A. A new genus of fossil Liliaceae..............-+- 211
Grarr, PauL W. Philippine Basidiomycetes—V.................-+- 223
DospaLL, LourisE. Occurrence of the pyncial stage of Puccinia Taraxaci 235
BARTRAM, Epwin B. Midwinter botanizing in southern Arizona...... 237
Britton, N. L., and Ross, J. N. Two new genera of Cactaceae...... 251
Ruspy, H. H. New species of trees of medical interest from Bolivia... 259
AsHE, W. W. Notes on trees and shrubs of the southeastern United
ies p C. Commelinantia, a new genus of Commelinaceae (plates
Oo ei eek ON Rad be ne ee Uaik epee Ue ee oh Ge es 269
ae W. W. The genus Costus in Central America (plates 12-15).. 283
SHIMEK, B. Quercus Lge m Jowd’ (plates 16, 17) 2000. eset as 293
PORTERPIELD, W. M., References to the algae in the Chinese classics. 297
Dopce, B.O. A Flam with a botryose conidial stage............. 301
ili
iv DATES OF PUBLICATION
Doran, W.L. Effect of external and internal factors on the germination
GE TUNGOUS SPOTES rye a GS Fe LS eee ree ee S13
Mouwnz, Pamir A., and JoHNsToN, IvAN M. Miscellaneous notes on
plants of Southert Cairne sce ik ee os ve ae ee oe 349
MACKENZIE, KENNETH Kent. Notes on Carex—XII..............4.. 361
OLDENBUSH, CARRIE. Stimulation of plants by carbon disulphide... 375
INDEX TO AMERICAN BOTANICAL LITERATURE, 23, 45, 65, III, 159, 185, 215,
253. 277) 397, 341, 391
INDEX. TO VOLUME 403060) o5:c. G8 oped oe ee ee 399
Dates of Publications
No. 1, tor January. Pages I- 30. Issued March 21, 1922.
No. 2, for February. 3I- 50. April 4, 1922.
No. 3, for March 51- 74 April 12, 1922
No. 4, for April 75-122 May 18, 1922
No. 5, for May 123-162 June 15, 1922
No. 6, for June 163-188 July 8, 1922
No. 7, for July 189-222 August 10, 1922
No. 8, for August 223-258 August 31, 1922
No. 9, for September. 259-282. September 22, 1922.
No. 10, for October. 283-312. October 11, 1922.
No, 11, for November. 313-348. November 9, 1922.
No. 12, for December. 349-408. January 27, 1923.
Errata
Page 2, line 8, for “‘arboreum”’ read “arborum.’
Page 6, footnote, for “ Caraguta”’ read ‘ ‘ Caraguata.”
PLATE 5”’’ read ‘' PLATE I.
Page 14, line 2 from bottom, for ‘ oy read ‘ aoe
Page 32, line 11 from bottom, for ‘‘muticus”’ read ‘mutica.
Page 38, line 13 from bottom, for penasseee> read asin. eo
Page 76, line 1, for “‘gareement” read “agreement.”
Page 77, line 12 from bottom, for “genera” read ‘‘general.”
Page 92, line 4, for “ertracted”’ read ‘‘retracted.” -
Page 100, line 4, for “ AMsINKIA”’ read ‘ AMSINCRIA.”’
Page 101, line 13, for ‘‘StBERICA”’ read ‘‘SIBIRICA.””
Page 109, line 6, for “ wartius’’ read ‘‘ Martius.”
Page 109, line 10 from bottom, for “‘falsl’’ read ‘‘false.’
Page 125, line 7 from bottom, insert the word “are”’ after ‘‘sure.”
Page 127, line 16, insert a semicolon after “ posterioribus.
ag ee. ee ae
Errata (Continued)
Page 127, line 23, for ‘‘zsooporis”’ read ‘‘zoospor
Page 143, Fig. 7, the lower right poe oe should be “St”
| Page 156, line 12, for “‘1902’’ read “
_ Page 166, lines 1 and 6 from bottom, a “hyalina” tead “hyalinus.”’
Page 167, line 14, for “HYALINA”’ read ‘ HYALINUS.’
Page 180, line 12, for “ PENTSTEMON”’ ee ‘ PENSTEMON,’
Page 189, line 9, for ‘‘ Pentstemon”’ a ‘* Penstemon.”’
Page 195, line 5, for “ Bermudionum bia ' Bermudiana.”
Page 197, line 8 from bottom, omit “ with.’
*)
0g
”
_ Page 224, line 16, for “corrugatus’’ read “‘ gala.
Page 226, line 9, for Bchacasteccqens® read ‘ ‘Pebpaistie”
_ Page 228, line 18, for “MULITPLEX”’ read “ MULTIPLEX.
Page 241, line 16, for “‘ NoTHOLEANA”’ read ‘‘ NOTHOLAENA.”’
Page 242, line 1, for ‘‘MACHROSTACHYA”’ read a
Page 242, line 16, for “‘RINGENS”’ RIGE
Page 244, line 8 from bottom, for Py aioe sunk” read ip. - cepa
Page 245, lines 24 and 26, for ‘‘ PAROSELLA”’ read ‘‘ PAROSELA
Page 247, line 7, for ‘‘CHOLOROTICA”’ read ‘‘CHLOROTICA.
Page 248, line 12 from bottom, for ‘‘' PENTSTEMON”’ read *‘ PENSTEMON. ”’
Page 271, line 22, for “‘spathe’’ read “sheath of leaf.”
Page 271, line 6 from bottom, for ‘‘leaves”’ — a .
Page 275, line 4, for ‘ Tinantia” read “‘ Trades ie
Page 291; line 15 from bottom, sea “herbarium = Cornell University” read
“U.S. National Herbar
Page 292, line 19, for sehen ite ne “‘Oersted.”’
Page 320, lines 5 and 13 from bottom, for “ Entothia”’ read ** Endothia.””
Page 323, line 7, for ‘ Plasmopora”’ read “‘ Plasmopara.
Vol. 49 No. 1
BULLETIN
TORREY BOTANICAL CLUB
JANUARY, 1922
Epiphyllous plants of certain regions in Jamaica*
Louis J. PESSIN
(WITH PLATE I AND ONE TEXT FIGURE)
The great variety and luxuriant growth of the epiphyllous
plants seen in Jamaica in the summer of 1919 led the writer to
study them from two points of view: first, their distribution in
relation to climatic conditions; second, their histological relation
to the leaves bearing them, in order to determine whether these
epiphytes are to any degree parasitic. The study of the distri-
bution and environment of the epiphyllous plants was made in
the field. The relation of the’epiphyll to its supporting leaf was
studied by histological methods in the laboratory.
HISTORICAL §,
Very little is known about epiphyllous plants today, in spite
of the fact that they have long interested botanists. As early as
1875 Berkeley (1) reported the occurrence of epiphyllous lichens
growing on the leaves of the tea plant; some of these apparently
epiphyllous forms showed marked parasitic adaptation. Such
forms were found in abundance in warm moist regions. Two of
the forms observed by Berkeley were regarded as species belonging
to the genus Strigula.
A few years after Berkeley’s paper appeared, Cunningham (2)
reported on an epiphyllous lichen and an epiphyllous alga, My-
coidea parasitica, parasitic on leaves of Mangifera indica, Croton
and Rhododendron.
[The BULLETIN for December (48 : 315-363) was issued February 28, 1922.]
od Botanical Contribution No. 69 from the Johns Hopkins University.
i
2 PEssIN: EpIPHYLLOUS PLANTS OF JAMAICA
Marshall Ward (6) worked on an epiphyllous lichen which he
believed to be the one worked on by Cunningham. He observed
no penetration of the tissues of the supporting leaf by the lichen
and therefore decided that it was not parasitic. He did find,
however, one or more cross walls in the palisade cells of the sup-
porting leaf beneath the epiphylls, which formed three or four
layers of these cells instead of the normally single layer.
Schmidle (5) found species of Trentepohlia arboreum and T.
aurea as well as Phycopeltis microcystis quite common as epi-
phyllous forms in Samoa. Massart (3) found that some orchids
pass their entire existence on a single leaf. He found also that
the epiphyllous floras differ in different regions. Algae and
liverworts are the most common epiphyllous forms, while mosses
acquire the epiphyllous habit only in very humid ravines.
OBSERVATIONS AND METHODS
Specimens of epiphyllous plants were collected in various —
regions in the Blue Mountains of Jamaica. The field observations
were made during June and July, 1919, in the following places:
Cinchona Plantation (5,000 feet), Morce’s Gap (4,934 feet),
New Haven Gap (5,600 feet), Green River Valley (3,000 feet),
Markham Hill (5,800 feet), Vinegar Hill Road (4,500 feet), Clyde
River Valley (4,500 feet) and Blue Mountain Peak (7,428 feet).
Material was either pressed or preserved in glycerine in 4 per
cent formaldehyde, or in 60 per cent alcohol. For careful histo-
logical study, material was fixed in chromoacetic acid and pre-
served in 70 per cent alcohol until finally dehydrated, embedded,
cut and stained with the Flemming’s triple stain. In some cases
material was left unstained for comparison.
The Cinchona Plantation: is situated on a spur running oe.
_ from the Blue Mountains at five thousand feet above sea level.
The temperature varies between the extremes of 48° and 82° F.
The vegetation of the garden about the residence of the Cinchona
Plantation consists largely of introduced plants, many of which
are Himalayan, Australian and Cape of Good Hope species.
There are also native tree ferns, orchids, bromeliads, and many
other transplanted native species. The banks along the trails —
outside the garden are covered mostly by native ferns interspersed
j
PESSIN: EPIPHYLLOUS PLANTS OF JAMAICA 3
with, or on the north side of Cinchona Hill largely replaced by,
Hedychium coronarium.
During the wet season Canchonn is enveloped in clouds for days
at a stretch and the rains are very heavy and frequent. In spite
of this abundance of moisture at certain times of the year, epi-
phyllous plants are rather scarce on the south side of Cinchona
Hill. Only one hepatic, two species of algae and a few kinds of
lichens were found growing there. Of these Trentepohlia grows
in damp or shaded places on the ground, on the bark of trees and
on the leaves of certain plants. Lichens, the most common of
which belong to the genus Sticta, grow abundantly on certain
plants. Some of the leaves of Callistemon lanceolatum, for instance,
are almost completely covered by this lichen. In a few cases the
leaves of Hedychium were found sprinkled with a reddish disk-
like alga belonging to the genus Phycopeltis. The epiphyllous
flora of this region is evidently not a rich one (see TABLE I, A).
At Morce’s Gap, two miles north of Cinchona and at about
the same level, is found a characteristic tropical montane rain
forest. Here moisture is abundant at all seasons of the year. On
the windward side of the mountains everything reeks with mois-
ture; clouds drift constantly through these dense forests and rain
falls frequently during eight months of the year. The dense
vegetation, except on the ridges, is usually dripping wet. Very
rarely do the sun’s rays penetrate to the floor of these forests.
Ferns carpet the floor and cover the walls of the steep-sided
ravines. Tree ferns, such as Alsophila and Cyathea, are very
common, many of them having trunks four or five inches thick and
thirty feet high or more. Species of the genera Marattia, Danaea,
Polypodium and Elaphoglossum are very abundant. Such forms
as Blechnum attenuatum and many species of T rich , Hymeno-
phyllum, Polypodium and Elaphoglossum have been compelled to
adopt either the climbing or the epiphytic habit in the intense
competition for sunlight. Podocarpus, Hedyosmum, species of
Piper and many other dicotyledonous trees and shrubs, together
with numerous herbaceous plants, are scattered through these
woods, increasing the struggle for existence and making the
competition for light more keen.
In such an environment epiphyllous plants are very abundant.
TABLE I
EPIPHYLLOUS PLANTS OF JAMAICA
Supporting plant
Region Algae Lichens Hepaticae Musci
A. Cinchona Plan- | Callistemon lanceolatum| Parmelia sp. Cololejeunea diaphana
—— 5000 ft. alt Sticta sp.
uth exposure. Usnea sp.
; |A form of Pilocar-
| paceae
Hedychium coronarium | Phycopelti Lopadium sp.
Tren secpokife'ss aurea
Pittosporum tobira Phyco peltis sp.
B. Morce’s Gap, | Elaphoglossum latifo- | Desmidium sp. Cololejeunea
4934 ft. alt. north lium Nostoc sp. Colotejeunea diaphana
exposur Phycopeltis sp. Crossotolejeun
oe eg Digatiicens
Scyto ellucida
reeset sp.
Metzgeria furcata
Microlejeunea sp. (?)
Hedychium coronarium
Phycopeltis sp.
Crossotolejeunea sp.
pellucida Schiffn.
Hedyosmum arbor-
escens Sw.
Crossotolejeunea sp.
Lejeunea flava
TABLE I—Continued
Region Supporting plant Algae Lichens Hepaticae Musci
B. Morce’s Gap—| Mangifera indica Phycopeltis sp. Undetermined species*
Continued.
Podocarpus coriaceus Phycopeliis sp. Ph a ag eg sp.
Lejeunea flav
_Polypodium sp. Undetermined species* Clastobryum ameri-
canum
C. New Haven Gap, | Stelis ophioglossoides Phycopeliis sp.
5600 sk — north
expos
Trentepohlia aurea
Trichomanes sp.
Crossotolejeunea sp.
campan
Cololejeunea diaphana
Lejeunea
Meizgeria furcata
Microlejeunea sp.
| Blechnum attenuatum | Phycopeltis P: Cololejeunea wea Mittenothamnium
Pinnularia s Crossotolejeu reptans
ostoc sp. Di pasieicdions
Seaemena sp. ida
Drepanolejeunea
campanulat
phocolea
Metzgeria furcata
Taxilejeunea sp. —
Clusia havelioides Pinnularia sp. Parmelia sp. A — sp. Mittenothamnium ‘
. ostoc sp. A phano nolejeu: reptans
Scytonema sp. sicaefolia
Diplasiolejeunea
pellucida
Drepanolejeunea
* Up to the present the writer has been unable to have some of the epiphyllous lichens definitely determined.
Region
TABLE I—Continued
Supporting plant
Alene
st patos
Hepaticae
Musci
New Haven Gap—
Continued.
Monoclea Gottschet
Mittenothamnium
reptans
Plagiochila sp.
Riccardia sp.
Mittenothamnium
repians
Rhododendron sp.
Phvlloporina epiphylla
Undetermined species
D. Vinegar Hill
Road, 4500 ft. alt.:
n and west
exposures.
Asplenium sp.
|
|
|
|
|
|
|
| Cololejeunea sp
| Cololejeunea diaphana
| Crossotolejeunea sp.
ee rays
cida
Dreponoeeunca
campan
M aie’ Pirbake
Clusia havetioides
Nost
Phycopeltis sp.
Pinnularia sp.
Parmelia
Cololejeunea sp.
Diplasiolejeunea
tzgeria furcata
Pachcidiies + sp.
Elaphoglossum
latifolium*
Chroococcus sp.
— oa sp.
phen peltis sp.
Scytonema sp.
Cololejeunea ceca
Cros. wines
Lejeunea flav
Moleaeli Gaiccla i
Mittenothamnium
reptans
TABLE I—Continued.
Region : Supporting plant Algae Lichens Hepaticae Musci
4), Hill) Epidendrum sp. Undetermined species. | Drepanolejeunea
eure a d. campanulata
Polypodium Phyllitidis Cololejeunea s Fissidens sp.
sp.
Cololejeunea diaphana
axilejeunea sp
E. Blue Mountain
veer zt —
rthea
Podocarpus coriaceus
Phycopeliis sp.
F. Clyde River bed
ley, 4000 ft.
north and
Citrus Limonium
tiesencions sp.
Scyton sp.
Treatepohtin aurea
Po peacie sp.
Collema
Laat ap.
ss ; ees
Pyrenula sp. 2
Undetermined species.
A re sp.
Cololejeunea
Diplasiolejewnen
pell:
Drepanolejennea
ampan:
ton ‘eunea flava
Metzgeria furcata
8 PEssIN: -EPIPHYLLOUS PLANTS OF JAMAICA
Here one finds plants of many types growing on the soil; on these
grow various plants as epiphytes and on these in turn grow other
epiphytes. A remarkable diversity of epiphytic forms can be
found on one plant, and some of the forms are wonderfully adapted
to their epiphytic mode of life. These epiphytes occur on every
patch of dead or living plant surface that receives even a moderate
supply of light. One finds on a single leaf here such forms as
algae, lichens, hepatics, mosses and, at times, even seed plants,
not to mention occasional fungi and minute epiphyllous animals.
It is rather strange that not a single fern prothallus has been
found on any of the leaves of this region, though numerous spo-
rangia were present and probably spores also on many different
kinds of leaves. In Taste I, B, are listed the different kinds of
epiphyllous forms found near Morce’s Gap.
The climatic conditions of New Haven Gap do not differ
markedly from those of Morce’s Gap. The temperature is nearly
the same, ranging annually from 40.5° to 83° F., and the floor of
the forest is constantly moist and shaded. Here again one finds
the characteristic rain forest vegetation. The epiphyllous plants
here are quite as abundant as at Morce’s Gap and consist mainly
of algae, hepatics, and mosses. The leaves of the climbing plants
of this locality are well inhabited by epiphyllous forms. The
leaves of Clusia havetioides and of such ferns as Blechnum atten-
uatum, Elaphoglossum latiyolium and Polypodium Phyllitidis serve
most frequently as the substratum for epiphylls (see PLATE 1).
A glance at TaBLE I, C, gives one a fair notion of the groups of
epiphyllous plants found in the New Haven Gap region.
Going northward from Morce’s Gap along the upper eastern
slopes of the Mabess Valley the trail is particularly interesting as
it presents a splendid view of the ravines. The crowns of the tree
ferns carpet many side ravines of the valley. The huge lianes,
the numerous orchids, the wild pines and the purplish filmy ferns
mantle the branches of many trees and the species of Asplenium,
Blechnum and Polypodium, along with the walking fern (Poly-
stichum plaschnichianum), spread over the banks. One does not
have to walk off the trail to find numerous epiphyllous plants.
The upper leaves of many of the low trees on the Vinegar Hill
Road are completely covered by them. Tasxe I, D, shows the -
great variety of epiphyllous forms found in this region.
PESSIN: EPIPHYLLOUS PLANTS OF JAMAICA 9
On Taste I, E and F, are indicated the epiphyllous plants
observed on the Blue Mountain Peak and in the Clyde River
Valley. In the former, though there is always an abundance of
moisture, the epiphyllous flora is not a rich one, owing probably
to the frequent heavy rains, the rather low temperature and the
occasional strong winds. Those of the Clyde River Valley occur
chiefly on trees growing along streams, where there is sufficient
shade and a humid atmosphere.
iii E
i. Part of transverse section of a leaf of Elaphoglossum latifolium (EF)
a sa rhizoids (Rh) of an epiphyllous liverwort (L!) pressed into its cuticle. This
liverwort in turn bears another epiphyllous liverwort (L*), which is likewise attached
by its rhizoids (RA)
In general, observations in these Blue Mountains of Jamaica
show that the epiphyllous members of each group of plants are
limited to certain regions where climatic conditions are suitable
for such growth. Thus, Algae, Hepaticae, and Musci usually
thrive best on the windward side of the mountains and are abun-
dant only in very damp and very shady woods. Epiphyllous
lichens, on the other hand, do not require very warm, humid, and
shady places but are also found in relatively dry regions. It was
rather difficult to ascertain with any degree of definiteness the
exact age of the leaves which bore epiphyllous plants, but it is
safe to state that none are present on very young leaves. As a
rule they do not appear on leaves until the latter are one or two
seasons old. :
In order to determine the relation between the epiphyllous
forms and the plants on which they grow, cross sections were
made of the leaves of Elaphoglossum latifolium, Clusia havetioides,
10 PEsSIN: EPIPHYLLOUS PLANTS OF JAMAICA
Polypodium Phyllitidis and Mangifera indica. The leaves of
Elaphoglossum latifolium were those chiefly used because of the
abundance on them of epiphyllous liverworts in various stages of
development. Sections of leaves of Mangifera indica were also
studied to determine the relation of an epiphyllous lichen to the
mango leaf.
Sections of the leaves bearing epiphyllous hepatics showed
characteristic thickened, club-like rhizoids arising in groups from
the stem of the creeping liverwort. These rhizoids are sunken
into the cuticle of the supporting leaf as if pressed in by some
external force. The rhizoids broaden at their lower ends and
flatten out as is shown in TEXT-FIG. I. This is true even of the
rhizoids of one epiphyllous liverwort when growing upon another,
for one sometimes finds two or more tiers of epiphylls growing
upon a fern or a dicotyledon. . The rhizoids of the upper hepatic
are in such cases decidedly flattened against the delicate leaves
of the lower liverworts. This too is shown in the figure. The
important point to be noted is that the rhizoids of the epiphyllous
form were not observed to penetrate the tissues of the supporting
leaf. Nor was there evident any abnormality in the internal
structure of the supporting leaf, such as might conceivably be
produced by the shade or by the weight of the epiphylls on the
leaf or, possibly, by some substance such as an organic acid
excreted by the rhizoids on the liverwort.
The leaves of Mangifera indica bore numbers of orange-red
disk-like plants of a species of Phycopeltis. Among these orange-
red specks there were also minute grey patch-like lichens resem-
bling those described by Cunningham (3) and Ward (9). The
relation between these lichens and the mango leaf on which they
grew has not been definitely determined. The sections of the
mango leaf thus far studied showed no modifications of its tissues
and no penetration by the lichens.
DISCUSSION
The problem of the evolutionary origin and individual develop-
ment of the epiphyllous plants is a complex one which still requires
much careful investigation. Schimper (4) observed that epiphytic
seed plants commonly originate on their support through seeds.
PEsSIN: EPIPHYLLOUS PLANTS OF JAMAICA 11
The scattered seeds germinate on the trunk, branches, or leaves
of a tree or shrub and, with adequate moisture and other nutritive
material, develop into epiphytes. He divides the fruits and seeds
of epiphytes into three categories: first, those transported by
birds, monkeys and other tree-inhabiting animals; second, those
which are very light and may be transported by the wind and
which may readily catch in the crevices of the bark and germinate
there; third, those seeds which are especially adapted to be carried
by the wind. Such seeds usually possess either hairs or wings or
inflated coats which aid their distribution.
It is conceivable that a great many of the epiphyllous forms
here being considered arise from spores, for spores have been found
on many leaves. The epiphyllous liverworts evidently arise
often from gemmae, for many such gemmae in the process of
development have been found on leaves. The spores of liverworts,
on the contrary, have rarely been found germinating on leaves.
The observations thus far made indicate that many an epiphyllous
hepatic or moss makes its start on the host by creeping up the
stem of the support from the ground or up the petiole from the
stem. As it climbs upward it dies off from below and when its
younger part reaches the leaves it spreads over the surface and
becomes strictly an epiphyllous form. Secondly, epiphyllous
forms may be lodged upon the host by the agency of water.
When during a rain the water flows down over the leaves and
branches of a tree in the rain forest, it breaks off the hepatics and
mosses and other plants from the surface of the stem or leaf and
washes them down on the leaves below, where they continue to
grow, covering the leaf in the course of time with a dense mat.
Such is probably the explanation of the fact that in most cases
the lower leaves possess the greatest number of epiphyllous plants,
though, of course, lower leaves are older, better shaded, and are
surrounded by moist air. It is probable also that the epiphyllous
plants found on the dorsal sides of certain overturned leaves arise
in the manner just described. The fact that no epiphyllous
plants were observed on the lower sides of leaves still in normal
position is evidence that these plants are propagated largely by
these fragments. If it were commonly by spores we should
expect these, at least occasionally, to stick to the underside of a
12 PEssIN: EpIPHYLLOUS PLANTS OF JAMAICA
leaf. Such cases, however, are extremely rare. In a third possible
method of dispersal, spores, gemmae, or seeds may be washed down
by the dripping water from above, or may be blown from adjacent
plants by the wind. This method is particularly common among
algae and lichens. The first two types of vegetative propagation
seem to be the commoner ones, for in many instances the epiphyl-
lous forms growing on the stem have actually been observed to
spread over the surface of the leaf. Moreover, many of the forms
found on the leaves occur likewise on the stem of the supporting
plant. This is particularly true of hepatics and mosses. In
lichens, however, the case may be different. There the alga, as
well as the component fungus, may often be present separately on
the leaf; each may reach the leaf in the form of loose spores or
fragments, and there combine to give rise to the lichen itself.
Massart (3) failed to find in Java any epiphyllous pteridophytes
and phanerogams. In the tabulated lists of epiphyllous plants of
Jamaica in this paper it will be noted that no pteridophytes have
been found. Although many fern sporangia were observed among
the epiphylls by the writer, no germinating fern spores or fern
prothallia were seen. The writer, however, found a few epiphyl-
lous phanerogams. These were very young plants of the genus
Caraguata attached to the surface of the leaf, and apparently doing
well. How much further their development would continue is
hard to tell, for the substratum is not firm enough to support a
mature wild pine; before the epiphyllous wild pine could have time
to mature, the supporting leaf would probably be shed. The fact
is, at all events, that no mature wild pines were found growing on
leaves. According to Massart, the epiphyllous plants do not
constitute.a similar flora everywhere; in some localities this consists
solely of thallophytes, in others of bryophytes. This is also true
for Jamaica, where in certain regions only one type occurs, while
in other regions numerous diverse forms are present.
Massart’s suggestion that the leaf-inhabiting fungus has been
derived from the epiphyllous plant is at least favored by the rela-
tion to its host of the alga Cephaleuros, which may have either the
parasitic or the epiphytic mode of life. On the other hand, there
are still some fungi which possess the epiphyllous habit, such as
Metiola astrina and a few others. There seems little doubt that
PEsSIN: EPIPHYLLOUS PLANTS OF JAMAICA 18
a careful study of epiphyllous plants would throw some light on
the origin of parasitism, such as that of leaf-infesting fungi.
SUMMARY
. Epiphyllous plants of diverse selected regions in Jamaica
were ase in the field and collected for laboratory study in
Baltimore.
2. The results of the field observations were tabulated so as
to give at a glance a general idea of the distribution of these
epiphyllous plants in relation to the altitude, average temperature,
and average humidity of their various habitats.
3. The possible modes of attaining the epiphyllous position
are:
(a) Continual growing forward and dying off below, of hepatics
and mosses, until they reach and cover the leaf.
(6) By distribution of spores, gemmae or seeds through the
agency of rain and wind. This is especially true for algae and
fungi.
(c) Propagation by displaced fragments af mature lichens,
hepatics, and mosses.
. An abundance of humidity, gistoraie temperature, and a
raletivaly calm atmosphere encourage an otis epiphyllous
growth.
\ 5. Histological studies of epiphyll-bearing leaves of Elapho-
glossum latifolium, Clusia havetioides and Mangifera indica revealed
no apparent penetration of — leaf by any part or organ of the
epiphyll.
6. The rhizoids of epiphyllous hepatics are especially adapted
for the epiphyllous habit by becoming thickened, by flattening
out and by being pressed into the cuticle of the leaf, thus firmly
attaching the epiphyll to the leaf and preventing its being blown
or washed off easily.
7. It is conceivable that the shade produced by the epiphyll,
the accumulation of the organic material, the possible excretion
of acids and other substances by its rhizoids may, have some
physiological effect on the supporting leaf. |
8. A careful study of the epiphyllous mode of life will, it is
believed, bring out some close relations between parasitism and
e
14 PEssIN: EPIPHYLLOUS PLANTS OF JAMAICA
the epiphyllous habit. Parasitism is possibly an advanced,
specialized phase of epiphytism.
The writer is indebted to the Botanical Department of the
Johns Hopkins University for the opportunity of going to Jamaica.
The writer wishes to thank particularly Professor Duncan 5.
Johnson for suggesting this problem, for taking the photographs
reproduced on PLATE 5, and for helpful advice in carrying on the
work. Acknowledgments are due Professor Alexander W. Evans
for the determination of most of the epiphyllous hepatics; to
Mrs. N. L. Britton of the New York Botanical Garden for the
determination of some of the epiphyllous mosses; and to Pro-
fessors C. C. Plitt and Lincoln W. Riddle for the determination
of certain epiphyllous lichens.
LITERATURE CITED
. Berkeley, M. J. On the thread blight of tea. Quart. Jour. Microsc.
Sci. 15: 130-133. f. 7-4. 1875.
_
2. Cunningham, D. D. On Mycoidea parasitica. Trans. Linn. Soc.
London II. 1: 301-316. pl. 42, 43. 1877.
3. Massart, J. Les végétaux épiphylles. Ann. Jard. Bot. Buitenzorg.
Suppl. 2: 103-108. 1898. é
4. Schimper, A. F. W. Botanische Mittheilungen aus den Tropen 2.
Die epiphytische Vegetation Americas. Jena. 1888.
5. Schmidle, W. Einige Baumalgen aus Samoa. Hedwigia 36: 277-
287. pl. 6-9. 1897.
6. Ward, H. M. On the structure, dsyeloomuut and life history of a
tropical epiphyllous lichen. Trans. Linn. Soc. London II. 2: 87-
119. pl. 18-27. 1883.
Explanation of — x
Fic. 1. - Leaf of nSubed peated cma: h Il h osse:
Fic. 2. Leaf of Elaph atifolium : showing a ‘tusuriant growth of ..
lous plants; a small wild pie; pei Stiteues. can be seenon the right hand
margin.
The genus Catinella
Exuias J. DuraND
The genus Catinella was established by Boudier* for a single
species, of rather indefinite relationship, which has commonly
been identified as Peziza olivacea Batsch. While Batsch’s plant
is indeterminable when tested according to present day standards
of specific limitations, the fungus which has gone by that name is
- well known, being clearly described by Massee,} and nicely figured
by Boudier.{ The synonymy listed below emphasizes not only
the truth of Boudier’s statement that the species has been in-
cluded sometimes in one genus, sometimes in another, but that
it has been referred by various writers to several different families
as well. The inoperculate asci and intense reaction to caustic
potash remove it from the Pezizaceae, where the bright color and
fleshy consistency of the fresh young plants at first seem to place
it, and to which it has been referred by Saccardo, Rehm and others.
The somewhat gelatinous nature of the excipulum has. suggested
the Bulgariaceae to Fries, Karsten and others, but that character
is certainly not conspicuous enough to associate the plant with
Bulgaria inquinans or Sarcosoma rufum. Moreover, a truly
gelatinous tissue does not become friable on drying. The present
writer is inclined to the opinion of Berkeley, Phillips, Massee and
Boudier that its affinities are more properly with the Patellari-
aceae, such, for example, as Karschia. The genus may be charac-
terized as follows:
A genus of the Patellariaceae. Ascomata fleshy and somewhat
gelatinous when fresh, becoming friable when dry, sessile, attached
to the substratum by radiating, dark fibers; excipulum entirely
parenchymatous, becoming deep violet with KOH; asci opening
by a pore, not blue with iodine; spores eight, continuous, pale
brown; asci and paraphyses agglutinated at the tips to form an epi-
thecium.
* Hist. Class. Disc. Eur. 150. 1907.
+ Brit. Fungus-Fl. 4:94. 1895.
} Icon. Myc. 3. pl. 452.
15
16 DuRAND: THE GENUS CATINELLA
Species two, as follows:
Spores 7-It X 4-5 p. ; 1. C. nigro-olivacea.
Spores 13-15 X 6-7 n. 2. C. elastica.
1. Catinella nigro-olivacea (L. v.S.) Durand, comb. nov.
?Peziza olivacea Batsch, Elench. Fung. 127. pl. 12. f. 51. 1783.
Peziza nigro-olivacea L. v.S. Syn. Car. 121. 1822.
Patellaria pulla 8 nigro-olivacea Fr. Syst. 2: 160. 1822.
Bulgaria nigrita Fr. Elench. 2: 16. 1830.
Lemalis rufo-olivacea L. v. S. Syn. N. Am. n. 1089. 1834.
Rhizina nigro-olivacea Curr. Trans. Linn. Soc. 24: 494. pl. 51.
f. 10-12 (fide Phillips).
Peziza viridi-atra B. & C. Jour. Linn. Soc. Bot. 10: 369. 1868.
Patellaria violacea B. & Br. Jour. Linn. Soc. Bot. 14: 108. 1875
(fide Massee I.c.).
Patellaria hirneola B. & Br. pk Linn. Soc. Bot. 14: 108.
1875 (fide Massee l.c.). ~
Patellaria applanata B. & Br. Jour. Linn. Soc. Bot. 14: 108.
1875. :
Peziza fuscocarpa Ell. & Hol. Jour. Myc. 1:5. 1885.
Patellaria olivacea Phill. Brit. Disc. 361. 1887 (Batsch ?).
?Humaria olivacea Sacc. Syll. 8: 148. 1889 (Batsch ?).
Pezicula viridi-atra Sacc. Syll. 8: 315. 1889.
Phaeopezia fuscocarpa Sacc. Syll. 8: 474. 1889. -
Bulgariella pulla 8 nigro-olivacea Sacc. Syll. 8:638. 1889.
Bulgariella nigrita Sacc. Syll. 8:638. 1889.
Patinella violacea Sacc. Syll. 8: 770. 1889 ( fide Massee).
Patinella olivacea Sacc. Syll. 8: 770. 1889 (Batsch ?).
Patinella hirneola Sacc. Syll. 8: 771. 1889 (fide Massee).
Patinella applanata Sacc. Syll. 8: 771. 1889.
Humaria marchica Rehm, Rabenh. Kypt.-Fl. 1°: 952. 1894.
Phaeopezia marchica Sacc. Syll. 11: 415. 1895.
Aleurina marchica Sacc. & Syd. Syll. 16: 739. 1902.
Humaria fuscocarpa Morgan, Jour. Myc. 8: 189. 1902.
Aleurina fuscocarpa Sacc. & Syd. Syll. 16: 739. 1902.
Catinella olivacea Boud. Hist. Class. Disc. Eur. 150. 1907
(Batsch ?).
A scomata solitary or two or three together, sessile, attached to
‘the substratum by numerous, radiating, dark brown fibers most
DuRAND: THE GENUS CATINELLA 17
conspicuous in young plants; at first spherically closed, then open-
ing out so as to become successively cup-shaped, saucer-shaped,
finally applanate, with a permanently upturned margin; ver
young plants greenish yellow, becoming deeper and darker green,
finally blackish olive when old, fleshy and somewhat amigete
when fresh, 3-10 mm. in diameter, the majority 3-5 m On
drying the disk becomes olive-black, the exterior eowiishis some-
what furfuraceous or vertically striate, and the texture friable.
Excipulum about 400 yu thick at the base, entirely parenchymatous,
the ectal cells 18-20 » in diameter, polygonal, with dark brown ~
walls, becoming gradually smaller and yellow toward the hy-
menium, the superficial cells of the margin and sides projecting to
form short, stout, obtuse, septate, flexuous, brown pili causing the
surface to appear furfuraceous, those at the base much longer and
radiating so as to form a small subiculum covering the substratum
2-3 mm. beyond the margin. On treatment with 1 per cent
caustic potash the hymenium or whole flesh becomes deep violet
and frequently yields a violet solution. Hymenium about 120-
140 w thick. Asci narrowly cylindric-clavate, apex rounded, not
blue with iodine, 65-90 x 3-6 uw (mostly 70-80 yn). Spores eight,
uniseriate, elliptic or elliptic-clavate, usually slightly narrowed
near the middle so as to appear slipper-shaped, continuous, two-
guttulate, pale olive-brown to deep brown, 7-11 x 4-5 uw (majority
8-10 yw); Paraphyses cylindric, septate, rarely branched, somewhat
longer than the asci, the tips agglutinated with amorphous matter
to form an epithecium
On rotten wood, especially of old, moist, carious, decorticated
logs, not abundant but widely distributed in eastern North America
and the West Indies; also in Europe and Ceylon.
This is one of our most peculiar and attractive discomycetes.
“In its young state it is truly Peziza-like, and very beautiful.”
While easily recognizable at sight when once understood, it has
been described as new over and over again until very few fungi
have a greater multiplicity of names. While this may be the
species to which Batsch applied the name Peziza olivacea, his brief
description and crude figures are much too inadequate to justify
more than surmise, and may apply equally well to numerous other
species. The apparent rarity of our plant in central Europe makes
it still more improbable that Batsch had seen it. /P. olivacea is
simply one of the numerous names of fungi handed down from the
pre-Persoonian period, the mycological stone age, which are ab-
solutely indeterminable according to present taxonomic standards,
18 DuRAND: THE GENUS CATINELLA
and which should therefore be dropped completely and no longer
allowed to encumber the mycological literature.
The first certainly identifiable name applied to the plant under
consideration is the one adopted in the present paper. While
Schweinitz’s type, from North Carolina, is missing from his her-
barium, there is a specimen marked ‘‘rotten wood. Beth.’’ under
the name ‘‘ Lemalis pulla B nigro-olivascens Schw. No. 1089.”’ This
label involves a curious switching of names and an error in spelling
as follows: Peziza nigro-olivacea L. v. S. was included by Fries in
his Systema as “Patellaria pulla B nigro-olivacea.”” In 1834,
Schweinitz transferred the species to the genus Lemalis, and
changed the specific name so that it appeared as ‘“‘1089. 4. L.
rufo-olivacea, L. v. S., Syn. Car. 1220, Peziza nigro-olivacea; Salem
“et Bethlehem differt specifice a priori [L. pulla Fr.]’’. At a later
date, Dr. Michener, in mounting and arranging Schweinitz’s fungi,
restored the original specific name (nigro-olivacea) to the label,
but misspelled it ‘ igro-olivascens.”’ The presence of numbers and
references in each case leaves no doubt that the specimen now in
the Schweinitzian Herbarium is the one referred to in 1834 as from
Bethlehem.
Although Fries stated that he had seen a specimen of Pesziza
nigro-olivacea, there seems to be no Schweinitzian material so
labeled in the Friesian Herbarium, at Upsala. However, there is a
specimen called ‘‘ Lemalis rufo-olivacea Schw.,’’ from Curtis, as well
as one marked “ Peziza applanata ex herb. Schweinitz.’’ While
both are typical Catinella nigro-olivacea, the latter is quite dif-
ferent from the specimen in Schweinitz’s own herbarium called
P. applanata.
In 1830, Fries described as Bulgaria nigrita a plant collected
in Russia by Weinmann. So far as the writer can discover, no
more recent collection has been referred to this species, which has
remained practically unknown for nearly a century. In the
Friesian Herbarium is a specimen marked “Bulgaria nigrita,
Petersburg,” in Elias Fries’s own hand, which is in all probability
the original type. When the writer saw it, in 1904, he was at once
impressed with its gross resemblance to the American plant. Sub-
sequent microscopical examination removed all doubts as to their
specific identity.
DuRAND: THE GENUS CATINELLA 19
During the Berkeleyan period this species was collected occa-
sionally in various parts of the world, and almost as often described
as new. Material from Cuba (Wright, No. 369) was given the
name Peziza viridi-atra B. & C. Examination of the type, at
Kew, shows it to be identical with the Schweinitzian plant.
Saccardo placed it in Pezicula. Specimens from Ceylon were
called Patellaria applanata B. & Br. The writer has not seen the
type, but Massee, on examination, declared it to be Patinella
olivacea (C. nigro-olivacea). On the basis of the same evidence
the same writer included P. violacea B. & Br. and P. hirneola
B. & Br., also from Ceylon, as synonyms. In the original descrip-
tion of P. applanata the authors mentioned a “United States
specimen.’ Material so labeled in the herbaria of Fries and the
Philadelphia Academy of Science, collected in South Carolina,
by Curtis, has been examined, and certainly belongs here. Addi-
tional collections reported from Connecticut (Wright), Pennsyl-
vania (Michener), and North Carolina (Curtis) have not been
seen.
No specimen of Rhizina nigro-olivacea Curr. has been available
to the writer. But since both Phillips and Massee (the latter
after examination of the type) place it as a synonym of P. olivacea,
there seems to be no doubt of its identity with C. nigro-olivacea.
Peziza fuscocarpa Ell. & Hol. was described from material col-
lected in lowa by Holway. It has been referred to Phaeopezig by
Saccardo, to Humaria by Morgan, and to Aleurina by Saccardo &
Sydow, and by Rehm. This specific name is the one under which
the plant here considered has commonly been reported or dis-
tributed in America. The type is identical with Peziza nigro-
olivacea L. v.S. The writer has received it from many localities
and has had abundant opportunity to study it in the fresh con-
dition in all stages of development.
Of all the references of the present species made by Saccardo,
the most curious is that where Peziza nigro-olivacea L. v. S. is
made a synonym of Mollisia umbonata (Pers.) Sacc. To Per-
soon’s specific description are appended verbatim Fries’s observa-
tions comparing Schweinitz’s plant with Patellaria pulla Fr., a
totally different plant from Persoon’s. This confusion may be
due to a mixing of notes. o
20 DURAND: THE GENUS CATINELLA |
Finally, the plant here considered was described as Humaria
marchica by Rehm, from German material distributed by Sydow
and examined by the writer.
While it has been demonstrated that Catinella nigro-olivacea
has passed under about thirty different names, it may well be that
others have escaped the writer’s attention.
In the citation of specimens the following abbreviations have
been used: CU, Herbarium of Cornell University; D, Durand
Herbarium; F, Fries Herbarium; K, Kew Herbarium; Mo, Her-
barium of the Missouri Botanical Garden; NY, Herbarium of the
New York Botanical Garden; ‘Phil, Philadelphia Academy; S,
Schweinitz Herbarium.
MATERIAL EXAMINED
Russia: Petersburg (F, type of Bulgaria nigrita Fr.).
GERMANY: Brandenburg, Sydow (NY, cotype of Humaria
marchica Rehm, in Sydow, Myc. March. 2958); Moravia, Petrak
(D, as Aleurina olivacea [Batsch] v. Héhn.).
ONTARIO: Carleton Place, Macoun (NY); London, Dearness
(CU, D, E. & E. N. Am. Fungi 2325). |
New HAmpsuireE: Warren, L. W. Riddle (D).
New York: Ithaca, Atkinson (D); Varna, Reddick (D);
Honeoye, Durand (D); Adirondack Mountains, Catskill Moun-
tains, Old Forge, Kasoag, Big Indian, South Pond, Elisabethtown,
Peck (A, as Patellaria olivacea).
PENNSYLVANIA: Bethlehem, Schweinitz (S); Springtown, Witte
(NY); Mauch Chunk (Phil); Chester County, Michener (Phil).
Nortu Caro_rna: Blowing Rock, Durand (D).
SouTH CAROLINA: Curtis (F, Phil).
LouIsIANA: Cloutierville & Abita Springs, Langlois (NY).
ARKANSAS: Camden, C. J. Humphrey (D)
Onto: Preston, Morgan (NY).
MinneEsora: Lakeville (D); St. Louis River, Holway (NY, Mo).
Nortu Dakota: Fargo, Seaver (NY, Mo, N. Dak. Fung. 28,
as Phaeopezia fuscocarpa).
Iowa: Mt. Pleasant, Seaver (D); Decorah, Holway (NY, type
of Peziza fuscocarpa).
CuBa: Wright 369 (K, type of Peziza viridi-atra)
DuRAND: THE GENUS CATINELLA 21
2. Catinella elastica (Pat. & Gail.) Durand, comb. nov.
Phaeopezia elastica Pat. & Gail. Bull. Soc. Myc. Fr. 4: 99.
1888.
Aleurina elastica (Pat. & Gail.) Sacc. & Syd. Syll. 16: 7309.
1902; Rehm, Ann. Myc. 1: 515. 1903.
Ascomata sessile, solitary or in groups, I-3 mm. in diameter,
rich dark brown externally, black within, attached at the base by
radiating brown fibers consisting of septate hyphae; margin
. thickened. Excipulum parenchymatous, of polygonal cells, the
ectal ones with brown walls, those at the sides of the cup giving
rise to short, septate, flexuous, brown pili, those at the base to
stout, brown, radiating hyphae. Asci cylindric-clavate, apices
rounded, not blue with iodine. Spores eight, uniseriate, elliptic,
or elliptic-clavate, or slipper-shaped, continuous, brown, two-
guttulate, 13-15 x 6-7 4. Paraphyses cylindric, hyaline, septate,
tips agglutinated with amorphous matter.
MATERIAL EXAMINED
VENEZUELA: on dead wood, Mapire, May, 1887, A. Gatllard 6
(NY).
This species seems to differ from the preceding only in the
larger spores. In the herbarium of the New York Botanical
Garden is a specimen of what appears to be the original and only
collection. The above description is that of the authors supple-
mented by notes drawn from the above mentioned specimen.
In the original description the tissue is spoken of as elastic, some-
what like that of Bulgaria. The dried material is friable. It
breaks up so that no complete ascus has been seen. The material
becomes deep violet on treatment with KOH.
UNIVERSITY OF MINNESOTA
INDEX TO AMERICAN BOTANICAL LITERATURE
1908-1921
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,
mance ae products of vegetable origin, or labora atory me 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 mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
rs or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to sub-
scribers at the rate of three cents for each card. Selection of cards is not permitted:
each subscriber must take all cards published during oy term of his subscription.
Correspondence relating to the card issue should be addressed to the Treasurer of
the Torrey Botanical Club.
Andres, H. Piroleen-Studien. Verh. Bot. Ver. Brandenburg 56: 1-70.
6 Je 1914.
Arber, A. Leaves of the Helobieae. Bot. Gaz. 72: 31-38. pl. I-
16 Jl 1921. :
Arthur, J.C. Specialization and fundamentals in botany. Am. Jour.
Bot. 8: 275-285. 30 Je 1921.
Bakhuizen Van den Brink, R.C. Revisio generis Avicenniae. Bull.
Jard. Bot. Buitenzorg III. 3: 199-226. pl. r4-22. My 1921.
Barnhart, J. H. Biographical notices of persons mentioned in the
Schweinitz-Torrey correspondence. Mem. Torrey Club 16: 290-
300. 16 Jl 1921.
Blake, S. F. A new Aspilia from Trinidad. Proc. Biol. Soc. Washing-
ton 34: 119, 120. 30 Je 1921.
Blake, S. F. New Meliaceae from Mexico. Proc. Biol. Soc. Washing-
ton 34: 115-118. 30 Je 1921
New species in Cedrela (1) and Guarea (2).
Bradshaw, R. V. Oregon willows. Am. Bot. 27: 56-59. My 1921.
23
24 INDEX TO AMERICAN BOTANICAL LITERATURE
Britton, N. L. The cacti of Trinidad. Bull. Dept. Agr. Trinidad &
Tobago 19: 81-87. 1921.
Carano, E. Nuove ricerche sulla embriologia delle Asteraceae. Annali
di Botanica 15: 97-196. pl. 4-12 + f. 1, 2. 28 F 1921. |
Campbell, D. H. Professor H. Bruchmann. Science II. 54: 67, 68.
22 Jl 19at.
Campbell, D.H. The eusporangiate ferns and the stelar theory. Am.
Jour. Bot. 8: 303-314. f. 1-7. 30 Je 1921.
Campbell, D.H. Die Verbreitung gewisser Lebermoose der malaiischen
Region. Jahrb. Wissensch. Bot. 56: 365-373. 1915.
Chamberlain, C. J. Helmut Bruchmann. Bot. Gaz. 72: 45-47. 16
Ji 1921. [With portrait.]
Child, C. M. Certain aspects of the problem of physiological correla-
tion. Am. Jour. Bot. 8: 286-295. 30 Je 1921.
Clark, S. P. Sweet clover in Arizona. Univ. Arizona Coll. Agr.
Circ. 347 4-7: Ap i921.
[Clute, W. N.] A new form of Dentaria. Am. Bot. 27: 69. My 1921.
Dentaria laciniata var. dichronum from lowa.
Collins, J. L. Inbreeding and crossbreeding. Jour. Hered. 12: 89-93.
fc ZF-g4e 214 J teen.
“The effect of inbreeding and crossbreeding in a wild plant [Crepis capillaris}
of the sunflower family.”
Conard, H. S. An outline for vascular plants. Science II. 54: 15, 16.
1 Jl 1921.
Cook, M. T., & Martin, W. H. Potato diseases in New Jersey. New
Jersey Agr. Exp. Sta. Circ. 122: 1-39. f. 1-21. F 1921.
Cook, M. T., & Poole, R. F. Diseases of sweet potatoes. New Jersey
Agr. Exp. Sta. Circ. 123: 1 1-24. f. 1-17. Ap 1921.
Douin, C. La famille de Cephaloziellacées. Soc. Bot. France Mem.
29: 1-90. f. f-102. 26 D 1030.
Includes as American novelties, the genus Evansia, and species in Prionolobus (1),
Evansia (1), and Cephalozielia (9).
East, E. M., & Jones, D. F. Round tip tobacco—a plant ‘‘ made to
order.”’” Jour. Hered. 12:. 51-56. 61. 1 +f. 1-5. 11 Jl 1921.
Felt, E. P. Key to American insect galls. New York State Mus. Bull.
200: 1-310. pl. I-16 + f. 1-250. 1 Au 1917.
Fernald, M. L. The geographic distribution of hybrids. Science II.
54: 73, 74. 22 Ji 1921. :
Folsom, D. Potato leafroll. Maine Agr. Exp. Sta. Bull. 297: 37-52.
f. 26-35. Ap 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 25
Fromme, F. D., & Wingard, S. A. Varietal susceptibility of beans to
rust. Jour. Agr. Research 21: 385-404. pl. 69-73. 15 Je 1921.
Glover, W.O. Blister canker of apple and its control. New York Agr.
Exp. Sta. Bull. 485: 1-71. pl. 1-15 + f. 1-8. Ja 1921.
Hansen, A. A. Protect our vanishing wild plants! Conservationist 4:
67-70. My 1921. [Illust.]
Harms, H. Uber die Fluorescenz des Aufgusses der Rinde von Prunus
serotina. Verh. Bot. Ver. Brandenburg 59: 146-150. 15 Ja 1918.
Harms, H. Uber Fluorescenz-Erscheinungen bei dem Holze der
Leguminose Eysenhardtia amorphoides H. B. K. Verh. Bot. Ver.
Brandenburg 56: 184-197.° 18 Ja 1915.
Harms, H. Nachtrage und Verbesserungen zu meinem Aufsatze tiber
Fluorescenzerscheinungen. Verh. Bot. Ver. Brandenburg 57: 191-
202. 1 Mr-1976,
Harms, H. Ernst Ule. Verh. Bot. Ver. Brandenburg 57: 150-184.
1 Mr 1916. [With portrait.]
Harris, J. A. Tissue weight and water content in a tetracotyledonous
mutant of Phaseolus vulgaris. Proc. Soc. Exper. Biol. and Med. 18:
207-209. 1921.
Harvey, E. M., & Murneek, A. E. The relation of carbohydrates and
nitrogen to the behavior of apple spurs. Oregon Agr. Coll. Exp.
Sta. Bull. 176: 1-47. f. I-12. Ap 1921.
Hayes, H. K., & Gardner, R. J. Breeding crop ; planta: Pp. i-xvii
+ 1-328. f. I-66. New York. 1921.
Hume, A. M., Loomis, H., & Hutton, J.G. Water as a limiting factor
in the growth of sweet clover (M[elilotus] alba). South Dakota
Agr. Exp. Sta. Bull. 191: 257-298. pi. r, 2. Jl 1920.
Hutchinson, C. B. Heritable characters of maize—VII. Shrunken
endosperm. Jour. Hered. 12: 76-83. f. 20-24. 11 JI 1921.
Hutchinson, J. The genera of Fumariaceae and their distribution.
Kew Bull. Misc. Inform. 1921: 97-115. 1921.
Illick, J. S. The white pine. Am. For. 27: 422-426. Jl 1921.
[Ilust.]
Jones, L. R., Walker, J. C., & Tisdale, W. B. Fusarium resistant
cabbage. Univ. Wisconsin Agr. Exp. Sta. Research Bull. 48: 1-34.
f. 1-10. N 1920.
Judd, C. S. The Alahee tree. Hawaiian For. & Agric. 18: 133-137-
Je 1921. [Illust.]
26 INDEX TO AMERICAN BOTANICAL LITERATURE
Kraemer, H. Plant colors. Am. Jour. Pharm. 93: 414-416. Je 1921.
Kraemer, H. Some experiments on the modification of color in plants.
Am. Jour. Pharm. 93: 416-418. Je 1921.
Kunkel, L. O. A possible causative agent for the mosaic disease of
corn. Bull. Exp. Sta. Hawaiian Sugar Pl. Assoc. 3: 1-15. pl. 4-15
+ fe. Fy. Bex > Jt 19a:
Lee, H. A. The relation of stocks to mottled leaf of Citrus trees.
Philipp. Jour. Sci. 18: 85-93. pl. 1-3. Ja aig2t.
Loesener, T . Plantae Selerianae—IX. Verh. Bot. Ver. Brandenburg
58: 129-157. 1 My 1917.
Includes new American species in Nemastylis (1), Iresine (1), Daphnopsis (1),
and Aniseia (1).
Lonay, H. Sur quélques genres rares ou critiques de Renonculacées.
Bull. Soc. Bot. Belgique 45: 191-204. 4 D 1908.
Long, W. H. Notes on new or rare species of rusts. Bot. Gaz. 72:
39-44. 16 Jl 1921.
Includes new species in Gymnosporangium (1), and Ravenelia (3).
MacDougal, D. T. Water deficit and the action of vitaniines, amino-
compounds, and salts on hydration. Am. Jour. Bot. 8: 296-302.
30 Je 1921.
Martin, W. H. Studies on tomato leaf-spot control. New Jersey
Agr. Exp. Sta. Bull. 345: 1-42. pl. 1 +f. 1. N 1920.
Maxon, W. R. A neglected fern paper. Proc. Biol. Soc. Washington
34: Tlt-1i4."' 30 Je fozt.
Refers to Watt’s Catalog of Ferns, published in Canadian Naturalist if, 33:
157-160. :
Merrill, E. D. New Philippine Moraceae. Philipp. Jour. Sci. 18:
49-69. Ja 1921.
Merrill, E. D. New Philippine Myrtaceae. Philipp. Jour. Sci. 18:
289-308. Mr 1g2t.
Mulford, F.L. Oaks for ornamental planting. Am. For. 27: 461-467-
Jl 1921. [Illust.]
Nash, G. V. Acacia armata. Addisonia 6: 13. pl. 199. 29 Je 1921.
Nash, G. V. Acacia leprosa. Addisonia 6: 7. pl. 196. 29 Je 1921.
Nash, G. V. Acacia lineata. Addisonia 6: 15. pl. 200. 29 Je 1921.
Nash, G. V. Acacia longifolia floribunda. Addisonia 6: 11. pl. 198.
29 Je 1921.
Nash, G. V. Acacia Nabonnandi. Addisonia 6: 9. pl. 197. 29 Je
1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 27
Nash, G. V. Acacia pubescens. Addisonia 6: 3, 4. pl. 194. 29 Je
1921.
Nash, G. V. Acacia pulchella. Addisonia 6: 5. pl. 195. 29 Je 1921.
Nash, G. V. Acacia scapuliformis. Addisonia 6: 1, 2. pl. 193. 29 Je
1921. ‘
Nelson, E. W. Lower California and its natural resources. Mem.
Nat. Acad. Sci. 161: 1-171. pl. 1-35. 1921.
Neville, H.O. Pineapples in Cuba. Cuba Rev. 19: 13-28. Jl 1921.
[Illust.]
Osterhout, W. J. V., Thaxter, R., & Fernald, M. L. Lincoln Ware
Riddle. Science II. 54: 9. 1 Jl 1921.
Parish, S.B. A reminiscence of roses. Am. Bot. 27: 60,61. My 1921.
Patouillard, N. Clathrotrichum, nouveau genre d’hyphomycétes. Bull.
Soc. Mycol. France 37: 33-35. .15 Ap 1921.
Parasitic on Setaria in Ecuador.
Pittier, H. Acerca del genero Gyranthera Pittier. Bol. Com. e Indust.
[Venezuela] 13: 42 — Ja 1921.
Includes G. caribensis sp. n
Pittier, H. Arboles y ee nuevos de Venezuela—I. Bol. Com. e
Indust. [Venezuela] 13: 417-428. Ja 1921
Includes new species in Inga (1), Raputia (1), Esenbeckia (1), Trichilia (1),
Stillingia (1), Gustavia (1), Mouriria (1), Oxythece (1), Pouteria (1), and Sider-
xylum (1).
Rapp, C. W. Bacterial blight of beans. Oklahoma Agr. Exp. Sta.
Bull. 131: 1-39. f. 1-17. Jl 1920.
Reinking, O. A. Citrus diseases of the Philippines, southern China,
Indo-China, and Siam. Philipp. Agric. 9: 121-179. pl. 1-14. F
1921.
Reinking, O. A. Diseases of economic plants in Indo-China and Siam.
Philipp. Agric. 9: 181-183. F 192.
Rolfe, R. A. Wercklea lutea [Malvaceae]. Kew Bull. Misc. Inform.
1921: 118, 119. 1921. ;
New species from Costa Rica.
Rydberg, P. A. Two new species from Arizona. Am. Bot. 27: 61-63.
My 1921.
Eriogonum Clutei Rydb. and Forestiera arizonica (A. Gray) Rydb.
Sands, W.N. Plants common to the West Indies and Malaya. Agric.
News 20: 163; 182, 183. 28 My 1921.
28 INDEX TO AMERICAN BOTANICAL LITERATURE
Schmitz, H., & Daniels, A.S. Studiesin wood decay—I. Laboratory
tests on the relative durability of some western coniferous woods
with particular reference to those growing in Idaho. School For-
estry Univ. Idaho Bull. 1: 1-11. Jl 1921.
Schmitz,H. Studiesin wood decay—If. Enzymeactionin Polyporus
volvatus Peck and Fomes ignarius (L.) Gillet. Jour. Gen. Physiol. 3:
795-800. 20 J] 1921.
Shear, C. L., & Stevens, N. E. (Editors). The correspondence of
Schweinitz and Torrey. Mem. Torrey Club 16: 119-300. pl. 6, 7.
16 Jl 1921.
Sherman, H. Respiration of dormant seeds. Bot. Gaz. 72: 1-30.
f. 1-4. 16 Jl 1921.
Soth, M. E. Goat-weed in Idaho. Am. Bot. 27: 70. My 1921.
Sousa Brito, E.C. de. Algumas plantas uteis do Brasil, 0 hybridismo
de uma dellas. Arch. Esc. Sup. Agric. e Med. Vet. .4: 25-54. pl.
10-16. D 1920.
Stadler, L. J., & Helm, C. A. Corn in Missouri—I. Corn varieties
and their improvement. Missouri Agr. Exp. Sta. Bull. 181: 1-51.
Jj. 1-7. Mr 1021.
Stahel, G. De Sclerotium-ziekte van de Liberiakoffie in Suriname
veroorzaakt door Sclerotium coffeicolum nov. spec. Dept. Landb.
Suriname Bull. 42: 1-34. pl. 1-12. Ja 1921.
Stevens, F. L. The relation of plant pathology to human welfare.
Am. Jour. Bot. 8: 315-322. 30 Je 1921.
Stout, A. B. Types of flowers and intersexes in grapes with reference
to fruit development. New York Agr. Exp. Sta. Techn. Bull. 82:
1~16. pl. 1-7. Ja 1921.
Sydow, H. & P. Novae fungorum species—XVI. Ann. Mycol. 18:
154-160. Ap 1921.
ncludes new American species in Septobasidium (1), widen dog? (1), Aecidium (1),
Catacauma (1), Taphrina (1).
Taubenhaus, J. J. A study of the black and the yellow molds of ear
corn. Bull. Texas. Agr. Exp. Sta. 270: 3-38. f. 1-10. O 1920.
Taubenhaus, J. J.. & Mally, F.W. Pink root disease of onions and its
control in Texas. Texas Agr. Exp. Sta. Bull. 273: I-42. f. f+3.-
Ja 1921.
Taylor, W. R. Additions to the flora of ee Desert, Maine. Rhodora
23: 65-68. 20 Ap Ig2t.
INDEX TO AMERICAN BOTANICAL PITERATURE | 29
Thatcher, R. W. The chemistry of plant life. Pp. i-xvi + 1-268.
New York. 1921.
. G. E. Hegari in Arizona. Univ. Arizona Coll. Agr. Circ.
1-4. Ap iga2t. [lIllust.]
whee, R.C. Botrytis rot and wilt of tomato. Bull. Ohio Agr. Exp.
Sta. 6: 59-62. Ap i921. [Illust.]
Thurston, H. W. A note on the corrosive sublimate treatment for
the control of Rhizoctonia. Phytopathology 11: 150, 151. r
1921.
Tiffany, L. H. Algal food of the young gizzard shad. Ohio Jour. Sci.
21: 113-122.. FB igea
Tiffany, L. H. New forms of Oedogonium. Ohio Jour. Sci. 21: 272,
273. $f. Ts Je oe
2 new species from Ohio.
Tisdale, W. H., & Jenkins, J. M. Straighthead of rice and its control.
U.S. Dept. Agr. Farm. Bull. 1212: 1-16. f. 1-6. Je 1921.
Toumey, J. W. Damage to forests and other vegetation by smoke,
ash and fumes from manufacturing plants in Naugatuck Valley,
Connecticut. Jour. For. 19: 367-373. f. 1-6. Ap 1921. :
Trelease, W. The Jack Oak (Quercus ellipsoidalis). Trans. Illinois
Acad. Sci. 12: 108-118. pl. 139-143. 1919.
Trelease, W. North American pipers of the section Ottonia. Am.
_. Jour. Bot. 8: 212-217. pl. 5-8. 30 Ap 1921.
Tucker, E. M. Bibliographical notes. Jour. Arnold Arbor. 2: 181-
184. 1921.
Discussion of issuance of Nouveau Duhamel and of Torrey and Gray’s Flora of
North America.
Tucker, E. M. Catalogue of the library of the Arnold Arboretum of
Harvard University. Publ. Arnold Arbor. 6. 1: 1-782. 1914;
I-542. I917.
Compiled under the direction of Charles Sprague Sargent.
Ule, E. Unter den Indianern des nérdlichen Amazonas. Verh.
Naturw. Vereins Hamburg III. 21: LX XVI, LXXVII._ 1914.
A botanical and anthropological journey.
Uphof, J. C. T. Die Waldflora im Staate Washington. Vegations-
bilder XIII. 7: 17 pp. + pl. 37-42. 1920.
Urban, I. Novitates haitienses. Notizbl. Bot. Gart. und Mus. Berlin-
Wipe 7: 495-499. 1 Mr 1921
new species in ei (3), pie (1), Cassia (1), Sebastiana (1).
‘sue na and Eugenia (1).
30 INDEX TO AMERICAN BOTANICAL LITERATURE
Urban, I. Uber die Malvaceen-Gattung Montezuma Moc. et Sessé.
Notizbl. Bot. Gart. und Mus. Berlin-Dahlem 7: 543, 544. 1 Mr
1921.
Wakefield, E. M. Mosaic diseases of plants. West Indian Bull. 18:
197-206. [1921.]
Walker, E.R. The gametophytes of Equisetum laevigatum. Bot. Gaz.
471: 378-391. pl. 23, 24 +f. 1-3. 16 My 1921.
Walker, J.C. Onion smudge. Jour. Agr. Research 20: 685-721. pl.
80-85 +f. I-10. 1 F 1921
Walker, J.C. Rust of onion followed by a secondary parasite. Phyto-
pathology 11: 87-90. f. 1, 2. F 1921.
Weatherby,C.A. Aformof Jlexopaca. Rhodora 23:118, 119. 19 My
1921.
Weir, J. R. Thelephora terrestris, T. fimbriata, and T. caryophyllea
on forest tree seedlings. Phytopathology 11: 141-144. pl. 5. Mr
1921.
Wells, B. W. Evolution of zoocecidia. Bot. Gaz. 71: 358-377. pl. 21,
22. 16 My 192I. :
Wells,B. W. A phenomenal shoot. Science II. 54: 13,14. 1 Jl 1921.
Paulownia tomentosa.
Weston, W. H. Another conidial Sclerospora of Philippine maize.
Jour. Agr. Research 20: 669-684. i. 76-79. 1 ¥F 1921.
Whaley, W. Properties of wood. A chapter from Lucien Chancerel’s .
Botanique Forestiére. Jour. For. 19: 414-420. Ap-1921.
White, O. E. The pollination of flowers. Brooklyn Bot. Gard. Leafl.
Qn. 3,4: 1-15. 4 My 1921.
Wiegand, K. M. Amelanchier amabilis, a new name. Rhodora 23:
48. 5 Ap Ig2t.
Wiegand, K. M. Echinochloa in North America. Rhodora 23: 49-
65. 20 Ap 1921.
Includes E£. paludigena sp. nov. and several new varieties.
Wieland, G. R. Monocarpy and pseudomonocarpy in the cycadeoids.
Am. Jour. Bot. 8: 218-230. pl. 9-12. 30 Ap 1921.
Williams, C. B. Report on the Froghopper Blight of sugar-cane in
Trinidad. Mem. Dept. Agr. Trinidad and Tobago 1: 1-170. i.
I-11 +f. 1-32. Ja 1921.
Considers also ‘‘The Root Disease of Sugar Cane” caused by fungi.
Yunker, T. G. A Curious abnormality in Cuscula cuspidata. Am. Bot.
27: 48,49. My 1921. [Illust.]
BuL_. TORREY CLUB VOLUME 49, PLATE I
’
ae *g
AR, tg
SMEAR:
PLANTS OF JAMAICA
PESSIN «. EPIPHYLLOUS
Vol. 49 No. 2
BULLETIN
OF THE
TORREY BOTANICAL CLUB
FEBRUARY, 1922
Miscellaneous notes on plants of Southern California—l
Puitip A. Munz AND IvAN M. JOHNSTON
Recent botanical collecting has added to our knowledge of the
flora of Southern California and has made possible the presenta-
tion of the notes given in this paper. Collections cited as having
been made by ‘‘M & H” were by Munz and Harwood, those by
“M, J & H” were by Munz, Johnston and Harwood; other collec-
tions are not abbreviated. Specimens of all the plants mentioned
are in the C, F. Baker Herbarium of Pomona College.
NOTHOLAENA SINUATA var. INTEGERRIMA Hook.
Notholaena sinuata var. integerrima Hook. Sp. Fil. 5: 108. 1864.
Locally abundant on a rocky hillside in a gulch back of the
Bonanza King Mine, Providence Mountains, Mohave Desert,
M & H3550and M,J@H 4224. Our material is a fine match for
the figure given by Eaton (Ferns of North America pl. 39, f. 1.
1879). New to California.
WoopsIA SCoPpULINA D. C. Eaton .
Woodsta scopulina D. C. Eaton, Canadian Nat. II 2:90. 1865.
This fern, infrequent in the higher mountains of the middle
and northern parts of the state, must be accredited a place in the
fern-list of Southern California, as a few plants were collected in
the Providence Mountains several miles south of the Bonanza
King Mines. The single colony discovered, M, J & H 4212, was
growing in the shelter of boulders in the bed of a dry gulch in the
lower portions of the pinyon belt. The soil in which the plant
{The BuLLETIN for January (49: 1-30. pl. z) was issued March 21, 1922.]
51
32 MuNz AND JOHNSTON: PLANTS OF CALIFORNIA
grew was damp, not saturated, gravel; a water hole was situated
a few hundred feet down the canyon and there was apparently a
meager supply of water from underground sources. On the
adjacent canyon walls were such typical xerophilous ferns as
Notholaena tenera, M, J & H 4232; N. Parryi, M, J & H 4236;
Chetlanthes Feet, M, J & H 4215; C. Covillei, M, J & H 4o15;
Gymnogramme triangularis, M, J & H 4021; and Pellaea compacta,
M, J & H 4ozs.
NOTHOLAENA CALIFORNICA D. C. Eaton
Notholaena californica D. C. Eaton, Bull. Torrey Club 10: 27.
188
The junior author collected a few plants of this fern high upon
a precipitous south-facing cliff at Deadman Point, about ten miles
southeast of Victorville. The collected plants were associated
with an abundance of Notholaena Parryi and with fewer Cheilanthes
Covillei and Gymnogramme triangularis. This collection now
marks the most northern station for this species, at least in Calli-
fornia, and at the same time makes the species first known as an
element of the Mohave Desert flora. Maxon in his recent dis-
cussion ( Contr. U. S. Nat. Herb. 17:603. 1916) has cited many
specimens that show the distribution of this neat little plant.
Johnston (Bull. So. Cal. Acad. 17: 64. 1918) has recorded a
second coastal and what is at the same time the most westerly
known station on the mainland.
TRIODIA MUTICA (Torr.) Benth.
Tricuspis muticus Torr. U.S. Rep. Expl. Miss. Pacif. 4: 16: 1857-
Triodia mutica Benth.; Wats. Proc. Am. Acad. 18 : 180. 1883.
Abundant on a eke pinyon-clad hillside near the Bonanza
King Mine, in the Providence Mountains, M, J & H 4132. Pre-
viously known in California only from a collection made in the
central Sierra Nevada (cf. Jepson, Fl. California 141. 1912).
Determination by Mrs, Chase.
AGAVE UTAHENSIS Engelm. —
A gave utahense Engelm. Bot. King. Exp. 497. 1871.
Common in rocky places in the pinyon belt of the Providence
Mountains, M, J & H 4302. This is the “‘ Agave sp.” reported by
Munz AND JOHNSTON: PLANTS OF CALIFORNIA 33
Brandegee (Zoe 5: 153. 1903). Mulford has cited a collection
made by Parish at Ivanpah (Rep. Missouri Bot. Gard. 7: 78.
1896).
Yucca BACCATA Torr.
Yucca baccata Torr. Bot. Mex. Bound. Surv. 221. 1859.
The presence of this species in the Providence Mountains was
reported recently by Mr. Parish (Bot. Gaz. 65: 336. 1918) and at
’ a much earlier date by Dr. Engelmann (Trans. St. Louis Acad. 3:
44. 1873), so that all that remains for us to do is to add a few
notes on its habits. This species and Y. mohavensis are very
common in the vicinity of the Bonanza King Mine but, although
growing in the same region, differ widely in habitats and
habits, as well as in technical characters. Y. baccata grows on
rocky, sunny hillsides in the pinyon belt and is a very late-flowering
plant; for but few individuals were in flower during our visit to
the mountains from May 21 to 24, 1920. On the other hand,
Y. mohavensis grows along the foot of the mountains, below the
pinyon belt, and flowers very early; for on the May visit fruit
was set, flowering having been observed on an earlier visit during
the last week in March. Our observations fully substantiated
Mr. Parish’s statement that Y. baccata was recognizable in the
field by the lighter color of the leaves and the universal acaulescent
abit.
Structurally the two species are very distinct. Y. baccata has
large campanulate flowers, whose perianth segments do not spread
from the base but remain erect and appressed about the ovary
for a short distance before spreading, thus suggesting the tubular
condition characteristic of the genus Samuela Trel. In
mohavensis the shorter segments spread from the very base and
are more curved, thus making the flowers globose in shape.
The meager material that is available of Y. baccata suggests
that there may be a major and a minor flower form. Mr. Brande-
gee’s collection, reported as Y. mohavensis (Zoe 5: 153. 1903),
Lemmon’s 1884 collection at Ft. Mohave, and our M, J & H 4174
all represent plants with conspicuously large flowers, the perianth-
segments of which vary between 8 and 9.5 cm. in length. On the
other hand, Parish 10281 from the north end of the Providence
Mountains and our M, J & H 4113 from the south end of these
34 Munz AND JOHNSTON:’ PLANTS OF CALIFORNIA
mountains have strikingly smaller corollas, the perianth-segments
measuring 6-7 cm. in length. Under the original description
Dr. Torrey quotes Bigelow to the effect that this species has
perianth lobes 24 or 3 inches (6.25-7.5 cm.) long, while Wooton
and Standley (Contr. U.S. Nat. Herb. 19: 135. 1915) give them as
5-8 cm. long, and Trelease (Rep. Missouri Bot. Gard. 13: I10.
1902) says that they are “about 75 mm.” long; these facts would
indicate that our large-flowered plants have flowers large for the
species. It is to be hoped that future collections will be made
with especial attention to this species, in order to ascertain the
exact nature and extent of the floral variation here indicated.
In passing, it should be noted that the style-characters given by
Trelease (Rep. Missouri Bot. Gard. 13: 46. 1902; and Contr. U.S.
Nat. Herb. 23: 91. 1920) and the flower- and foliage-characters
given by Sargent (Gard. & For. g: 104. 1896) seem to be il-
lusionary,and that the distinctness between our western Y. mo-
havensis and the Texan plant originally called Y. baccata var.
macrocarpa (Torrey, l.c.) is open to strong doubt. We would use.
the name Y. macrocarpa (Torr.) Coville (Contr. U. S. Nat. Herb.
4: 202. 1893) in preference to Y. mohavensis Sarg., were it not
for Engelmann’s Y. macrocarpa (Bot. Gaz. 6: 224. 1881); the de-
notation of Y. macrocarpa Engelm. is uncertain from the literature
(Trelease, op. cit.98. 1902; Sargent, /.c.; and Engelmann, Bot.
Gaz. 7:17. 1882); and it is thought best to make no change in >
our usage until it has been definitely decided whether Engel-
mann’s species be a synonym of Y. Schottii or actually identical
with Torrey’s var. macrocarpa, as Wooton and Standley have
treated it. In other words we use Y. mohavensis in a wide sense
and include therein Y. baccata macrocarpa Torr., though appreci-
ating the fact that Y. mohavensis may not be the correct name for
such a concept
Dr. Merriam (N. Am. Fauna 7: pl. 12, I4. 1893) has given
two plates that show the very different forms which characterize
Y. baccata and Y. mohavensis. Although his plate of the latter
represents the dominant and ordinary phase of that species, yet
we observed a phase with unbranched trunks and usually longer
leaves, whose habit is strongly suggested by the plants of a very
different species, figured in pl. 3 of the third report of the Missouri
Botanical Garden.
MuNz AND JOHNSTON: PLANTS OF CALIFORNIA — 35
NAIAS FLEXILIS (Willd.) Rostk. & Schmidt
Caulinia flexilis Willd. Abh. Akad. Berlin 95. 1803.
Natias flexilis Rostk. & Schmidt, Fl. Sedin. 382. 1824.
Previously reported by Davidson from Soldiers’ Home near
Los Angeles; collected in 1919, Street & Williams 2688, at Laguna
Lakes near Laguna Beach, about fifty miles southeast of earlier
records. The shallow pools known as the Laguna Lakes have a
rather interesting lot of aquatic plants, such as Zannichellia
palustris, Echinodorus cordifolius, Marsilea vestita, and Elatine
californica.
ERIOGONUM APICULATUM Wats.
Eriogonum apiculatum Wats. Proc. Am. Acad. 17: 378. 1882.
This species, said by Jepson (FI. California 407. 1914) to
occur on “Mt. San Jacinto, 7800 to 8200 feet’? and on ‘“Cuya-
maca Mt.,”’ has been collected at 5,500 feet, August 5, 1918, Mrs.
Mary F. Spencer 989, in open woods on Palomar Mountain in San
Diego County. This station, in addition to being of altitudinal
interest, is considerably west of the others mentioned.
MOLLUGO VERTICILLATA L.
Mollugo verticillata L. Sp. Pl. 89. 1753
Said by Jepson (FI. California 460. 1914) to be ‘‘sparingly
naturalized”’ and given as collected by Davidson in Los Angeles.
Abrams (FI. Los Angeles 127. 1917) names also Laguna in Orange
County. Wecan report this plant from near Beaumont, on the
line between Riverside and San Bernardino Counties, Munz 2614,
March 1919, growing on a dry hillside in the chaparral at about
2000 feet.
HERNIARIA CINEREA DC.
Herniaria cinerea DC. FI. Fr. Suppl. 375. 1815.
This species, naturalized from southern Europe, can now be re-
ported from Southern California, Munz & Johnston 4406, Pomona
College Campus, Claremont, where it is fairly well established, but
threatened by impending grading and building. Our plants are
well matured and some of them had ripened their seeds by the
end of March.
ANEMONE TUBEROSA Rydb.
Anemone tuberosa Rydb. Bull. Torrey Club 29: 151. 1902.
Previously recorded from California as collected in the Pana-
mint Mountains, 1891, Coville & Funston; and in 1849 by Fremont,
36 Munz AND JOHNSTON: PLANTS OF CALIFORNIA
locality not given. It was found abundant at the eastern base of
the Providence Mountains on rocky slopes just below the pinyon
belt, M. & H 3555 and M, J. & H 4207.
Myosurus CUPULATUS Wats.
Myosurus cupulatus Wats. Proc. Amer. Acad. 17: 362. 1882.
Canyon side in the Upper Sonoran Zone at 4,000 feet, Provi-
dence Mountains, M, J & H 4152, May 22, 1920; at this time
the plants were quite dead, having matured their seeds. Pre-
viously known only from a few collections in Arizona.
JEPSONIA PaRRyI (Torr.) Small
Saxtfraga Parryi Torr. Bot. Mex. Bound. Surv. 69. pl. 25. 1859.
Jepsonia Parryi Small, Bull. Torrey Club 23:18. 1896.
In January, 1921, Mr. Gordon Nicholson detected a very
populous colony in a small side canyon running up into the Santa
Ana Mountains from the Santa Ana Canyon, and very near the
Orange-Riverside County line. The plants grew at about 1,000
ft. altitude in clay soil, on a rather open grassy slope facing west.
We have only specimens in leaf, but these check closely with plate
25 in the Mexican Boundary Survey Report. The type came
from San Diego County where it has been collected several times
since, but outside of that county it has been known from but two
collections, both in Riverside County. Dr. Hall found it at
Double Butte near Winchester, and Mr. Parish has reported it
from Palm Springs (Muhlenbergia 3: 123. 1907).
LINUM SPERGULINUM Gray
Linum spergulinum Gray, Proc. Am. Acad. 7: 333. 1868.
This plant, previously known from Middle California, has been
collected in Pine Hills, San Diego County, Mrs. Mary Spencer
15906, at 4,200 ft., on June 25, 1920. The appendage on the petal .
is as described by Gray.
HoLacanTHA Emoryi Gray
Holacantha Emoryi Gray, Pl. Nov. Thurb. 310. 1855.
Two collections were made: one, M, J & H 4295, was eight
miles west of Ludlow in a sandy wash along which it occurred for
several miles; the other, M, J & H 4300, was five miles east of
Amboy on flat stony desert. Both these specimens are more
‘MUNZ AND JOHNSTON: PLANTS OF CALIFORNIA 37
compact than Emory’s figures in the Mexican Boundary Survey,
but a specimen sent us by Mr. S. B. Parish, collected by a Mr.
Childs at the Hayfields in the Chuckawalla Valley, is of more
open habit than the one in Emory’s plate. Both of our collec-
tions were very near the road and from large conspicuous shrubs,
three to four feet high, forming wide gray masses of interlacing
thorns; it seems strange that such conspicuous plants so near the
road have been collected so seldom. Our Ludlow station is no
doubt the one reported by Mrs. Ferris (Bull. So. California Acad.
18: 13. 1919), but the one near Amboy is some distance from
those reported by Davidson (ibid. 19: 55. 1920).
TRAGIA RAMOSA Torr.
Tragia ramosa Torr. Ann. Lyc. N. Y. 2: 245. 1826.
Locally common in a gravelly and stony canyon bottom in the
Lower Sonoran Zone at the eastern base of the Providence Moun-
tains, M, J & H. 4219. The first collection that has been re-
ported from California.
ABUTILON PARVULUM Gray
Abutilon parvulum Gray. Pl. Wright. 1:21. 1852.
In rocky ground at the base of the Providence Mountains near
a deserted mining camp several miles south of the Bonanza King
Mine; rather common locally; M, J & H 4206. First record
from California.
FRASERA Parryl Torr.
Frasera Parryt Torr. Bot. Mex. Bound. Surv. 156. 1859.
Collected June 7, 1919, in the foothills of the San Gabriel
Mountains between Live Oak and San Dimas Canyons, by Mrs.
Fitch, C. F. Baker Herbarium No. 6594. Long ago reported from
“east of . . . Los Angeles’”’ by Brewer (Bot. California 1: 484.
1876), but not again detected on the south slope of the San
Gabriel range until the present. Frequent in the San Bernardino
and San Jacinto ranges, where it inhabits pine-clad slopes of the
Transition Zone. The present collection was a single plant and
grew in the lower chaparral belt at about 1700 feet altitude.
SALVIA MOHAVENSIS Greene
Audibertia capitata Gray, Proc. Am. Acad. 7: 387: 1868.
Salvia mohavensis Greene, Pittonia 2: 235. 1892.
38 Munz AND JOHNSTON: PLANTS OF CALIFORNIA
Collected on the crest of a rocky outlying spur of the Turtle
Mts. southwest of Needles, M & H 3509. It was also found rather
commonly on rocky places in the Providence Mountains, which
constitute the type locality. The Parish brothers long ago col-
lected it in the mountains near Camp Cady. The Turtle Mountain
station was at only about 1,200 feet altitude; plants were beginning
to flower there on the first of April. In the Providence Moun-
tains the species was observed from the base to near the crest,
from 3,000 to 7,000 feet altitude, where only a few plants were in
bud in the last week in May. Field observation would indicate
that the species is restricted to rocky places, paying little attention
to the lines between the Sonoran Zones.
CRYPTANTHA MARITIMA Greene
Krynitzskia ramosissima Greene, Bull. Cal. Acad. 1: 203. Au 1885.
Not Krynitzkia ramosissima Gray, Proc. Am. Acad. 20: 277.
Ja 1885.
Krynitzkia maritima Greene, Bull. Cal. Acad. 1: 204. Au 1885.
Cryptantha maritima Greene, Pittonia 1: 116. 1887.
Attention should be called to the fact that the plants currently
going under the name of C. ramosissima Greene should properly
be designated as C. maritima. As Coville indicated twenty-nine
years ago (Contr. U.S. Nat. Herb. 4: 165. 1893) K. ramosissima is
an untenable name for the plant in question, Gray having originally
proposed it as a substitute, on the grounds of applicability, for
the specific name racemosa. Gray clearly indicated that Eri-
trichium racemosa Wats. and K. ramosissima Gray were synony-
mous, but Greene took the latter name and gave it a wholly new
connotation, making it cover certain misdetermined specimens
cited by Gray under the name K. ramosissima. K. ramosissima
Greene is a redefinition of K. ramosissima Gray, and as this latter -
unquestionably a synonym of C. racemosa, the former, technically
having the same type as the latter, since it is merely a redefinition
of it, cannot be considered more than an improper interpretation
of K. ramosissima Gray. Neither the American nor the Vienna
Code will justify the use of K. ramosissima in the sense that it is
currently used in today!
C. maritima was described from Guadalupe Island specimens,
but despite its remote insular habitat, like many other species,
Munz AND JOHNSTON:,PLANTS OF CALIFORNIA 39
it is identical with the mainland form. Gray (in the supplement
of the Synoptical Flora, p. 428), attempted to distinguish between
the insular plant and C. ramosissima Greene, but when applied to
a series of any size, his characters utterly fail to divide the two
forms and certainly do not segregate the material into anything
suggesting geographical lines. Vasey and Rose (Proc. U. S.
Nat. Mus. 11: 532. 1888) and Brandegee (Bot. Gaz. 27: 453.
1899) have expressed doubts as to the distinctness of C. ramosis-
sima and C. maritima, but now with a fine series for study, we
feel that their doubts can be increased to the point of certainty.
Every character presented by the Guadalupe specimens, or for
that matter by any of the coastal plants, can be exactly duplicated
in specimens from the desert interior. Though it is unfortunate
that a name such as ‘‘ maritima” should be applied to a plant so
characteristic of the driest portions of the desert area, yet there
is no technical reason for coining a new name or for accepting any
other old one.
Among the California representatives of the genus, C. maritima
is unique in its possession of dark reddish stems. ‘This species,
along with C. recurvata, possesses but two ovules, a condition dif-
ferent from that found in all other species of the genus (cf. Bran-
degee, /.c.).
’ Cryptantha gracilis var. Hillmanii (Nels. & Ken.) comb. nov.
Cryptantha Hillmaniui Nels. & Ken. Proc. Biol. Soc. Washington
19: 157- 1906.
This plant must now be accredited a place in the California
flora. It would seem that it enters the state only in the region of
the Providence Mountains, for the only California collections come
from them; Mrs. Brandegee got it at Barnwell and we made a
fine collection, M, J & H 4222, near the Bonanza King Mine.
C. Hilimanii is very close to C. gracilis Osterhout (Bull. Torrey
Club 30: 236.. 1903) and at most is but a poor variety of the
latter species. Abundant collections of the species may cause
C. Hillmanii to be reduced outright, but the single isotype of
C. gracilis in the University of California collection seems to differ
from all the specimens of C. Hillmanii in its lower and more slender
habit and less congested inflorescence; therefore, for the time
40 MuNZz AND JOHNSTON: PLANTS OF CALIFORNIA
being, C. Hillmann can be retained in varietal rank. The type
of C. gracilis came from Glenwood Springs in the mountains of
Colorado and it is entirely possible that the differences detected
are environmental in origin. Among California cryptanthas,
C. Hillmanii can be recognized by its single smooth nutlet and
usually fulvous, ——- pilose and inconspicuously bristly “—
v Panctomon Munzii Johnston, sp. nov.
Plants with several coarse, erect, loosely tufted glabrate stems
that become at least five dm. high; leaves all opposite, entire and
glabrate; basal leaves ovate. or Tanicecla te: spatulate with winged
petioles that about equal the blade, becoming 7 cm. long and
2.5 cm. wide; the lower cauline leaves oblanceolate, the upper
ones broadly sessile and lanceolate, while the leaves of the in-
florescence are minute and linear-subulate; inflorescence narrow,
the flowers in strict one- to three-flowered cymules; corolla bright
red, 2 cm. long, narrowly funnelform-tubular, evenly though but
slightly ampliated upward, strongly and conspicuously keyed
glabrous within; upper two lobes of the corolla erect, about
long, united foi about two-thirds of their length, lower hee
lobes of the corolla strongly reflexed, about 6 mm. long, united
for nearly ‘half their length; anther-sacs glabrous, iia apets rugu-
lose or papillose, adnate if at all only near the base, ovate-oblong,
2-2.5 mm. long, their inner sides paralleling each other or forming
a small angle; dehiscent by a slit extending between two-thirds
and three-fourths the way to the base, sharply dentate along the
line of dehiscence; sterile filament glabrous, somewhat flattened,
emarginate; sepals broadly ovate, acute, 3-5 mm. long, scarious
margined; pedicels about as long as the sepals; fruit unknown.
Known only from the type specimen, collected in May, 1920,
Munz, Johnston, & Harwood 4271, on a high exposed ridge in the
pinyon belt near the Bonanza King Mine on the east slope of the
Providence Mountains, Southeastern California. The type is
No. 7534 in the Baker Herbarium of Pomona College.
Certainly of the Barbati and probably nearest P. barbatus
Torreyi (Benth.) Gray, but very different from it in its color,
which is a brighter, lighter, and more yellowish red, in the meas-
urements of the corolla, and in the size as well as non-divergence
of the anther-sacs. P. Munzii is a lower, stouter, and less grace-
ful plant than most of the forms of P. barbatus (Cav.) Roth.
From P. Eatoni Gray and from P. subulatus Jones, the other
two red penstemons found in this part of the desert area, both of |
MUNz AND JOHNSTON: PLANTS OF CALIFORNIA 41
which are in the Centranthifolii, this species can be quickly sep-
arated by its unique tone of red and by its smaller, stouter, de-
cidedly bilabiate corollas. It isa well marked species and a
fitting addition to the interesting penstemon flora ef the Providence
Mountains.
» Penstemon Clevelandii var. Stephensi (Brandegee) comb. nov.
Penstemon Stephensi Brandegee, Zoe 5: 151. 1903.
During one of our trips to the Providence Mountains we made
a small collection of this little known plant, which is apparently
rare in the type region; for Mr. Brandegee informs us that he
collected only enough for the type sheet, a full one, while our
collection, M, J & H 4274, was enough for only a few duplicates.
A critical study of our collection and of the type seems to in-
dicate that P. Stephensi finds a very close relative in P. Cleveland
Gray, a species restricted to the canyons about the Colorado Des-
ert and in the mountains to the west of it. Our collection of P.
Stephensi, as well as the type, is so similar to certain collections
of P. Clevelandi (e.g., Hall 1160 and 2149 and Parish Bros. 1216)
that as far as gross aspect, size and vegetative characters are con-
cerned, they might be of one and the same collection. There is,
however, one character which separates the two collections of P.
Stephensi from all of the many collections of P. Clevelandi; this is the
lack of bearding on the sterile filament. Though the corollas of
the two forms are alike in size, color, and shape, the sterile filament
in P. Stephensi is absolutely bald, while it is densely bearded in
P. Clevelandi. Appreciating, therefore, that we are concerned
here only with a unit-difference, we feel that P. Stephensi is noth-
ing but a geographic race of P. Clevelandi and, because of it, worthy
only of a subordinate rank.
PENSTEMON PALMERI Gray
Penstemon Palmeri Gray, Proc. Am. Acad. 7: 379. 1868.
We found this species to be the most common and most
spectacular penstemon in the Providence Mountains, M, J & H.
4276. About the Bonanza King Mine it was common about the
foot of the mountains and especially so in small draws, where it
grew in large colonies. This being our first acquaintance in the
42 MuNz AND JOHNSTON: PLANTS OF CALIFORNIA
field with the typical plant, and knowing quite intimately the
low, much-branched, few-flowered montane plant, currently going
under the name of P. Palmeri, we were much surprised to discover
that this tall, virgate, many-flowered, desert plant was the typical
form of P. Palmeri. While unquestionably, the montane plant
is a close ally of the taller interior form, and as herbarium material
shows, intergrades with it, yet we feel that so pronounced an ex-
treme ought to be given nomenclatorial recognition, and so, for
us, the mountain form becomes:
” Penstemon Palmeri var. Grinnellii (Eastw.) comb. nov.
Penstemon Grinnellu Eastw. Bull. Torrey Club 32: 207. 1905.
The type of this form came from Mt. Wilson and is the form
rather common in rocky ground in the pine belt, and to a less ex-
tent in the upper chaparral belt, in the mountains of Southern
California from the San Jacinto Range northward. With this
restriction the species itself becomes strictly deserticolous. In
its extreme the variety is characterized by its looser, shorter and
fewer flowered inflorescence, lower and more slender stature and
non-glaucous leaves.
PENSTEMON CALCAREUS Brandegee
Penstemon calcareus Brandegee, Zoe 5: 152. 1903. Not Jones,
1908.
Penstemon desertorum Jones, Contr. W. Bot. 12:59. 1908.
We succeeded in making a fair-sized collection of this peculiar
and interesting little plant, M, J & H. 4154. Thespecies is known
from but two other collections, all of them from the Providence
Mountains, where it grows in crevices of rocks on vertical cliffs.
P. desertorum Jones, the type of which came from Kelso on the
west slope of the mountains, is identical with Mr. Brandegee’s
species; for isotypes of the former plant are exact matches for
plants on the type sheet of the latter; in fact, Mr. Brandegee’s
type, Mr. Jones’ Kelso collection and our plants form a remarkably
uniform series in aspect and size, as well as in structural characters.
P. calcareus Jones (Contr. W. Bot. 12: 60. 1908) is a homo-
nym and appears to be a synonym of P. petiolatus Brandegee
(Bot. Gaz. 27: 455. 1899).
MuNz AND JOHNSTON: PLANTS OF CALIFORNIA 43
’ Penstemon antirrhinoides var. microphyllus (Gray) comb. nov.
Penstemon microphyllus Gray, Pac. R. R. Rep. 4: 119. 1856.
Penstemon Plummerae Abrams, Bull. Torrey Club 33: 445. 1906.
In California this interior form of P. antirrhinoides Benth. is
known only from the Providence Mountains. Mr. Brandegee
collected and reported it (Zoe 5: 151. 1903) as P. antirrhinoides,
while we found it to be rather common on the rocky sides of a
small canyon back of the Bonanza King Mine. Although quite
familiar with P. antirrhinoides, as represented on the coastal slopes,
we could not detect in this desert plant any peculiarities of habit
or size, though while observing the plant in the field we agreed
that there was some intangible difference, a difference which was
later discovered to reside in the grayish hue resulting from a
cinereous pubescence.
This shrub was described by Gray from flowerless specimens
collected by Bigelow on the ‘‘Williams Fork of the Colorado.”
Based on insufficient material, the plant has remained an obscure
one: It is not surprising, therefore, that Dr. Abrams, upon
meeting:so obscure a plant, treated it as though it were unde-
scribed. It can hardly be doubted that P. Plummerae and P.
microphyllus are synonyms, for Dr. Gray’s meager description
applies completely to P. Plummerae, while the types of both are
from the same phytogeographical area. Significant evidence is —
found also in the fact that this is the only shrubby Penstemon re-
vealed by recent collecting in Arizona.
Dr. Abrams has indicated the characters separating this form
from the typical P. antirrhinoides, which ranges on the hillsides
near the coast from San Pedro Martir Mountains northward to
near San Bernardino, and which is characterized by its very
broadly ovate sepals and glabrate or puberulent twigs. The
variety microphyllus, on the other hand, is strictly an interior
form, ranging from the Providence Mountains eastward into
western Arizona, and is marked by its distinctly cinereous twigs
and oblong-ovate or ovate-lanceolate sepals. While the three-
fold difference residing in the sepals, pubescence and range, argues
much for specific distinctness of these plants, the relationship is
so close and so obvious that we feel it best to treat them only
as well marked geographic forms.
44 MuNnz AND JOHNSTON: PLANTS OF CALIFORNIA
PENSTEMON ALBOMARGINATUS Jones
Penstemon albomarginatus Jones, Contr. W. Bot. 12:61. 1908.
We are able to report the first collection in California of this
distinct species. We obtained the plant near Lavic, where it
grew in the deep sand of a wash that ran up between the tongues
of the lava-flow surrounding Mt. Pisgah. The wash was a stone’s
throw from the Sante Fe Trail and near the grade crossing west of
Lavic. The species is very aptly named, for the white margining
to the leaves and sepals is very striking. It is apparently rare
and we can get trace of but three other collections: Good Springs,
Nevada, and Yucca, Arizona, reported by Professor Jones; and
Jean Station, Nevada, collectcd by Mrs. Brandegee (specimen
in the herbarium of the University of California). Professor
Jones collected the type in sand and, as our plant grew in similar
material, the species is probably arenicolous and is to be looked for
in sandy places. The type and our collection, M, J & H 4204,
were both collected in May and were at the time setting seed.
vy Aplopappus Gooddingi (Nels.) comb. nov.
Sideranthus Gooddingi Nels. Bot. Gaz. 37: 266. 1904.
A single plant, M & H 3603, collected on a gravelly hillside
about eleven miles west of Needles. The present collection is the
first one from California, but is remarkably like the typical plants
from Nevada. It is characterized by its perennial root and tall
strict stems and large and multiradiate heads, which are up to
16 mm. wide. A. gracilis the nearest relative of the desert regions,
is a lower annual with ascending stems, smaller (1 cm. or less wide)
heads and fewer rays, eight to sixteen; its leaves are more setose
than in A. Gooddingi.
POMONA COLLEGE,
CLAREMONT, CALIFORNIA
INDEX TO AMERICAN BOTANICAL LITERATURE
1910-1921
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 o: — origin, or laboratory methods are not included, and
no attempt is made to index the literature of SE a An occasional iga.tine is
made in favor of some eae appearing in an American periodical which is devoted
wholly to botany. Reprints are not men aaa unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to — or omissions, their kindness will be appreciated.
Index is ee monthly on cards, and furniched in coi form to subscribers
at ‘i co of three cents 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
Knuth, R. Zwei nieue Dioscorea aus Brasilien. Notizbl. Bot. Gart.
und Mus. Berlin-Dahlem 7: 538, 539. 1 Mr 1921.
Korstian, C. F., Hartley, C., Watts, L. F., & Hahn, G. G. A chlorosis
of conifers corrected by spraying with ferrous sulphate. Jour.
Agr. Research 21: 153-171... 1-4. 2 My 10921.
Kranzlin, F. Orchidaceae Dusenianae novae. Arkiv Bot. 168: 1-30
6 Ap 1920.
Includes new species from Brazil (23), and Mexico (4), the latter collected by
Bourgeau.
Lee, H. A. The increase in resistance to citrus canker with the ad-
vance in maturity of Citrus trees. Phytopathology 11: 70-73.
F 1921.
Lehman, S. G. Soft rot of pepper fruits. Phytopathology 11: 85-
87. F 1921.
Lindstrom, E. W. Concerning the inheritance of green and vellane
pigments in maize seedlings. Genetics 6: 91-110. Ja 1921.
Livingston, B. E., & Hawkins, L.A. The water-relation between plant
and soil. Carnegie Instit. Washington Publ. 204: 1-48. Gf I-3.
10 Mr 10915. ;
45
46 INDEX TO AMERICAN BOTANICAL LITERATURE
Lloyd, F. E. Guayule (Parthenium argentatum Gray). A_rubber-
plant of the Chihuahuan desert. Carnegie Instit. Washington
Publ. 139: 1-213. pl. 1-46 + f. 1-20. 27 Jl 1911.
MacDougal, D. T. A new high temperature record for growth. Sci-
ence II. 53: 370-372. 15 Ap 1921.
MacDougal, D. T., & Spalding, E. S. The water-balance of succulent
plants. Carnegie Instit. Washington Publ. 141: 1-77. pl. 1-5 +f.
1-16. 29D 1910.
Macpherson, G. E. Comparison of development in dodder and morn-
ing glory. Bot. Gaz. 71: 392-398. pl. 25-27. 16 My 1921.
Malme, G. O. A.-N. Asclepiadaceae riograndenses adjectis notulis de
ceteris Asclepiadaceis in Brasilia extratropica, Uruguay et Misiones
collectis. Arkiv Bot. 16: 1-34. 23 D 1920.
Includes new species in Roulinia (1), Calostigma (1).
Mameli de Calvino, E. Estudios anatomicos y fisiologicos sobre la
cafia de azucar en Cuba. Estac. Exper. Agron. Santiago, Cuba,
Bol. 46: 1-49. f. 1-21.
Marsh, C. D., & Clawson, A. B. Poisonous properties of the whorled
_ milkweeds Asclepias. pumila and A. .verticillata var. Geyeri. U.S.
Dept. Agr. Bull. 942: 1-14. pl. 1-3 +f. 1, 2. 4 My 192%.
Mason, T. G. A note on some recent researches on the cotton plant in
the West Indies, with special reference to St. Vincent. West Indian
Bull. 18: 184-197..pl..1,,2. [1921.]
Mason, T. G. The water-balance of the plant and its significance in
crop production. West Indian Bull. 18: 157-184. [1921.]
Massey, L. M. Experimental data on losses due to crown-canker ot
_rose. Phytopathology 11: 125-134. Mr 1921.
Matz, J. Infection and nature of the yellow stripe disease of cane
(mosaic, mottling, etc.). Jour. Dept. Agr. Porto Rico 3°: 65-32.
f. 1-11. O1919. xe
Matz, J. Obeservaciones en la gomosis de la cana en Puerto Rico.
Revista Agr. Puerto Rico 64: 33-39. 30 Ap 1g2t. [Illust.]
[Maxon, W. R., & Killip, E. P.] Botanical exploration in Jamaica
[1920]. Smithson. Misc. Coll.-72°: 49-54. f. 61-66. 1921.
Mayes, J. The practical control of weeds in Missouri meadows and
pastures. Bull. Missouri Board Agr. 18: 1-36. Je 1920. [Illust.]
McAtee, W. L. Notes on Viburnum and the assemblage Caprifoli-
aceae. Bull. Torrey Club 48: 149-154. f. r. 25 My 1921.
McDougall, W. B. Thick-walled root hairs of Gleditsia and related
genera. Am. Jour. Bot. 8: 171-175. f. 1-3. 3 Ap 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 47
McLean, F. T. A study of the structure of the stomata of two species
of Citrus in relation to citrus canker. Bull. Torrey Club 48: 101-
106. f. r. 4 Ap 1921.
McLean, F. T., & Lee, H. A. The resistance to citrus canker of Citrus
nobilis and a suggestion as to the production Of resistant varieties in
other Citrus species. Phytopathology 11: 109-114. f. z. Mr 1921.
McNair, J.B. <A study of Rhus diversiloba with special reference to its
toxicity. Am. Jour. Bot. 8: 127-146. pl. 2+/f.1, 2. 3 Ap 1921.
McNair, J. B. The morphology and anatomy of Rhus diversiloba.
Am. Jour. Bot. 8: 179-191. pl. 3, 4. 30 Ap 1921.
McNair, J. B. The transmission of Rhus poison from plant to person.
Am. Jour. Bot. 8: 238-250. 24 My 1921.
Menaul, P. Note on the formation of hydrocyanic acid in plants.
Jour. Biol. Chem. 46: 297. Ap 1921.
Merriman, M. L. The receptacle of Achillea Millefolium L. Torreya
21: 21-24. f. 5: “Ap 192i.
Morse, W. J. The transference at potato late — ig insects.
Phytopathology 11: 94-96. F 1921.
Mousley, H. Further notes on the orchids of Hatley, Stanstead Co.,
Quebec, 1920. Canad. Field Nat. 34: 169-173. 12 My-1g21.
Muttkowski, R. A. Copper in animals and plants. Science II. 53:
453, 454. 13 My tgat.
Needham, J. G. A biological examination of Lake George, N. Y.
Sci. Mo. 12: 434-438. My 1921.
Nelson, J. C. Additions to the flora of western Oregon during 1920.
Torreya 21: 24-28. p 1921.
Oakley, R. A., & Evans, M. W. Rooting stems in timothy. Jour.
Agr. Research 21: 173-178. pl. 39, 40. 2 My 1921
Orton, C. R. Lightning injury to potato and cabbage. Phytopathol-
ogy 11: 96-98. f. z. F 1921.
Palmer, E. J. Botanical reconnaissance of southern Illinois. Jour.
Arnold Arbor. 2: 129-153. Ja 1921.
Peattie, D. C. An interesting habitat. Rhodora 23: 69-71. 20 Ap
Pennell, F. W. ‘‘Veronica’” in North and South America. Rhodora
23: I-22, 29-41. _ 1921.
Includes Veronica catenata sp. nov., and several new combinations.
Pieters, A. J., & Kephart, L.W. Annual white sweet clover and strains
of the biennial form. U.S. Dept. Agr. Dept. Circ. 167: I-21. f.
6. Ap eoat.
48 INDEX TO AMERICAN BOTANICAL LITERATURE
Piper, C. V. Two new legumes from Mexico and Costa Rica. Proc.
Biol. Soc. Washington 34: 41, 42. 31 Mr 1921
Phaseolus chiapasanus and Calopogonium ferrugineum sp. nov.
Pool, R. J., Weaver, J. E., & Jean, F.C. Further studies in the ecotone
between prairie and woodland. Univ. Nebraska Stud. 18: 1-47.
Fe taT7 1918.
Pritchard, F. J., & Porte, W. S. Collar-rot of tomato. Jour. Agr.
Research 21: 179-184. pl. 41-45. 2 My 1921.
Includes Verticillium Lycopersici sp. nov.
Puttemans, A. Gloeosporium Bombacis, n. sp. Bull. Soc. Path. Veg.
Perch vere. S 1920.
Puttemans, A. Sur /’Oidium du Chéne au Brésil. Bull. Sec. Path.
Veg. 7: 37-40. 1 Je 1920.
Rathbun, A. E. Methods of direct inoculation with damping-off fungi.
Phytopathology 11: 80-84. f. 1-3. F 1921.
Record, S. J. Further notes on intercellular canals in dicotyledonous
woods. Jour. For. 19: 255-266. Mr 1921.
Reed, H. S. Growth and sap concentration. Jour. Agr. Research
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Rehder, A. Azalea or Loiseleuria. Jour. Arnold Arbor. 2: 156-159.
Ja 19021.
Rehder, A. Philadelphus verrucosus Schrader spontaneous in IIlinois.
_ Jour. Arnold Arbor. 2: 153-156. Ja 1921. ;
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Ap rg2t. [Illust.] |
Richards, H. M. Acidity and gas interchange in Cacti. Carnegie
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Rusby, H. H. Guide to the Economic Museum of the New York
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Includes a few new combinations, such as Acyntha Kirkii (Baker) Rusby, page 23-
INDEX TO AMERICAN BOTANICAL LITERATURE 49
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-
50 INDEX TO AMERICAN BOTANICAL LITERATURE
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192Ts
Vol. 49 | : No. 3
BULLETIN
OF THE
TORREY BOTANICAL CLUB
MARCH, 1922
Quadripartition by furrowing in Sisyrinchium
CLIFFORD H. Farr
(WITH PLATE 2)
Since the publication of the author’s paper (5) on Magnolia
there have appeared a few contributions to the subject of the
quadripartition of pollen-mother-cells by furrowing. In 1918
Tackholm and Séderberg (24) presented an interesting discovery
of the occurrence of quadripartition and bipartition respectively
in two species of Aristolochia, a genus of Dicotyledons; Aristo-
lochia fimbriata was found to have successive bipartition, just
as Samuelson (19) had reported in this genus, but Aristolochia
Sipho was found to have quadripartition of the pollen-mother-
cells after the homoeotypic mitosis. It is stated that there is
a slight thickening of the spindle fibers after the heterotypic
division in the latter species, but no figures of quadripartition
are shown in this paper. In Vinca rosea the authors conclude
that quadripartition must occur because of the tetrahedral
arrangement of the microspores within the mother wall. It
will be remembered that Frye and Blodgett in 1905 (8) found
successive bipartition in the sister genus Apocynum, and the
microspores here are sometimes arranged tetrahedrally. The
authors (24) state that after the homoeotypic division in Aris-
tolochia Sipho walls are laid down as fine plates on the equator
of the spindle, and that in Albizzia lophanta, a Dicotyledon,
and Dioscorea quinquefolia, a Monocotyledon, quadripartition
[The Buttetin for February (49: 31-50) was issued April 4, 1922.]
2 i
52 FARR: QUADRIPARTITION IN SISYRINCHIUM
is accomplished by equatorial plates on the spindles. Quadri-
partition by cell-plates has never been satisfactorially described
and figured in any of the higher plants in connection with a
study employing modern methods of microtechnique; it is there-
fore much to be desired that the authors present these instances
in detail as soon as possible.
Florin (7) in 1918 found that quadripartition occurs in the
spore-formation of one of the acrogynous Jungermanniales but
does not give the details of the process as to whether it is by
furrowing or cell-plates. In 1919, Miss Digby (3) published
an extensive chromosomal study of Osmunda in which she refers
to the formation of cell-plates after the first reduction division.
These apparently persist through the second mitosis without
forming completed partitions, and then other cell-plates are
formed at right angles to these. The figures of these stages
are drawn from material of O. palustris var. aurea although the
description seems to apply to all four of the types studied,
including O. regalis. Smith (20) in 1900 described and figured
these stages of the latter species. He concluded that the cell-
plate of the first division was ephemeral and that the division
was finally accomplished by the simultaneous formation of six.
cell-plates on the six spindles respectively, thus accomplishing
quadripartition by cell-plates. During 1920 Yamaha (27) pub-
lished a study of cell-plate formation in the archesporial and
spore-mother-cells of Psilotum triquetrum. He found that the
nearly complete cell-plate which is formed after the first meiotic
division breaks up into a granular mass, which persists until
the close of the second nuclear division. At this time it becomes
transformed into connecting fibers upon which a cell-plate is laid
down between the non-sister nuclei, while other cell-plates are
being formed on the connecting fibers between the sister nuclei
of the second division. In this way a quadripartition of the
cell is accomplished by cell-plates. Yamaha concludes that the
cell-plate has a duplex nature from the first, and that it does not
split after formation as Timberlake believed. It is to be noted
that in quadripartition by furrowing the partition is duplex from
the first. Yamaha suggests that the invagination of the plasma-
membrane may play some part in the later stages in these cells
of Psilotum. It therefore seems to me that on account of this
duplex nature of the partition and in the light of the recent work
on quadriparition by furrowing that further work on cell-div-
ision of Psilotum would be warranted.
FARR: QUADRIPARTITION IN SISYRINCHIUM 53
Guérin (10) has made a study of the development of the
anthers and pollen of the labiates, but passes over the matter
of division of the mother-cell into the four microspores as ex-
hibiting nothing of particular interest. Valleau (26) in connec-
tion with his study of sterility in the strawberry has published
a careful account of pollen-formation. He describes in detail
the formation of the material about the protoplast which in
other forms has been considered as the thickening of the cell-
wall. However, in Fragaria, it seems to be a secretion from the
protoplast rather than the thickening of the wall. With regard
to the formation of the partitions the author does not commit
himself. He states:
he daughter nuclei are soon formed, and walls are laid down between
them, dividing the cytoplasm evenly. The cells gradually split apart,
separating the four microspores and allowing the entrance between them
of the viscous material.
It might be concluded from this description that cell-plates
accomplish the partitioning and that the invagination of the
peripheral material is simply incidental to the rounding. up
process. No figures of these stages are shown. Quite recently
R. R. Gates (9) has published a preliminary account of reduction
divisions in the pollen-mother-cells of Lactuca sativa. In this
paper there is described a quadripartition by furrowing rather
than by cell-plates, just as the writer (4) had previously reported
for Helianthus and Ambrosia, as well as in other groups of
Dicotyledons. No drawings are given by Gates, but a para-
graph is devoted to a description of the process. He mentions
that furrows may be formed either in the presence or the absence
of spindle fibers but does not describe the latter instance any
farther. It is stated that an ephemeral cell-plate occasionally
is found after the heterotypic division, but it never functions
and no cell-plate is present after the homoeotypic mitosis.
The most careful recent study of this question is that of
Mrs. Wanda K. Farr (6), in which she gives figures and des-
criptions of quadripartition in the pollen-mother-cells of Cobaea
scandens. These support the writer’s contention (4) that
cell-division by furrowing is common in the pollen-mother-cells
of Dicotyledons. It is interesting to note that the first descrip-
tion of cell-division in any plant is probably that by Brongniart
in 1827 on the pollen-mother-cells of Cobaea scandens (2), in
which it is indicated that the process is furrowing, though, of
54 FARR: QUADRIPARTITION IN SISYRINCHIUM
course, he had no realization of the internal mechanism of the
cell. This work was followed by that of Mirbel (17) who
presented his paper in 1832, though it was not published until
later. He described and figured quadripartition by furrowing
in the pollen-mother-cells of Cucurbita Pepo. A little later
Von Mohl (18) published figures and descriptions of the same —
process in other plants. It now seems that the interpretation
given by these first observers was more nearly correct than that
which has been given by most students of cell-division in higher
plants in more recent years.
Up to the present no detailed study has been made of quadri-
partition in Monocotyledons, and it was with the idea of com-
paring the situation in this group with that found in Dicotyledons
that the present study was undertaken. Several cases of
quadripartition in Monocotyledons are known. In 1915, L.
Guignard published two papers (12, 13) on the occurrence of
quadripartition in Monocotyledons. In the first paper he
includes a comprehensive review of the literature on that subject, —
as well as on bipartition in the reduction-divisions of Dicotyledons.
Quadripartition in Monocotyledons and bipartition in Dicoty-
ledons at the time of reduction-divisions are unquestionably
the exception in the method of pollen formation in these two
groups respectively. So rare is their occurrence in fact that
Van Tieghem (25) used them as characters of prime importance
in distinguishing the two classes of Angiosperms, thereby
including the Nympheaceae with the Monocotyledons. Guig-
nard had previously (11) reported quadripartition in six genera
of the Orchidaceae, and in the first (12) of his recent publications
adds four genera (six species) of Liliaceae and six genera (sixteen
species) of the Iridaceae. To this list the second paper (13)
contributes three more genera (three species) to the number
of the Iridaceae having this method. To the Liliaceae may be
added Strasburger’s (21, p. 151) findings in Asphodelus, and it will
be remembered that Hofmeister found it in Naias (14, p. 636).
In 1917, Tackholm and Séderberg published a paper (23) in
which they discuss tetrad formation in Monocotyledons. In
addition to its occurrence in the above-mentioned groups they
refer to quadripartition being reported by Hofmeister in Trades-
cantia, by Rosenberg in Anthericum and by Von Mohl and others
in the Juncaceae. So that they find it reported: in six families
of the Monocotyledons, namely: Liliaceae, Juncaceae, Iridacaea,
FARR: QUADRIPARTITION IN SISYRINCHIUM 55
Commelinaceae, Orchidaceae, and Cyperaceae. The last-named
is included because of the work of Juel on Carex, of which it
it may be questioned whether the term quadripartition is
applicable. In 1918 the same authors (24) published again,
adding another species of Monocotyledons to this list, namely,
Dioscorea quinquefolia. This brings the total number of genera
up to nineteen or twenty. In Canna they report a tetrahadral.
arrangement of the microspores but do not take this as evidence
of quadripartition.
Guignard (12) states that in four species of Iris there is
occasionally a suggestion of a partition after the heterotypic
nuclear division but this is always ephemeral. After the
homoeotypic mitosis in Sisyrinchium, Antholyza, Freesia, Ixia,
and Monbretia, when the fibers of the central spindle are formed
between the four nuclei, it is stated that rather frequently there
is noticeable a faint thickening on the internal face of the mother
wall at the places which are to become the points of insertion
of the partitions. Further than this no details of quadripartition
are given, nor are drawings or photomicrographs shown to give’
evidence as to whether the process is accomplished by cell-plates
or by constriction furrows. It is not unlikely that the ephemera!
equatorial differentiations which Guignard found in these
Iridaceae, and which Tackholm and Séderbeg reported for
Aristolochia Sipho are orange zones, such as the writer (5) has
shown to occur in Magnolia. Tackholm and Séderberg (24)
do not describe the process of quadripartition in Dvzoscorea
except to state that it is accomplished by equatorial plates
being formed on the spindles.
It thus appears desirable that a study be made of the details
of the process of quadripartition in a Monocotyledon. The
writer has discussed at some length the literature on this subject
in his first paper in 1916 (4). At that time he presented a
description and drawings of quadripartition in Nicotiana and
other Dicotyledons which led to the conclusion that no cell-plates
are formed, but that division is accomplished by furrows very
much as in animal cells. Mrs. Wanda K. Farr (6) has more
recently shown that the same type of cytokinesis occurs in
Cobaea scandens as was found in Nicotiana. In 1918, the writer
(5) supplemented the work of Guignard on Magnolia, in which
the latter showed incipient furrows during interkinesis, but did
not present the stages following the origin of the tetranuclear
56 FARR: QUADRIPARTITION IN SISYRINCHIUM
condition. It was found that after the disappearance of the
ephemeral orange zone a furrow developed on the margin of the
equator of the heterotypic spindle. However, the development
of this furrow is arrested when it is only about one-fourth
completed and the homoeotypic nuclear division ensues. The
division of the cell is finally accomplished by the completion
of this furrow and the formation of two other furrows at approxi-
mately right angles to it. Although Nymphaea has not as yet
been thoroughly investigated the work of Lubimenko and Maige
(16) indicates that it is similar in this respect to Magnolia.
The following study was made upon material of Sisyrinchium
Bushii Bicknell, collected near College Station, Texas, and
identified by Dr. A. S. Hitchcock. This species proved especi-
ally fine for study, both because of the excellent fixation and
because of the distribution of the mitotic figures in the anthers.
In Nicotiana the mother-cells of a given anther are all in very
nearly the same stage of division, and hence to secure an un-
broken series of stages it is necessary to take material from several
different anthers. In Magnolia the mother-cells within the
anther were at different stages but there was no special arrange-
ment of them. In either of these cases the objection might
be raised that a cell-plate was overlooked due to the study of
an incomplete series of stages. It was therefore highly desirable
that a form be found in which the stages were arranged in order
within the anther from one end to the other, so that by studying
a number of anthers it could be determined with certainty that no
steps have been omitted. Such a situation is that which Sisy-
rinchium Bushii presents. The mother-cells at one end of the
anther may be in metaphase of the heterotypic division and
those at the other end in metaphase of the homoeotypic. This
is the condition shown in Fic. 1.. In Fic. 2, the cells at the inner.
end are in metaphase of the homoeotypic division and those at
the outer end are in the tetranuclear stage. It is evident that
the study of a number of such anthers is sure to reveal every
stage, however transitory. It will be remembered that a similar
condition exists in the testes of Batracoceps, and among plants
it was found by Strasburger (22) in the anthers of Fritillaria and
by D’Angremond (1) in the banana.
e pollen-mother-cells of Sisyrinchium are smaller than
those characteristic of many Monocotyledons. Miss Kliene-
berger (15) has recently published a study of the size of nuclei
FARR: QUARDIPARTITION IN SISYRINCHIUM 57
of Monocotyledons, in which she finds that contrary to the
general opinion the nuclei of most Monocotyledons are small.
It is only in the Iridaceae and parts of the Liliaceae, Amary]-
lidaceae and Convallariaceae that large nuclei are found. How-
ever, her study is not concerned with pollen-mother-cells
especially. It would be interesting to find how great the
differences in size of pollen-mother-cells may be among Mono-
cotyledons, and their relation to the size of the nuclei. The
tetranucleate pollen-mother-cells of Sisyrinchium just before
division are 36-40 microns in diameter, which is just about the
size of those of Nicotiana.
The mother-cell wall is likewise thickened to about the same
extent as is that of Nicotiana, that is, on the average about one-
tenth of the diameter of the cell. The process of thickening
of the cell-wall begins about the time of synapsis, but the greatest
development is during interkinesis. The cells are more compact
within the anther than they are in either Nicotiana or Magnolia,
but nevertheless are quite loosely disposed during diakinesis
and later stages of reduction. They are somewhat closer
together toward the end of the process than in the earlier stages
doubtless due to the enlargement of the cells and the thickening
of the walls. There seems to be no evidence that the gelatinous
substance enveloping the protoplast during the later stages of
reduction-divisions is a secretion from the protoplast, as Valleau
(26) suggests, rather than a product of the cell-wall. The
middle lamellae remain quite distinct throughout these stages
and careful observation of the material both of Sisyrinchium
and of the other forms previously studied leads the writer to
the conclusion that it is the secondary lamellae of these pollen-
mother-cells which during reduction-divisions take on the
colloidal property of imbibition and swell to many times their
original thickness.
The heterotypic division results in two discoid nuclei being.
organized at opposite ends of a rather long central spindle. No
indication of a cell-plate or other equatorial differentiation is
to be discerned at any time during interkinesis. Nor is there
any indication of furrowing. The nuclei gradually become more
nearly spherical (Fic. 1) and finally, before the second nuclear
division begins, they are almost perfect spheres. They appar-
ently do not come into contact with the plasma membrane. The
fibers of the central spindle disappear almost entirely during in-
58 FARR: QUADRIPARTITION IN SISYRINCHIUM
terkinesis, and the integrity of the spindle as such becomes en-
tirely lost. The fact that all stages of interkinesis may be found
within the extent of a single anther (Fic. 1) makes it possible
to arrive at an index of the relative time required for that stage.
Judging from the extent of the various stages within the anther
it would seem that the time involved in interkinesis is about
equal to that involved in either the first or the second reduction-
division.
In the metaphase of the homoeotypic nuclear division the
spindles are in some cases parallel and in some at right angles
to each other, and in still others at positions intermediate
between these two extremes. When the nuclei are reconstituted
they may occasionally be found in a single plane as is shown in
the outer end of the anther in Fic. 2. But more frequently the
nuclei are tetrahedrally arranged (Fic. 3). Quite a number of
cells show intermediate orientations. It is interesting to note
that the type of division of the cell is the same, regardless of
the orientation. of the nuclei, just as was previously reported
for Magnolia (5).
he nuclei become reorganized in much the same manner
as after the heterotypic division. They, however, do not become
as large as in the previous division and they move apart so that
they become closely appressed to the plasma membrane. Al-
most as soon as the nuclear membranes appear spindles are
organized connecting the nuclei which are not already connected
by the two homoeotypic spindles. There are thus six spindles
in all within the cell, but the integrity of some of these seems
alirest lost especially in cases where the four nuclei are in one
plane.
The time elasping between the conclusion of nuclear division
and the beginning of cytokinesis is evidently relatively Jong.
This is indicated by the fact that all of the cells of a single anther
may be in these stages, those at one end just coming out of
karyokinesis and those at the other end not yet showing signs
of the formation of partitions. It thus appears that this period
is at least as long as that involved in either the heterotypic or
homoeotypic mitosis; which is quite in harmony with the writer's
previous suggestion (5) that cell-plate formation and furrowing
o not occur at corresponding periods in the cell-cycle, but that
cell-plate formation follows very soon after nuclear division, if
indeed the two processes may not frequently overlap. Furrow-
FARR: QUADRIPARTITION IN SISYRINCHIUM 59
ing on the other hand takes place only after a considerable time
has elasped following the re-formation of the nuclei. If cell-
plates appeared in this division we would most certainly expect
to find them in such an anther as that shown in FIG. 2, but none
such are present.
The process of cytokinesis in the quadripartition of the
pollen-mother-cells of Sisyrinchium is by furrowing and is
practically identical with that found in Nicotiana. It does not
seem necessary to repeat the details of the description of the
process here, inasmuch as there is no difference from that which
has been published before (4). In cells where the nuclei are
tetrahedrally arranged the plasma membrane invaginates most
rapidly at the points equidistant from the three nuclei and
therefore a central triangular area is formed (Fic. 3). In cells
in which the nuclei are in one plane no such central triangular
area appears, but the spindles have the same relation to each
other, as was shown in those cells of Magnolia (5) in which the
arrangement was of this type.
LITERATURE CITED
1. D’Angremond, A. Parthenokarpie und Samenbildung be
Bananen. Flora 107: 57-110. pl. 4-11. 5.
. Brongniart, Adolph. Mémoire sur la génération et le dé-
veloppement de l’embryon de la végétaux phanérogames.
Ann. Sci. Nat. 12: 14-53, 145-172, 225-296. fl. 34-44.
1827.
. Digby, L. On the archesporial and meiotic mitoses of
Osmunda. Ann. Bot. 33: 135-172. pl. 812. 1919.
Farr, C. H. Cytokinesis of the pollen-mother-cells of certain
’ Dicotyledons. Mem. New York Bot. Garden 6: 253-
217. Dl. 27-29. IGIG.
- — Cell division by furrowing in Magnolia. Am. Jour.
Bot. 5: 379-395. pl. 30-32.
. Farr, Wanda K. Cell division of the pollen-mother-cells of
Cobaea scandens alba. Bull. Torrey Club 47: 325-338.
pl. 14. 1920. n
Florin, R. Cytologische Bryophytenstudien I. Uber Spor-
enbildung bei Chiloscyphus polyanthus (L.) Corda.
Ark. f6r Bot. 15'*: 1-10. pl.r+/f.1,2. 1918.
. Frye, T. C., & Blodgett, E. B. Contribution to the life-
history ‘il Apocynum androsaemtfolium. Bot. Gaz. 40;
49-53. pl. 2. 1905.
Ny
w
-
on
ON
“I
co
60
FARR: QUADRIPARTITION IN SISYRINCHIUM
9. Gates, R. R. A preliminary account of the meiotic phe-
Io.
Lal
—
Lal
oo
—
‘o
No
°
ty
=~
nomena in pollen mother-cells and tapetum of lettuce
(Lactuca sativa). Proc. Roy. Soc. London, B. 91: 216-
oe fi) 258.6 HOA:
Guerin, Paul. Développement de 1l’anthére et du pollen
des Labiées. Compt. Rend. Acad. Sci. Paris 168:
182-185. I919.
. Guignard, L. Recherches sur le développement de !|’anthére
et du pollen des Orchidées. Ann. Sci. Nat. Bot. VI.
14: 26-45. Ol. 2.
Sur la formation du pollen. Compt. Rend. Acad.
Sci. Paris. 160: 428-433. I9I5.
Nouvelles observations sur la formation du _ pollen
chez certaines Monocotylédones. Compt. Rend. Acad.
Sci. Paris. 161: 623-625. 1915.
. Hofmeister, W. Neue Beitrage zur Kenntniss der Embryo-
bilding der Phanerogamen. II. Monocotyledonen. Ab-
handl. K6énigl. Sachs. Ges. Wiss. 7: 629-760. pl. 1-25.
1861,
. Klieneberger, Emmy. Ueber die Grésse und Beschaffenheit
der Zellkerne mit besonderer Beriicksichtigung der
Systematik. Beih. Bot. Centralb. 35': 219-278. pl. r.
1918.
. Lubimenko, W., & Maige, A. Recherches cytologiques sur
le développement des cellules-méres du pollen chez les
Nymphéacées. Rev. Gén. Bot. 19: 401-425, 433-458,
474-595. 1907.
. Mirbel, M. Complément des observations sur le Mar-
chantia polymorpha, suivi de recherches sur les méta-
morphoses des utricules, et sur l’origine, les développe-
ments et la structure de l’anthére et du pollen des
végétaux phanérogames. Mém. Acad. Roy. Sci. 13:
375-436. pl. I-10. 1835.
. Mohl, Hugo von. Ueber die Vermehrung der Pflanzen-
zellen durch Theilung. Tiibingen. 1835.
- Samuelson, G. Ueber die Pollenentwickelung von Anona
und Aristolochia und ihre systematische ores ws
Svensk. Bot. Tidsk. 8: 181-189. 1914.
. Smith, W.R. The achromatic spindle in the spore mother
om of Osmunda regalis. Bot. Gaz. 30: 361-377. pl. 22.
: Sirastuigex. E. Zellbildung und Zelltheilung. Ed. 3. Jena.
1880.
FARR: QUADRIPARTITION IN SISYRINCHIUM 61
‘ Das botanische Praktikum. Jena. 1884.
23. Tackholm G., & Soderberg, E. Ueber die Pollenent-
wickelung bei Cinnomomum nebst Eroérterungen iiber
die phylogenetische Bedeutung des Pollentypus. Ark.
for Bot. 158: 1-12. 1917.
24. —— Neue Beispiele der simultanen und_ sukzessiven
Wandbildung in den Pollen-mutterzellen. Svensk. Bot.
Tidskr. 12: 189-201. f. 1-9. 1918
25. Tieghem, Ph. Van. L’oeuf du jileeniteas considére ns werk
de la classification. Ann. Sci. Nat. Bot. VI.
464. I9O1
26. Valleau, W. D. Sterility in the strawberry. Jour. Agr.
Research 12: 613-6.70 pl. B-E+ 35, 36. 1918.
27. Yamaha, Gihei. Einige Beobachtungen iiber die Zellteilung
in der Archesporen und Sporenmutterzellen von Psilotum
triquetrum Sw., mit besonderer Riicksicht auf die Zell-
plattenbildung. Bot. Mag. Tokyo 34: 117-129. ff. I-20.
1920.
Description of plate 2
Fic. 1. Photomicrograph of a longitudinal section of an anther of
Sisyrinchium Bushii. The cells in the outer end are in the metaphase of
the oe and those at the inner end are in metaphase of the homoeo-
typic a
Fic — of an anther at a slightly later stage of develop-
ment. The cells at the inner end are in metaphase of the homoeotypic
mitosis and those at the outer end are in the tetranucleate stage just prior
to furrow formation.
3. A drawing of a cell in cytokinesis. The furrows are partly
formed and the central triangular area is appearing.
JNn
QQ
BuLL. TORREY CLUB VOLUME 49, PLATE 2
FARR: QUADRIPARTITION IN SISYRINCHIUM
A modern plant fossil
Epa M. Rounp
(WITH ONE TEXT FIGURE)
While collecting botanical materials in East Killingly,
Connecticut, the writer came upon a so-called “cedar swamp”
in which ferns like Woodwardia angustifolia Smith and Aspidium
simulatum Davenp. grew, together with sphagnum and several
species of liverwort. Woodsmen had visited the locality recently
and cut the large hemlock trees, from the stumps of which
broad shelf fungi belonging to the Polyporaceae had developed.
One of these fungi had been removed from its host by a previous
visitor and thrown upon a rock, over which flowed the waters
of the peat bog. Examination of this water-soaked specimen
showed it to be tough and pliable like India rubber, due perhaps
to the resinous nature of the hemlock tree on which the fungus
grew and the preservative qualities of the water from the peat
bog. The specimen was in such good condition that the writer
contemplated drying it for further study, when a cluster of the
same fungi was found in situ upon a large hemlock stump,
somewhat overgrown by small black birch trees. Examination
of one of these specimens revealed not only the ordinary lines
upon its surface but ‘also several very clear tracings of black
birch leaves, the details of which were as exact as obtain on
many rocks with fossil leaves impressed thereon (FIG. 1).
In searching for an explanation of this phenomenon, it
appeared that the black birch leaves had touched the surface
of the shelf fungus when it was damp and growing rapidly, so
that the leaves had been completely overgrown by the fungoid
strands while still attached to the parent plant. Over the whole
surface could be found examples of leaves in process of being ab-
sorbed or incorporated into the fungus, parts of them being
encrusted while other parts of the same leaves were free, dry
and easily broken. Still other cases showed leaves scarcely
touching the fungus but evidently somewhat invaded by its
hyphae, as they were beginning to brown and seemed to be dying.
Many ideas are advanced to account for the formation
63
64 RounD: A MODERN PLANT FossIL
of fossils.* Given, however, materials of as tough a nature
as the shelf fungus from the hemlock stump, on which a leaf
had been encrusted or replaced by hyphae of the same tough-
%
Fic. 1. Black birch leaves “fossilized” on a_ shelf fungus, X 3%. Original
specimen now in the Paleontological Laboratory, Brown University.
ness, one has but to postulate the submergence of the fungus
in the surrounding peat bog and conditions are supposedly right
for its continued preservation.+
It appears, therefore, that the fungus above described with
its encrusted black birch leaves illustrates a method whereby
fossils may have formed in the past or may develop under mod-
ern conditions. It may therefore be popularly designated as
exemplifying a ‘‘modern plant fossil.’
* Pirsson & Schuchert, Text book of geology, p. 435, 1915.
Tt See Shimer, An introduction to the study of fossils, p. 3, 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE
1910-1921
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
America being used in the broadest sense.
Reviews, and papers that relate exclusively to foresty, agriculture, horti-
ee con pepe of oe origin, or laboratory methods
i nd is mad index the literature of bacteriology.
An occasional sasied is anide t in favor of some paper appearing in an Ameri-
can 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 three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Adams, J. F. Rustson conifers in Pennsylvania. Pennsylvania
Agr. Exp. Sta. Bul. 160: 3-30. f. 1-10. D 1919
Adams, J. F. Sexual fusions and development of the sexual
organs in the peridermiums. Pennsylvania Agr. Exp. Sta.
Bull. 160: 31-76. pl. 1-5 + f. 1-8. D 1919.
Albertson, A. O. Nantucket wild flowers. Pp. i-xlv + 1-442.
i921. [Illust.]
Arny, A. C. Further experiments in field technic in plot tests.
Jour. Agr. Research 21: 483-500. pl. 94. 1 Jl 1921.
Ashe, W. W. A new shrubby ged Jour. Washington
Acad. Sci. 3: 434° 468.2
Aesculus microcarpa, sp. nov., a. poe
Barnes, C. R. Physiology, in Coulter, J. M., Barnes, C. R., &
Cowles, H. C., A textbook of botany 295-484. f. 619-600:
1910.
Barnhart, J. H. Otto Kuntze. Bull. Charleston Mus. 9: 65-
68: DD: 1913.
Berry, E,W. An Eocene flora from Trans-Pecos, Texas. U.S.
Geol. Surv. Professional Paper 125-A: 1-9. pl. 1-3+f.
2 S 1919
Includes new species in Geonomites (1) and Ilex (1).
65
66 INDEX TO AMERICAN BOTANICAL LITERATURE
Berry, E. W. The teaching of paleobotany. Bull. Geol. Soc.
Amer. 31: 389-394. 30S 1920.
Blake, S. F. The American species of Maximilianea (Cochlo-
spermum). Jour. Washington Acad. Sci. 11: 125-132. f. 2,
19 Mr. 1921.
Includes I new species from Venezuela.
Blake, S. F. The anay, a new edible-fruited relative of the
avocado. Jour. Washington Acad. Sci. 9: 457-462. 4 0
1919.
Ee: 2 new species of Hufelandia from Central America.
Blake, S. F. A preliminary revision of the North American
and West Indian avocados (Persea spp.) Jour. Washington
Acad. Sci. 10: 9-21. f. 7, 2. 4 Ja 1920.
Includes 2 new species.
Blake, S. F. Revision of Ichthyomethia, a genus of plants used
for poisoning fish. Jour. Washington Acad. Sci. 9: 241-
252. 4 My 1919.
Includes 2 new species from tropical America.
Blake, S. F. Revision of the true mahoganies (Swietenia).
Jour. Washington Acad. Sci. 10: 286-297. f. z, 2. 19 My
1920
Includes I new species from Mexico.
Briggs, L. J. The living plant as a physical system. Jour.
Washington Acad. Sci. 7: 89-111. f. I-6. 19 F 1917.
Briggs, L. J., & Shantz, H. L. The wilting coefficient for
different plants and its indirect determination. Jour.
Washington Acad. Sci. 1: 228-232. 19 N 1911
Britton, E.G. Bahama mosses. Bryologist 24: 17-19. pl. I.
25 Jl 1921.
Britton, E.G., & Holzinger, J.M. The rediscovery of Physcomi-
trium pygmaeum James. Bryologist 24: 26, 27. 25 Jl
1921.
Britton, N. L. Further botanical studies in Trinidad. Jour.
New York Bot. Gard. 22: 93-102. My 1921.
Brown, B. Cretaceous Eocene correlation in New Mexico,
Wyoming, Montana, Alberta. Bull. Geol. Soc. Amer. 25:°
355-380. 15 S 1914.
Buchholz J. T. Studies concerning the evolutionary status of
polycotyledony. Am. Jour. Bot. 6: 106-119. f. 1-25. Mr
1919.
INDEX TO AMERICAN BOTANICAL LITERATURE 67
Carey, C. L. On the gross structure of an agar gel. Bull.
Torrey Club 48: 173-182. f. 7-4. 1 Au 1921.
Clements, F.E. Scope and significance of eae ecology. Bull.
Geol. Soc. Amer. 29: 369-374. 30 Je 1918.
Cobb, F., & Bartlett, H. H. On sushiee inheritance in
crosses between mass- mutating and non-mass-mutating
strains of Oenothera pratincola. Jour. Washington Acad.
Sci. 9: 462-483. 4 O 1919.
Collins, G. N. Intolerance of maize to self-fertilization. Jour.
Washington Acad. Sci. 9: 309-312. 4 Je 1919.
Collins, G.N. New place effect in maize. Jour. Agr. Research
1Z:. 231-243. 4 FP 1gts:
Collins, G. N. The origin of maize. Jour. Washington Acad.
Sci. 2: 520-530. 19 D 1912.
Collins, G. N., & Kempton J. K. A hybrid between Tripsacum
and Euchlaena. Jour. Washington Acad. Sci. 4: 114-117.
4 Mr 1914.
Collins, G. N., & Kempton, J. H. A _ teosinte-maize hybrid.
Jour. Agr. Research 19; 1-37. pl. 1-7 +f. 1-33. 1 Ap 1920.
Cook, O. F. Agriculture and native vegetation in Peru. Jour.
Washington Acad. Sci. 6: 284-293. f. z, 2. 19 My 1916.
Cook, O. F. Evolution through normal diversity. Jour. Wash-
ington Acad. Sci. 9: 192-197. 4 Ap 1919.
Cook, O. F. Glaucothea, a new genus of palms from Lower
California. Jour. Washington Acad. Sci. 5: 236-241. 4
Ap 1915.
Cook, O. F. Ivory palmsin Panama. Jour. Washington Acad.
Sci. 3: 138-143. 4 Mr 1913.
Includes 4 new species of Phytelephas.
Cook, O. F. Jointed leaves of Amygdalaceae. Jour. Washing-
ton Acad. Sci. 2: 218-220. 4 My I912
Cook, O. F. The Mascarene cabbage palm as a new genus.
. Jour. Washington Acad. Sci. 7: 121-127. 4 Mr 1917.
Linoma Cook, gen. nov.
Cook, O. F. Morphology and evolution of leaves.. Jour.
Washington Acad. Sci. 6: 537-547- 19S 1916.
Cook, O. F. A new omer name for the epee Jour.
Washington Acad. Sci. 3: 158-160. 19 Mr
Achradelphia Cook., nom. nov.
68 INDEX TO AMERICAN BOTANICAL LITERAURE
Cook, O. F. A new genus of palms allied to Archontophoenix.
Jour. Washington Acad. Sci. 5: 116-122. 19 F 1915.
Loroma Cook, from Australia.
Cook, O. F. Pomegranate flowers dimorphic. Jour. Washing-
ton sAcad: Sci..2: 434-497. fot, 2. 4. N_1912.
Cook, O. F. Quichua names of sweet potatoes. Jour. Wash-
ington Acad. Sci. 6: 86-90. 19 F 1916.
Cook, O. F. Seedling morphology in palms and grasses. Jour.
Washington Acad. Sci. 7: 420-425. 19 Jl 1917.
Cook, O. F. Tribroma, anew genus of tropical trees related to
Theobroma. Jour. Washington Acad. Sci. 5: 287-289. 19
Ap I0I5.
Based on. Theobroma bicolor H. & B. of tropical America.
Cook, O. F., & Cook, R.C. The maho, or mahogua, as a trans-
Pacific plant. Jour. Washington Acad. Sci. 8: 153-170.
19 Mr 1018.
Cook, O. F.,& Cook, R.C. Polynesian names of sweet potatoes.
Jour. Washington Acad. Sci. 6: 339-347. 4 Je 1916.
Coulter, J. M. Morphology, in Coulter, J. M., Barnes, C. R.,
. & Cowles, H. C., A textbook of botany 1-294. f. 1-618.
1910.
Coulter, J. M., Barnes, C. R., & Cowles, H.C. A textbook of
botany (for colleges and universities). Pp. 1-964 + A-L
+ a-q. f. I-1234. 1910-1911.
Coville, F. V. The formation of leafmold. Jour. Washington
Acad. Sci. 3: 77-89. 4 F 1913.
Cowles, H. C. Ecology, in Coulter, J. M., Barnes, C. R. & ©
Cowles, H. C., A textbook of botany 485-964. f. 700-
1234. 1611.
Cunningham, C. C. Study of the relation of the length of
kernel to the yield of corn (Zea Mays indentata). Jour.
Agr. Research 21: 427-438. pl. 80-87. 1 Jl 1921.
Dufrenoy, J. The biological significance of false witches’-
brooms in ericaceous plants. Jour. Washington Acad. Sci.
8: 527-532. 19.5 1918:
Fairchild, D. A hunter of plants. Nat. Geog. Mag. 36: 57-77:
Jl 191g. [Illust.] ;
Account of the work of Frank N. Meyer.
INDEX TO AMERICAN BOTANICAL LITERATURE 69
Fernald, M. L. The American representatives of Avrenaria
sajanensis. Rhodora 21: 12-17. Io F Igig.
Includes Arenaria marcescens, sp. nov., from Newfoundland and Quebec.
Fernald, M. L. American variations of Avrenaria verna.
Rhodora 21: 21, 22. 10 F 191g.
Fernald, M. L. The earlier names for Alsinopsis. Rhodora
21: 9-12. 10 F 1919.
Fernald, M. L. The specific identity of Arenaria groenlandica
and A. glabra. Rhodora 21: 17-21. 10 F 1919.
Fernald, ae L. The type of the genus Alsine. Rhodora 21:
7-9. 10 F 1919.
oo i L. The unity of the genus Arenaria. Rhodora
1: 1-7. 10 F 1919.
ae many new combinations.
Goodspeed, T. H., McGee, J. M., & Hodgson, R. W. Note on
the effects of illuminating gas and its constituents in causing
abscission of flowers in Nicotiana and Citrus. Univ.
California Publ. Bot. 5: 439-450. 28 D 1918. [Corrected
reprint.]
Griffiths, D., & Juenemann, H. E. Commercial Dutch-bulb
culture in the United States. U.S. Dept. Agr. Bull. 797:
I-50. f; 2-32: 22: N--ig1.-
Harper, R. M. The supposed southern limit of the eastern
hemlock. Torreya 19: 198, 199. O IgI9.
Harper, R. M. Water and mineral content of an epiphytic
fern. Am. Fern Jour. 9: 99-103. 24 Ja 1920.
Polypodium polypodioides (L.) Hitchcock.
Herre, A. W. C. T. Supplement to the lichen flora of the
Santa Cruz Peninsula, California. Jour. Washington Acad.
Sci. 2: 380-386. 19 S 1912.
Includes Thelocarpon albomarginatum, sp. nov.
Hitchcock, A. S. History of the Mexican grass, Ixophorus
unisetus. Jour. Washington Acad. Sci. 9: 546-551. 4 N
1919.
Hodges, R. S. Ringworm of the nails. Arch. Dermat. &
Syphil. 4: 1-26: f. 1-12. “Jl 1921.
Caused by species of Trichophyton.
Hollick, A. Quercus heterophylla in the Clove Valley [New
York]. Proc. Staten Id. Assoc. Arts & Sci. 7: 32-34. pl.
4. Ag18.
70 INDEX TO AMERICAN BoTANICAL LITERATURE
Hopkins, A.D. The bioclimatic law. Jour. Washington Acad.
Sci. 10: 34-40 19 Ja 1920.
Howe, M. A. A little-known mangrove of Panama. Jour.
New York Bot. Gard. 12: 61-72. f. 16-23. Ap I9gI11.
Jamieson, C. O., & Wollenweber, H. W. An external dry rot
of potato tubers caused by Fusarium trichothecioides,
Wollenb. Jour. Washington Acad. Sci. 2: 146-152. f. I.
19 Mr 1912.
Jennings, O. E. New or noteworthy plants from northwestern
Ontario. I. Jour. Washington Acad. Sci. 10: 453-460. 4
O 1920.
ncludes new species and varieties in Lysias (1), Kneiffia (1), Pyrola
(3), Scutellaria (1), Stachys (3).
Kearney, T. H. Plant life on saline soils. Jour. Washington
| Acad. Sci. 8: 109-125. 4 Mr 1918.
Kellerman, K. F. The effect of salts of boron upon the distri-
bution of desert vegetation. Jour. Washington Acad. Sci.
10: 481-486. 19 O 1920.
Kellerman, M. Phototypes, a means for wide distribution of |
type material. Jour. Washington Acad. Sci. 2: 346-348.
19 Au I9gI2.
Kempton, J. H. The ancestry of maize. Jour. Washington
Acad. Sci. 9: 3-11.. 4 Ja 19109.
Kempton, J. H. A correlation between endosperm color and
albinism in maize. Jour. Washington Acad. Sci. 7: 146-
149. 19 Mr 1917.
Kempton, J. H. Linkage between brachytic culms and peri-
carp and cob color in maize. Jour. Washington Acad. Sci.
11: 13-20. 4 Ja 1921.
Knowlton, F. H. A catalogue of the Mesozoic and Cenozoic
plants of North America. U.S. Geol. Surv. Bull. 696: I-
815. 1919.
Knowlton, F. H. Cretaceous-Tertiary boundary in the Rocky
Mountain region. Bull. Geol. Soc. Amer. 25: 325-340.
15 S 1914.
Knowlton, F. H. Description of a supposed new fossil species
of maize from Peru. Jour. Washington Acad. Sci. 9: 134-
136. 4 Mr Io1g.
INDEX TO AMERICAN BOTANICAL LITERATURE 71
Knowlton, F. H. Relations between the Mesozoic floras of
North and South America. Bull. Geol. Soc. Amer. 29:
607-614. 30 D 10918.
Lamb, W. H. Moreh oak, a new name for Quercus morehus
Kellogg. Jour. Washington Acad. Sci. 6: 657, 658. 19
IgI6.
Lewton, F. L. The Australian fugosias. Jour. Washington
' Acad. Sci. 5: 303-309. 4 My 1915
Lewton, F. L. The history of kidney cotton. Jour. Wash-
ington Acad. Sci. 10: 591-597. f. 1, 2. 19 D 192
[Lumsden, D.] Hand-list of trees and shrubs native and exotic,
including the Coniferae, grown in the arboretum [of Walter
Reed General Hospital, Washington, D. C.]. pp. 1-66.
1 Mr 1921.
Maxon, W.R. Anew Anemia from Mexico. Jour. Washington
Acad. Sci. 8: 199, 200. 4 Ap 1918.
Anemia Makriniti Maxon.
McCulloch, H. L. A_ bacterial disease of Gladiolus. Science
iI. 54: 115, 116... 5 Au 192%:
Bacterium marginatum, sp. nov.
McDougall, W. B. The classification of symbiotic phenomena.
Plant World 21: 250-256. O 1918.
McLaren, D. California’s flora. Parks & Recreation 4: 275-
279... fl -i92k. Tiiastj
McMurran, S. M. Walnut blight in the eastern United States.
U. S. Dept. Agr. Bull. 611: 1-7. pl. z, 2. to D 1917.
Metcalf, F. P. Notes on North Dakota plants. Jour. Wash-
ington Acad. Sci. 10: 188-198. 4 Ap 1920.
Moxley, G. L. Some vacation lichens. Bryologist 24: 24, 25.
25 Jl 1921.
Nichols, G. E. The vegetation of Connecticut. VI.
plant associations of eroding areas along the seacoast.
Bull. Torrey Club 47: 89-117. f. 1-6. 31 Mr 1920.
Norton, J. B. The eastern and the western migrations of
Smilax into North America. Jour. Washington Acad.
Sci. 6: 281-283. f. zr. 19 My I916.
Norton, J. B. Spring flowers in the fall. Jour. Washington
Acad Sci. 5: 285-287. f.z. 19 Ap 1915.
72 INDEX TO AMERICAN BOTANICAL LITERATURE
Parish, S. B. A supplementary bibliography of the Southern
California flora. Bull. S. California Acad. Sci. 19: 24-29.
a 1920.
Pearson, W. H. Notes on a small collection of hepatics from
Oregon. Bryologist 24: 21. 25 Jl Ig2t.
Piper, C. V. A new genus of Leguminosae. Jour. Washington
Acad. Sci. 10: 432,433. 19 S 1920.
Monoplegma, gen. nov., from Costa Rica.
Pittier, H. Inophloeum, a new genus of the mulberry family.
Jour. Washington Acad. Sci. 6: 112-114. 4 Mr 1916.
From Panama and adjacent Colombia.
Pittier, H. Notes on the genus Swartzia in Panama and
Guatemala. Jour. Washington Acad. Sci. 11: 155-160.
4 Ap 1921
Includes 2 new species.
Pittier, H. On the origin of chicle with descriptions of two
new species of Achras. Jour. Washington Acad. Sci. 9:
431-438. 19 S 1919.
New species from Guatemala and Panama.
Pittier, H. Two new species of Bursera. Jour. Washington
Acad. Sci. 11: 229, 230. 19 My 1921.
From Panama and Guatemala.
Pittier, H. The Venezuelan mahogany, a hitherto undescribed
species of the genus Swietenia. Jour. Washington Acad.
sci. 10: 32-34. 19 Ja 1020,
Swittenia Candollet, sp. nov.
Pritchard, F. J. Relation of horse nettle (Solanum carolinense)
to leafspot of tomato (Septoria may aoe Jour. Agr.
Research 21: 501-506. pl. 95-99. 1 Jl 19
Ramaley, F. The role of sedges in some a plant
communities. Am. Jour. Bot. 6: 120-130. fia, 2. Mr 191s.
Richards, B. L. Pathogenicity of Corticium vagum on the
potato as affected by soil temperature. Jour. Agr. Research
21: 459-482: pl. 88-93. 1 Jl 1921.
Ricker, P. L. A sketch of botanical activity in the District
of Columbia and vicinity. Jour. Washington Acad. Sci.
8: 487-498. 19 Au 1918; 516-521. 19 S. 1018.
Ricker, P. L. A synopsis of the Chinese and Formosan species
of Albizzia. Jour. Washington Acad. Sci. 8: 242-246. 19
Ap 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 7
Rose, J.N. A new Echeveria from Mexico. Jour. Washington
Acad. Sei. 1: 267-269. f; 1 4:1) 1911,
Echeveria lutea Rose.
Rydberg, P. A. Notes on Rosaceae—XIII. Bull. Torrey Club
48: 159-172... 1 Au. 1921.
Safford, W. E. Acacia cornigera and its allies. Jour. Washing-
ton Acad. Sci. 4: 356-368. 19 Jl 1914
Includes 10 new species from Mexico and ‘see America.
Safford, W. E. Chenopodium Nuttalliae, a food plant of the
Aztecs. Jour. Washington Acad. Sci. 8: 521-527. f. I-3.
19 S 1918.
A new species from Mexico.
Safford, W. E. Identity of cohoba, the narcotic snuff of ancient
Haiti. Jour. Washington Acad. Sci. 6: 547-562. f. I-3.
19 S 1916.
Safford, W. E. New or imperfectly known species of bull-horn
acacias. Jour. Washington Acad. Sci. 5: 355-360. 19 My
IQI5.
Includes 3 new species of Acacia from Mexico.
Safford, W. E. Notes on the genus Dahlia with descriptions
of two new species from Guatemala. Jour. Washington
Acad. Sci. 9: 364-373. f. 1-4. 19 Jl 1919.
Safford, W. E. VPapualthia mariannae, a new species of
Annonaceae from the island of Guam. Jour. Washington
Acad. Sci. 2: 459-463. f. 1, 2. 19 N Ig12.
Safford, W. E. Proposed classification of the genus Rollinia
with descriptions of several new species. Jour: Washington
Acad Sci. 6: 370-384. f. 1-3. 19 Je 1916.
Includes 3 new species from tropical America.
Safford, W. E. Rolliniopsis, a new genus of Annonaceae from
Brazil. Jour. Washington Acad. Sci. 6: 197-204. f. I, 2
19 Ap 1916.
Includes 2 new species.
Safford, W. E. Synopsis of the genus Datura. Jour. Washing-
ton Acad. Sci. 11: 173-189. f. 7-3. 19 Ap 1921.
Includes 1 new species from Colombia and 4 from Ecuador.
Shreve, F. A map of the vegetation of the United States.
Geog. Rev. 3: 119-125. F 1917 [with map.]
Solis,O. Importancia de los jardines y algunas especies vegetales
exoticas. Rev. Agricola 6: 78-84. Je. 1921 [Illust.]
74 INDEX TO AMERICAN BoraNicAL LITERATURE
Standley, P. C. Ammocodon, a new genus of Allioniaceae, from
the southwestern United States. Jour. Washington Acad.
Sci. 6: 629-631. 4 N 1916.
Standley, P. C. Blepharidium, a new genus a Rubiaceae from
Guatemala. Jour. Washington Acad. Sci. 8: 58-60. 4 F
1918.
Standley, P.C. Comparative notes on the floras of New Mexico
and Argentina. Jour. Washington Acad. Sci. 6: 236-244.
4 My 1916.
Standley, P. C. A new species of Rondeletia from Mexico.
Jour. Washington Acad. Sci. 8: 126, 127. 4 Mr 1918.
Rondeletia Rekoi Standley.
Standley, P. C. The North American species of Agonandra.
Jour. Washington Acad. Sci. 10: 505-508. 4 N 1920.
Includes 2 new species from Mexico.
Standley, P. C. The North American species of Genipa. Jour.
' Washington Acad. Sci. 8: 639-643. 4 D 1918.
Includes 2 new species from Panama.
Standley, P. C. Notes on Orthopterygium huaucui. Jour.
Washington Acad. Sci. 5: 628-631. f r. 19 N 1915.
Standley, P. C. Omiliemia, a new genus of Rubiaceae from
Mexico. Jour. Washington Acad. Sci. 8: 426, 427. 19 Jl
1918. ;
Standley, P. C. T briana. a new generic name. Jour.
Washington Acad. Sci. 6: 69, 70. 4 F 1916.
New name for Cladothrix S. Wats., 1880, preoccupied by Cladothrix
Cohn, 1875.
Stevens, F. L. Bacteriology in plant pathology. Trans. Am.
-Microscop. Soc. 36: 5-12. Ja 1917.
Swanton, J. R. Note on the aboriginal name ‘‘aje’’. —
Washington Acad. Sci. 6: 136, 137. 19 Mr 1916.
Swingle, W. T. Chaetospermum, a new genus of hard-shelled
citrous fruits. Jour. Washington Acad. Sci. 3: 99-102.
19 F 1913
From the Philippine Islands.
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386. S 1012;
Includes new American species in Rickia (1) and Trenomyces (4)-
Vol. 49 No. 4
BULLETIN
OF THE
TORREY BOTANICAL CLUB
APRIL, 1922
The phylogeny of the genus Brachiomonas
Tracy E. HAZEN
(WITH PLATES 3 AND 4 AND FIVE TEXT FIGURES)
Brachiomonas, one of the oddest and most interesting genera
of the Chlamydomonas group, has not until now been reported
outside of western Europe. The genus was established by
Bohlin (1) in 1898 with two species, differing in details of form,
found in Sweden near Stockholm. The more slender species,
B. gracilis, has not been reported elsewhere. The type species,
B. submarina, had been collected about three years earlier by
Lagerheim (6, p. 7) near Troms¢, Norway, and recorded as a
nomen nudum which was taken up by Bohlin. A somewhat
general distribution of this species on the Norwegian coast has
been indicated by Wille (15) and Printz (8, p. 21). It appears
that Dangeard (3, p. 74) had observed the same form about
ten years earlier than Bohlin, at Luc-sur-Mer, France, but had
postponed publishing it until gametes could be found. Chodat
(2, p. 143, f. 66) has recorded the species from Ajaccio, Corsica,
showing that it is not exclusively of northern habitat.
West (12) in 1908 reported a form collected in brackish
marshes at Sheerness, England, near the mouth of the Thames,
which he rather hesitatingly referred to B. submarina, since it
seemed to be somewhat intermediate between Bohlin’s two
species. In July, 1920, at Cullercoats, near Newcastle-upon-
Tyne, and in August at Plymouth, England, I found abundant
material in small rock pools, showing forms similar to those des-
cribed by West, together with specimens agreeing more with
Bohlin’s type. Earlier in the same season (June 25) I had
found on the coast of Norway at Valdersund, near Trondhjem,
[The BuLtetin for March (49: 51-74. pl. 2) was issued April 12, 1922.]
75
76 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
a few specimens which showed precise gareement with Bohlin’s
description of B. submarina. Again a few days later, at Aale-
sund, south of Trondhjem, I collected typical material, and here
in great abundance. At this station some pools were almost
exclusively filled with slender forms corresponding to Bohlin’s
B. gracilis; in other pools a mixture of the two species, with
perplexing intermediate forms, appeared. Observing pools con-
taining a natural pure culture, one would scarcely question
the distinctness of the two species: when both were found
together in the same pool, the variety of forms seen presented a
very strong suggestion of hybridization between the two species.
New material collected in New York in March and April,
1921, has furnished forms (Fics. 5-7) almost precisely like the
slender cells illustrated by West, together with others showing
every gradation to the broadest type of B. submarina. These
slender forms closely resemble Bohlin’s f. rb of B. gracilis,
but I am now convinced that West was entirely justified in
including them in the very variable broader species. We seem
not to have in America, and I did not find in England the ex-
treme forms represented by Bohlin’s f. za and rc of B. gracilis.
Whether they are sufficiently distinct to warrant the main-
tenance of the second species is a question which would be more
convincingly settled through studies by pure culture methods.
I first found Brachiomonas submarina in March, 1907, in
a small rock pool on the shore of Long Island Sound at Twin
Island, Pelham Bay, New York. It was found in almost pure
culture, coloring the water green from its abundance, and per-
sisted in the motile condition for some weeks when brought to
the laboratory. Although the same station was visited many
times during the following twelve years, no Brachiomonas was
found until-the middle of November, 1919, when it was again
abundant in the same pool, and occurred sparingly in neighbor-
ing rock hollows of similar character. I found it repeatedly
in the same pools until the first of December, in spite of hard
frosts and ice formation during these two weeks, The habitat
appears to be very like that of Bohlin’s type station, small
hollows so high above ordinary tide limits as to seem to be filled
only with rain water, which are nevertheless sometimes dashed
by waves so as to become brackish, as evidenced by taste and
by the salt incrustation left on the rock margin upon evaporation.
About the time of the collection in 1919, an unusually high tide
was reported, and it is possible that this may have washed the
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 77
Brachiomonas into the pool from the open sea. Such an occur-
rence might account for the sporadic appearance of the species
at this station, though it is much more probable that it is usually
present in a resting state and that conditions were not favorable
for the production of zoospores at the time of other visits to the
station. A strange point is that, although careful examination
of the specimens collected in the spring of 1907 shows them all
to be like Bohlin’s type, as were also the few specimens found
there in September, 1920, and abundant material from March to
June and in September, 1921, only a few such individuals were
found in the fall of 1919 (Fics. 1, 2), while most of the material
showed a distinct difference in form (Fics. 8-16). This can
hardly be interpreted as a mere seasonal variation. New mater-
ial collected the last of February of the present season, from
pools still coated with ice, consisted very largely of the new
obtuse form, but after a week or more in the laboratory this
form had mostly been replaced by individuals corresponding to
Bohlin’s type, present only in small numbers at the time of
collection. Whether this is a case of the new type actually
giving rise to the standard type through reproduction, or merely
an example of the regular succession of growth of different organ-
isms familiar in collections of protozoa, is a matter calling for
more extended observation. For the determination of the per-
manent status of the new form, pure culture methods are highly
desirable, but thus far I have been unable to grow Brachiomonas
on seawater agar, which succeeds admirably with other chlamy-
domonads of similar habitat. For the present it will be con-
venient to discuss this divergent plant under the name
BRACHIOMONAS SUBMARINA Bohlin, forma obtusa f. nov.
This new form is entirely like Bohlin’s type in its genera
shape; that is, it possesses a long posterior extension or horn
and four shorter median arms directed backwards, often with
depressions between them; the anterior portion might be roughly
likened to the shape of an umbrella with four ribs. There is
often a wedge-shaped beak through which the cilia emerge,
though sometimes the apex appears merely rounded. The
protoplast may fill the arms and posterior horn entirely (Fic. 8)
or partially (Fic. 12) or not at all. The massive chromatophore
may be equally variable; that is the protoplasm in the horn
and in the arms may be all green, as also in the typical form
(Fic. 2), or it may be colorless, sometimes appearing sharply
78 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
delimited from the central green mass (FIG. 8), while in other
cases the green and colorless regions merge into one another
almost imperceptibly (Fic. 1). The chromatophore is here
certainly less definitely or constantly organized than in most
species of Chlamydomonas; in fact it would perhaps be more
accurate not to speak of a definite chromatophore but merely of
cytoplasm with chlorophyll more or less diffused through it.
This variability in the extent of the distribution of the chlorophyll
in the protoplasm is very similar to the condition in Hydro-
dictyon, to which Harper (5, p. 179) has called attention. West
(12) states that in his specimens of B. submarina the chroma-
tophore never extended into the arms. This was undoubtedly
due to the fact that he never saw his material when freshly
collected, but only after it had been sent from near the mouth
of the Thames to Birmingham. I find that very generally in
freshly collected specimens the chromatophore fills the arms
and posterior horn, but often retracts from them soon after
being subjected to the less favorable light and aeration conditions
of captivity.
Imbedded in the chromatophore in a subparietal position,
regularly on the side next to the stigma, is a single relatively
large pyrenoid; in vigorous young specimens this is generally
situated in front of the middle of the cell; often in older individ-
uals it becomes posterior in position. The conspicuous red
eye-spot is linear in form; or, sometimes at least, it may be very
narrowly wedge-shaped, pointed in front and broader behind;
it lies in the depression midway between two arms. Most
frequently when the cell is at rest the pyrenoid and stigma lie
in a median ventral position (F1G. 6). On the other side of the
cell from the pyrenoid the nucleus may sometimes be detected
without staining, lying in a colorless mass of cytoplasm, best
seen in polar view (Fic. 13). When the pyrenoid lies in its ©
characteristic anterior position the nucleus is likely to be some-
what posterior (Fics. 15, 17); if the pyrenoid becomes posterior
the nucleus is then more often median in position. In older
stages vacuoles often develop in the chromatophore (FIG. 4)
as described by West (12)
In hanging drop cultures, or in jars kept for some time in
the laboratory, many individuals may be found which show little
or no trace of the four lateral arms (Fics. 14, 15). Bohlin has
shown such a form in one dividing individual. The cell wall is
of so firm a character that it is difficult to imagine how the arms
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 79
could be modified or suppressed after they are once formed;
nevertheless in practically all examples of division seen, I have
found the arms well developed before the escape of the daughter
cells (FIGs. 20, 21) so that such cells can hardly be looked upon
as a reversion to the simpler ancestral form.
. The main difference between this new form and Bohlin’s
type lies in the fact that the arms are here always rounded and
obtuse, while in the type form they are sharp pointed and often
more eae The characters of the form may be summarized
as foll
BSACHibEnNad SUBMARINA Bohlin, forma obtusa f
et chromatophoro alias i in extrema ipsa extendentibus, alias ex
eis plus minusve retractis.
Longit. cellularum 15-32 (saepius 20-25,); lat. cell, 15-
22u (saepius ca. 18,).
Hab. in aquis subsalsis lacunarum saxearum. Twin Island,
Pelham Bay, New York; Nov., 1919, Feb.—March, 1922.
Chodat’s figures (2, f. 66) indicate that the form collected
in Corsica may be identical with the one here described, though
it is possible that the difference between his sketches and those
of Bohlin may be due to the personal equation in drawing.
As indicated above, I have found this obtuse form only at
one station. Bohlin’s type, as represented by my FIc. I-55
have also collected early in September during each of the past
two years at Bass Point, Nahant, Massachusetts. It should
be looked for on the coast of Maine and New Brunswick as
well, at any points where rocky ledges occur.
In older cultures numerous cells were found which showed
the entire protoplast contracted into a globose mass (FIG. 16).
These showed no further development in this material; but in
fresh hanging drop cultures of B. submarina followed for a much
shorter time in Norway, such cells presently became quiescent
and developed a new spherical wall closely investing the pro-
toplast. Soon these aplanospores showed their chromatophore
turning ‘from green to a tawny tint, and finally the original
wall of the zoospores gradually disappeared.
Practically all genera allied to Chlamydomonas possess two
contractile vacuoles pulsating more or less in alternation,
situated near the base of the cilia. In Brachiomonas these
appear to be entirely absent. One of West’s figures (12, pl. 20,
f. 1; 14, f. ror c), it is true, suggested such vacuoles, but his
80 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
beautiful original sketches in color, preserved with his collection
of drawings in the South Kensington Museum, show that these
spots are brown colored granules suspended in the fluid of a
large permanent vacuole. The absence of contractile vacuoles
I have been inclined to regard as another mark of the highly
specialized character of the genus, since they are regularly to
be expected in primitive members of this group and its ancestors.
It is possible, however, that this peculiarity is merely an incident
due to the special habitat; for Biitschli has remarked the general
absence of contractile vacuoles in marine Ciliates and Rhizopods,
and D’Arcy Thompson (11, pp. 165 and 264) regards this
absence in such cases as ‘‘no more than a physical consequence
of the different conditions of existence in fresh water and in
salt.”” The marine species, Chlamydomonas brachyura West,
besides exhibiting other points of resemblance to Brachiomonas,
appears also to have no contractile vacuoles; this species was
discovered in association with a new Carteriaceous genus
Platymonas, in which West (13) found two small contractile
vacuoles, which, however, ‘‘could only be observed with dif-
ficulty.”” In two brackish water species of Platymonas which
I have investigated with considerable care, I am unable to
satisfy myself that I ever see contractile vacuoles. Lewis has
just published figures of one of these species, P. subcordiformis
(Wille) Hazen,* which likewise show no contractile vacuoles.
Furthermore, two new brackish water forms of the primitive
family Polyblepharidaceae, one a minute species of Pyramimonas
and the other a new genus most closely related to Polyblepharides,
which have turned up in my latest collections of Brachiomonas,
made while this paper was in the hands of the printer, very
certainly possess no contractile vacuoles. Over against these
several records of their absence, however, attention may be
called to their presence (though here very small) in Chlamy-
domonas caudata Wille, discussed below, which must be con-
sidered as regularly a marine organism.
Asexual reproduction is accomplished by internal division
into four or eight (very rarely perhaps two) daughter cells.
The division very generally begins while the cell is swimming
actively, and often the typical cell shape is acquired by the
daughter cells or zoospores and they move with their own cilia
inside the mother cell-wall while it continues in active movement
* Notes from the Woods Hole Laboratory,—1921. Platymonas subcordi-
formis (Wille) Hazen, Rhodora 23: 249-251. pl. 133, f. 1-19. Mr 1922
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 8I
by its own cilia. Bohlin states that the first division is longi-
tudinal, and certainly this is regularly the case. Chodat
presents a figure (2, f. 66), and I have seen two or three such
cases, where the first division appears to be transverse, but it
is quite possible that such an appearance is due to an unusual
shifting of the position of the cell contents after a proper longi-
tudinal division. The subsequent divisions have not been
described or figured by previous observers, and are ordinarily
difficult to follow because of the continual active movement
of the mother cell. I was so fortunate as to find one very
favorable case in B. submarina f. obtusa, and later many similar
ones in the type form, which I believe to be entirely representa-
tive. This individual became quiescent after the first division
was completed (FIGs. 22, 23) ;it presented its anterior pole upward,
so that it was possible to see easily that the second plane of
cleavage is also longitudinal and perpendicular to the first.
Immediately after the second cleavage a slight stretching and
shifting of the daughter portions took place, so that one of them
appeared somewhat underneath two others (Fic. 24). The
third cleavage appears to cut each of the four daughter portions
across its longer diameter, that is the plane seems to be essentially
transverse to the previous divisions and to the axis of the mother
cell. The pyrenoid disappears before the first cleavage occurs,
and the stigma fades out during this process. A new and
rather small pyrenoid appears in each daughter portion after
the second cleavage, and probably another pyrenoid arises de
novo in four of the eight daughter cells. About an hour after
the completion of the third cleavage the eight daughter cells
had acquired the typical form of the arms, and also the eye-
spot and cilia (Fic. 27); after about another hour they escaped
from the mother cell-wall. In certain cases it is clear that the
cilia of the mother cell remain connected by a strand of proto-
plasm to one of the daughter portions during division (FIG. 19)
or even until the eight zoospores acquire their final form.
Whether this is regularly the case appears doubtful. In mate-
rial identified as Brachiomonas gracilis Bohlin, at Aalesund,
Norway, I observed numerous cases of division which followed
the course outlined above.
In the case of the production of only four daughter cells
it is clear that the cytoplasmic polarity of the mother cell
would be directly transferred to each of the new zoospores.
But when division proceeds further it is less clear how the
82 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
polarity may be transmitted: for, as stated above, the third
plane appears to cut the four daughter portions transversely
in such a manner that half of the resulting eight would retain
the original anterior cytoplasm, while the four posterior zoospores
would seem to be under the necessity of regenerating (so to
speak) their anterior pole. It is possible that in the slight
shifting of the daughter portions following so quickly upon the
first two divisions there is actually more rotation than is apparent
so that the third division may effect a properly qualitative
bipartition, but that appears doubtful from the cases observed.
The whole problem of polarity in chlamydomonads is one of
great interest demanding a degree of care in observation which
hardly any species up to the present has received.
Brachiomonas simplex sp. nov.
This species was first discovered at Aalesund, Norway, in
the last days of June 1920, and was also collected three weeks
later at Cullercoats, near Newcastle-upon-Tyne, and in August
at Plymouth, England. It was found in small rock pools of
brackish water, sometimes in association with one of the other
species of Brachiomonas, then again in practically a natural
pure culture.
At first sight the organism looked like a Chlamydomonas,
many individuals (Fics. 31, 39) showing somewhat the general
oval, posteriorly pointed form characteristic of C. caudata
Wille (Fics. 46-50), which occurred near by, though never in the
same pools. But presently it was noticed that even the cells
most nearly approaching C. caudata in form were nevertheless
more like Brachiomonas submarina in internal organization,
showing a similar linear stigma and large lateral and anterior
pyrenoid; furthermore many individuals show small bumps or
protuberances slightly back of the middle region, immediately
suggesting the arms of Brachiomonas, though never really at-
taining that character (Fics. 28, 29). The posterior horn, often
rather elongated, shows a characteristic curve not seen in B.
submarina. Generally the protoplast was conspicuously fur-
nished with vacuoles, but whether this is a constant character
is somewhat doubtful, inasmuch as C. caudata in similar pools
was often similarly vacuolate at this time. The English ma-
terial, moreover, was much less vacuolate. There is great
variability in the development of the lateral protuberances; if
formed at all they appear before the escape of the zoospores
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 83
from the mother cell, and the form then attained persists
through the life of the individual. In newly formed zoospores,
usually, but not always, the protoplast and chromatophore ex-
tend into the posterior horn so as to fill it completely. In
older individuals the protoplast gradually retracts so that a
considerable portion of the horn, or finally all of it is empty.
The pyrenoid and red eye-spot regularly lie on the same side of
the cell, and the nucleus on the opposite side; as in B. sub-
marina, this is most clearly shown in cells momentarily resting
with one of the poles directed upward (Fic. 32).
As in the other species, asexual reproduction is accomplished
by division of the protoplast into four or eight daughter cells
which develop the characteristic form while the mother cell
still retains its cilia in active movement (FIGs. 35, 36).
In sexual reproduction, gametes may be formed to the
number of thirty-two in one mother cell, which retains its
motile condition until they are completely developed (Fic. 40)
just as in the case of asexual reproduction.. The gametes are
similar in form to the asexual cells, but much smaller, lacking
a cell wall, and furnished with relatively longer cilia: they
also seemed to be more apt to show angularity in the middle
region than the asexual cells. Usually there was a slight differ-
entiation between the pair, a smaller and more fusiform gamete
conjugating with one slightly larger and more angular (Fics.
41, 42). Meeting by the ciliated anterior ends, the fusion is
lateral, resulting in a quadriciliate zygote of ovoid-pyriform
shape which swims actively, and may for some time show the
two nuclei, two pyrenoids, and two linear eye-spots (FIG. 43);
when momentarily at rest the four long cilia trail backwards.
Finally the nuclei fuse, the cilia disappear (Fic. 44) the form
becomes spherical, and the thick wall of the resting zygospore
is developed (Fic. 45). After some days the cell contents
assume a reddish orange color similar to that of the aplanospore
of this species and of B. submarina
Brachiomonas simplex sp. nov. 8B. cellulis a acca
majoribus, Ree oviformibus quidem sed a
partem in curvatum productis, brachiis quae c ore
hujus Ge apetiet habent aut omnino a nae a evolutis
tantummodo in 2-4 gibbos undatos;
ad polum autem anteriorem globosis vel saalba cobich pantals
exornatis: chromatophoro cellulam prope complente aut non
84 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
plura praebente: puncto rubro (stigmate) ioe prope pyre-
noidem sito: nucleo ad latus oppositum cellulae intra partem
ovohanphow excavatam complexo: ciliis binis ca. 4% breviori-
bus quam cellula instructis.
Propagatio fit per cellulae vegetativae impigre natantis
divisionem in 4 vel 8 zoosporas, quae tegumenti scissura erum-
punt. Praeterea aplanosporae, primo virides, postea paulatim
ad colorem fulvum transmutatae, per contractionem protoplasti
in massam globosam ad cellulae partem anteriorem, ut in B.
abmurne: effictuntur
Generatio fit per c copulationem gametarum inter se subae-
u
longioribus praeditae, quaeque per divisionem cellulae vegeta-
tivae protoplasti etiam in 32 individua parturiuntur. Zygo-
sporae globose, tegumento levi vestitae, diametro ca. 12
Longit Spooner veg. 30-48, lat. 18-24. Longit. ganic
tarum 13-15n, t. 6-8,; longit. ciliorum ca.
Hab. in oe lacunarum saxearum a Gcelcia Aalesund,
Norway, 28 June, 1920; Cullercoats, Northumberland, England,
19 July, 1920; Plymouth, England, 11 August, 1920.
Because of the fact that unarmed individuals appear in
older cultures of B. submarina, one might be tempted to inquire
whether B. simplex is merely a form of that species. Never-
theless in the natural pure cultures seen both at Aalesund and
at Plymouth, B. simplex maintained its characteristic simplicity
of form from the earliest development of the daughter cells
to maturity, and even when it was found intermingled with one
of the other species it showed a distinctive character, parti-
cularly in the curving of the posterior horn, which rendered
it always recognizable. Again, the variability of form exhibited
by this new species might readily suggest for it a hybrid origin.
We know little of hybridization among algae, but a very inter-
esting preliminary report of its occurrence in Chlamydomonas
has recently been published by Pascher (7). It may be remarked
that Pascher’s heterozygous cells showed great diversity in
internal organization, while here in B. simplex only variability
of external form is found. From a careful consideration of all
the conditions of its habitat and its behavior, I am convinced
that B. simplex is to be regarded as a distinct species which
represents the transition from Chlamydomonas to the more
characteristic species of Brachiomonas.
There is some resemblance between this species and Chlamydo-
monas brachyura West (13) a form which developed in a tank of
seawater sent from Plymouth to Birmingham in 1915, and for
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 85
which I searched vainly at Plymouth in 1920. Nevertheless
since C. brachyura lacks a red eye-spot and is pointed instead
of rounded at the anterior end, we may perhaps find more
suggestion of the origin of Brachiomonas in the species mentioned
at the beginning of this section, namely,
CHLAMYDOMONAS CAUDATA Wille
This very interesting and apparently rare species was found
June 28, 1920, at Aalesund, Norway, in small rock pools of the
same character as indicated in the original account in 1903
(15, pp. 115-118, 135, pl. 3, f. g-11) and my determination of the
species was later confirmed by Professor Wille, from living mater-
ial which I carried to him in Christiania. In one pool was a practi-
cally pure culture, giving an intense green color to the brackish
water; in another pool close by was a considerable admixture
of C. subcaudata Wille, which was certainly the most abundant
species of the region, now as in 1902. Why C. caudata should
occur in only three or four out of many similar pools is an inter-
esting question. It was also surprising that it could not be
found on the English coast, where the conditions appeared to
be much the same, and where I did find several of the other
brackish-water species which were associated with C. caudata
at Aalesund. This species appears not to have been found
elsewhere than at this original station.*
The elongated oval posteriorly pointed form of the zoospores
plex is equally striking; so much so, in fact, as intimated above,
that upon superficial observation the two species might be
mistaken for closely related members of the same genus. Ordi-
narily in active individuals, especially in newly formed zoo-
spores, the protoplast conforms rather closely to the shape of
the cell-wall, being merely rounded at the point of insertion of
* Playfair, to be sure, has reported [Proc. Linn. Soc. New South Wales 43:
515. 1918] a form found appazently in fresh water of which he writes:
“The only tailed species of Chlamydomonas; it is impossible to make any
mistake in the identificacion.” Since, however, Playfair states that his
specimens lacked pyrenoid and stigma, and his figure shows a distinctly
different morphology of the anterior portion of the cell, including a different
position fof the nucleus, there can be no easiretpinse for trying. - identity his
is brought forward to support the attempt.
86 THE PH¥LOGENY OF THE GENUS BRACHIOMONAS
the cilia, and tapering to an acute point at the posterior end
Fic. 46). Older individuals, kept for two weeks or longer in
hanging drops, show the cell-wall much thickened, or the pro-
toplast contracted and rounded posteriorly and produced into
more or less of a beak at the anterior end (Fics. 47, 48). The
broad truncate anterior papilla of the cell-wall (Hautwarze)
impresses me as being rather more prominent than shown in
Fics. 46-50. CHLAMYDOMONAS CAUDATA Wille
46. Young cell, “dorsal view,’’ showing typica Iposition of nucleus, stigma,
and pyrenoid. _ a1 Older cell with thickened wall. 48. Similar cell in
“lateral view.’’ 49. Typical arrangement of daughter cells. 50. Unusual -
arrangement of daughter cells. All, x 720, approximately.
Wille’ s (15) figures, and the red eye-spot, first described as
langgestreckter’ and again as ‘oval oder stabférmig,’ was in
my mature specimens always a strongly oval disc, sometimes
with an anterior pointed end, showing its thinness in the fact
that the edge view was narrowly linear. These differences
may be readily accounted for by differences of magnification
during observation. The longitudinal streaking of the chro-
matophore noted in the original description, and even intro-
duced in Wille’s key to the species, was never distinguishable
in vigorous material of either this species or C. subcaudata in
1920; only certain individuals kept for ten days or more in
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 87
hanging drop cultures showed almost a lattice-like appearance
of the chromatophore. Although the chromatophore is hollowed
out about as far as the middle of the cell, or sometimes even
more deeply, nevertheless the position of the nucleus is notably
less central than might be expected, for it lies approximately
in the anterior third of the cell.
A point of considerable interest is the position of the stigma
and contractile vacuoles. Regularly the eye-spot is so placed
that when the cilia are stretched out in a plane parallel to the
stage of the microscope the stigma lies on the apparent right
side of the cell and in a slightly upper focus, or with the cell
revolved 180° the stigma lies on the apparent left side and at a
slightly lower focus; in such a position only one of the small
contractile vacuoles is visible. If the cell is revolved only 90°
from the first position, so that one of the cilia lies nearer the
observer and the other at a somewhat lower focus, the stigma
shows its disc-like shape in an upper central focus (FIG. 48)
‘ and the two contractile vacuoles may be seen side by side at
the same time, though one is always smaller than the other,
on account of the alternation in their pulsations. This position
of the stigma and cilia in one plane and the contractile vacuoles
in another plane perpendicular to it indicates a dorsi-ventral
character of the cell which appears to me to be the rule in many
species of Chlamydomonas, though it seems to have received
little or no attention from students of the group.*
The dorsiventral differentiation in Brachiomonas, as men-
tioned above and indicated in Fics. 6, 7, 13, 15, and 32, is
slightly different; there the cilia normally stretch out in a plane
perpendicular to that which passes through the stigma, pyrenoid,
and nucleus, and most commonly the zoospore comes to rest
with the stigma lying exactly in the middle on the ventral side,
or (less often) on the dorsal side, toward the observer. In
many of the figures which seem to indicate a different relative
position of the eye-spot and cilia, the latter were actually lying
in a plane oblique to that of the microscope stage, which could
not easily be shown in the drawings.
* There are, it is true, many published figures of Chlamydomonas which
show the two contractile vacuoles side by side apparently in the same plane
with the cilia; but in cells exhibiting so much movement it is very easy to
transfer what is seen in one view to a sketch representing in general a different
view, unless one is keenly on the lookout for such a point. Probably also
many of our somewhat classic figures are not made from camera lucida draw-
ings of quiescent individuals, as they should be to determine such features.
88 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
In the asexual reproduction of C. caudata, Wille describes
the division as longitudinal, after the disappearance of the cilia
of the mother cell, resulting in the formation of four zoospores.
On several occasions when I examined material with my travel
microscope in the field in late afternoon, or freshly collected
material in early evening, I found numerous individuals which
contained the four fully formed zoospores while the mother.
cell was still swimming actively. It was only in the more rarely
seen divisions of material kept for a day or two after collection
that I found non-motille cells in process of reproduction; such
cells may have lost the power of movement earlier because of
poorer aeration than in the normal habitat. This division of
actively motile cells I have found to be ‘normal in several other
species of Chlamydomonas. I was unable to find examples
which would actually show more of the details of the process
of division than indicated by Wille, who merely furnishes one
figure of completed division. I did, however, observe that a
large majority of cases showed two of the daughter cells with °
their anterior end directed forwards, and two with the anterior
end directed backwards (Fic. 49). In other species of Chlamydo-
monas such as arrangement of the daughter cells is character-
stically found when the first plane of division appears transverse
to the longitudinal axis of the mother cell, or when, though
fundamentally longitudinal, the plane becomes placed trans- _
versely by a rotation of the entire protoplast. During the
past season I have had the opportunity of observing this
latter method many times ina new species to be described in a
future paper: this species furnished one exceptional case in
which both divisions, carefully followed, were strictly longi-
tudinal, resulting in four zoospores all headed toward the
anterior end of the mother cell. This exceptional arrangement
I also found in rare instances in C. caudata (Fic. 50). Wille’s
figures (15, pi. 3, f. 8) which shows the same arrangement,
appears, therefore to be untypical, and I am obliged to believe
that the first division of this species, though properly longitudi-
nal, would regularly show as a transverse cleavage, in consequence
of a rotation of the protoplast to such an extent that its anterior
pole comes to lie in a position nearly in the middle of the length
of the cell.
Reichenow (9, 10, pp. 35-39) it is true, has expressed the view
that the division in all species of Chlamydomonas may be funda-
mentally Jongitudinal, an apparent transverse division being al-
THE PHYLOGENY OF THE GENUS BRACHIOMONAS 89
ways preceeded, in his opinion, by a rotation of the protoplast, so
that he believes the manner of cell division to have no phyloge-
netic significance. Attractive as this view is in its possibility of
simplifying the idea of direct transmission of polarity to the
daughter cells (and my own recent studies on Chlamydomonas
somewhat tend to support such a view) nevertheless we still
lack sufficiently definite and accurate observations to permit
the denial that an actual transverse division may occur in some
species.
If in C. caudata the first cleavage plane is actually or by
rotation of the protoplast transverse (as appears to me most
probable), and if we accept the current view derived from Dill
(4) that the most primitive species are those in which both
divisions are longitudinal, then C. caudata could hardly be re-
garded as an actual transitional form between the ordinary
ovoid Chlamydomonas type and the Brachiomonas group, since
all species of the latter genus appear to have retained the primi-
tive mode of division by two strictly longitudinal cleavages,
in spite of their advancement in other respects. evertheless,
in its general morphology, and in the similarity of its method of
formation of aplanospores (rarely found in other species of
Chlamydomonas), C. caudata must be very close to the ancestral
line which has given rise to the four forms of Brachiomonas
recognized above; and these present a rather close ascending
series, with perhaps a definite orthogenetic tendency beginning
with B. simplex, which is barely emerging from the Chlamydo-
monas stage, through B. submarina f. obtusa and then B. sub-
marina (type) to B. gracilis, an extremely slender form, so far
removed from any Chlamydomonas type as to justify thoroughly
the retention of Brachtomonas as a genus.
In certain graphic schemes of the phylogeny of the chlamydo-
monads—e. g. those of Wille (15) and West (14, p. 163)—the genus
Lobomonas Dangeard is indicated as representing an inter-
mediate stage between Chlamydomonas and Brachiomonas; but
the two known species of Lobomonas, as well as two new species
to be described by the writer in the next number of this journal,
are very minute forms, and in their general morphology exhibit
by no means so good a connection as is shown by C. caudata
and B. simplex. Lobomonas must be regarded as a special
offshoot from the Chlamydomonas line not leading to anything
higher, so far as we know at present.
ARNARD COLLEGE,
CoLUMBIA UNIVERSITY
—
WwW N
hed
—
a
—
i)
THE PHYLOGENY OF THE GENUS BRACHIOMONAS
LITERATURE CITED
. Bohlin, K. Zur Morphologie und Biologie einzelliger
Algen. Ofvers. K. Vet. Akad. why cea 54: 507-529.
1898.
. Chodat, R. Algues vertes de la Suisse. Berne. 1902.
. Dangeard, P. A. Mémoire sur les eae spears
ou l’histoire d’une cellule. Le Botaniste 6: 65-292. f. 2
ao. 1899:
Dill, O. Die Gattung Chlamydomonas und ihre nachsten
Verwandten. Jahrb. Wiss. Bot. 28: 323-358. 1895. .
. Harper, R. A. The structure of protoplasm. Am. Jour.
Bot. 6: 273-300. 1919.
. Lagerheim, G. Studien iiber ere Cryptogamen.
Troms¢ Mus. Aarhefter 17: 1-24.
. Pascher, A. Uber die Kreuzung eee haploider —
G
Organismen: Chlamydomonas.. Ber. Deutsch. Bot. Ges.
34: 228-246. f. 1-5. 1916.
Printz, H. Kristianatraktens Protococcoideer. Vid.-Selsk.
Skrift. I. Mat. Naturv. Klasse 1913%: 1-123. pl. 1-7 +f.
1, 2 19.
Reichenow, E. Untersuchungen an Haematococcus pluvi-
alis und eingen anderen Flagellaten. Sitz.-ber. Ges. Naturf.
Freunde Berlin 1909: 85-91. 1909
————. Untersuchungen an MHaematococcus pluvialis
nebst Bemerkungen iiber andere Flagellaten. Arb. Kais.
Gesundheitsamte 33: 1-45. pl. 7, 2. 1909.
. Thompson, D’Arcy W. On growth and form. Cambridge.
sg a
. West, G. S. Some critical green algae. 2 Brachiomonas
submarina. Jour. Linn. Soc. Bot. 38: 281-283. pl. 20.
4 9 -T0:..: 1908.
. Algological notes XX, XXI. Jour. Bot. 54:
ny Too ee ‘
. Cambridge Botanical Handbooks. Algae 1.
Cambridge. 1916.
Wille, N. Algologische Notizen XI. Uber die Gattung
Chlamydomonas. Nyt Mag. Naturvid. 41: 109-162. /l.
3: J. Fak 1905.
Description of plates 3 and 4
All drawings made with the aid of camera lucida from living matezial
in hanging drop cultures (or in some cases freshly fixed with osmic acid vapor):
PLATE 3 done with Leitz 1/12 in. oil immers. obj. and compens. oc. 6, magnifica-
THE PHYLOGENY OF THE GENUS BRACHIOMONAS gI
tion about 1440 diameters; plate 4 done with Spencer 2 mm. oil immers. obj.
and Leitz compens. oc. 6, magnification about 1300 diameters; both reduced
in reproduction to about 720 diameters.
PLATE 3
Fics, 1-7. BRACHIOMONAS SUBMARINA Bohlin, type form
Fic. 1. Vegetative cell; posterior horn filled with colorless cytoplasm.
BiG oe romatophore filling the extremities: Dec. 191
Fic. 3. Colorless cytoplasm in extremities; stigma and pyrenoid lying
on the ee side of the cell: March, 1921.
Fic. 4. Anterior polar view of similar cell; vacuoles in chromatophore.
Fic. 5. Slender form similar to Bohlin’s f. 7b of B. gracilis (fixed with
Oosmic wie vapor).
IG. [oe slender form: showing typical position taken by
living zoospore temporarily at rest, the pyrenoid and stigma lying in the
center of the under side (osmic v. sie): April 1921.
1G. 7. Anterior polar view of similar cell, showing cilia stretching out
n plane ae to that in which the nucleus, pyrenoid, and stigma
he pically lie
Bes 8-27. BRACHIOMONAS SUBMARINA forma OBTUSA Hazen
Fic. 8. Chromatophore orci s arms and horn filled with colorless
cytoplasm (osmic vapor): Novem 9.
Fics. 9, 10. Arms empty, ‘olen cytoplasm extending into posterior
horn (osmic vapor).
Fic. 11. Anterior polar view of similar cell, living.
Fic. 12. Arms and horn nearly empty (osmic vapor).
IG. 13. Posterior is view of similar cell, living, showing typical re-
ia positions of organs as in Fic. 7; a large colorless cytoplasmic region
und the nucleus.
1G. 14. Older cell showing no trace of arms.
Fic. 15. Cell from old culture with only a Snes of lateral arms;
stigma = pyrenoid lying underneath the nucleu
Fic. 16. Senescent = with contracted RES preparing to form an
pean hae December, 1919
Fic. 17. Young cell siete nucleus posterior to pyrenoid.
Fic. 18. Posterior polar view of similar
Fic. 19. Fi irst division of motile cell; 10:30 P. M. (osmic vapor).
ed esent
FIGs. 22-27. a. stages in division resulting in eight zoospores
linen IgIg): . 22 at 2:40 P. M.; FiG. 23 at 3:40; FIG. 24 at 4:50;
FIG. 25 at 5:30; a IG. oe at 6:20; Fic. 27 at 7:40. The zoospores escaped
Stig Petia
PLATE 4
BRACHIOMONAS SIMPLEX Haz
Drawn 29 June—4 July, 1920, Aslan: Norw.
Fics. 28, 29. Typical mature vegetative cells: in Fic. 29 ae stigma lies
slightly cade
Fic. 30. Anterior polar view of similar cell.
92 THE PHYLOGENY OF THE GENUS BRACHIOMONAS
Fic. 31. Cell on which the ‘‘bumps” representing !ateral arms are un-
developed.
Fic. 32. Anterior polar view of similar cell.
Fic. 33. Protoplast ertracted from one of the “‘bumps” (osmic vapor).
Fic. 34. Large older cell with more posterior Lees id.
Fic. 35 Eight zoospores in motile cell; 12:45 A
Fic. 36. Four zoospores in motile cell.
Fic. 37. Young zoospore free; chromatophore filling posterior horn.
Fic. 38. Three of the four zoospores have escaped through a triangular
rent in the wall of the mother cell.
Fic. 39 Similar to Fic. 31, but with more typical position of pyrenoid
(osmic vapor)
Fic. 40. Thirty-two gametes in motile mother cell; 10 A M
Fics. 41, 42. Gametes at beginning of conjugation (osmic vapor).
Fic. 43. Motile zygote, momentarily resting; nuclei not fused.
Fic. 44. Zygote, cilia having disappeared, nuclei fused; two stigmata still
present.
Fic. 45. Zygospore twenty-four hours after conjugation.
In the living = the pyrenoid a appears upon casual
observation to be n because imbedded in the green chrmatophore It
is believed, Laer tat the printing of ie pyrenoids the same sepia
as the cell-wall and protoplasmic outlines represents more truthfully their
real nature
Note. As fina] proof of this paper is received, the writer has
just made a mid-April excursion to the Massachusetts coast,
where he has collected typical Brachiomonas submarina Bohlin
from Cape Ann to Buzzards Bay, at Rockport, Gloucester,
Marblehead, Nahant, and Fair Haven. There has been no in-
dication of other species of Brachiomonas or of Chlamydomonas
caudata at these stations.
a.
Butt. Torrey Crus VOLUME-49, PLATE 3.
-7. BRACHIOMONAS SUBMARINA Boxun
8-27. BRACHIOMONAS SUBMARINA rorma OBTUSA Hazen
2 i. hase AS ee
—
Burs. Torrey C nee oe Y fos veare 2
ORREY Crue VOLUME 49, PLATE 4 _
BRACHIOMONAS SIMPLEX Hazen
New records and other notes on North Dakota plants*
O. A. STEVENS
The present paper is supplementary to Bergman’s Flora of
North Dakota (in Sixth Bien. Rept. Agr. Coll. Survey [1911-
1912]. 1918) and, as such, will follow the same arrangement and
nomenclature. The purpose is to record additional species,
designated by a dagger (t), and other notes of special interest.
During the years 1910 to 1920 I have had opportunity to visit
nearly every part of the state and, incidental to economic work,
to obtain many specimens of interest. Some of these were
noted in the Flora and others have been reported by Lunell
(Am. Mid. Nat. 4-6. 1917-1919). I have also included in the
paper a few corrections which have come to notice and revised
certain groups in which I have become especially interested.
Where not otherwise stated, the specimens listed have been
collected by myself.
LEPIpIum RAMOsIssIMUM A. Nels. I do not know why
Bergman reduced this to a synonym of L. densiflorum Schrad.
The two species are very different and may be distinguished as
follows:
L. densiflorum. Annual, stem usually simple below and with
several long racemes above; basal and lower stem leaves
serrate; pods distinctly margined and broadest toward the
apex.
L. ramosissimum. Biennial, bushy branched from the base;
asal and lower stem leaves pinnatifid, the lobes linear or
oblanceolate; pods not margined above, elliptic in outline.
The following specimens belong to L. ramosissimum: Tower
City, Bergman 870; Valley City, Aug. 13, 1912, (first year
rosettes); Courtenay, June 20, I9IT; Kensal, July 26, 1915,
(rosettes, also flowering stems) ; Manfred, July 12, 1913; Sykes-
ton, July 15, 1891, Bolley; Sheyenne, July 4, 1908, Lunell;
Leeds, Aug, I, 1912; Bathgate, July 22, 1892, Lee; Ganbetta;
Bell 264; Rugby, Bergman 2611. This species is quite common
throughout the central part of the state, growing about streets
and dooryards rather than in fields. At Langdon on July 19,
1918, I noticed the rosettes of L. ramoisssimum fairly covering
* Contribution from Department of Botany, North Dakota Agricultural
College and Experiment Station.
93
94 STEVENS: NORTH DAKOTA PLANTS
the ground. In a dooryard at Courtenay the species grew
mixed with L. densiflorum. The latter had racemes about 3 cm.
long with numerous well-developed pods, while the plants of L.
ramosissimum had only a few half-developed pods.
+Lepmrum Drapa L. Several years ago (Ann. Rept.
North Dakota Exp. Sta. 22: 80. 1912) I called attention to the
possibility of this being introduced in Turkestan alfalfa seed.
On June 6, 1918, a flowering plant was sent me by Mr. H. A.
Nelson of Ray. [A well established colony has since been found
along the railroad track at Moorhead, Minnesota, May 22, 1921.|
+ Sisymprium Lorsenit L. Determination verified by Paul
C. Standley and specimens deposited in herbaria of the U. 5.
Sy rocer Museum and the New York Botanical Garden.
Annu dm. high, sparingly hirsute with simple flattened
hairs, dene less abundant above but present on upper stems
and pedicels; leaves runcinate, the terminal segment hastate
and irregularly ceo lower stem leaves sometimes I dm.
long; racemes becoming 3-4 dm. long, pedicels widely spreate
one third to one half the length of the slender pods, which a
ascending and about 3 cm. long with three-nerved valves
Devil’s Lake, July 15, 1920 (common about the streets);
Tappen, July 12, 1919 (a single plant in the street; several in
an old field of Bromus inermis Leyss. several years before).
The description was drawn from the growing plants and speci-
mens collected at Devil’s Lake. The plant has somewhat the
aspect of Sophia intermedia Rydb., but the leaves are quite
different and the racemes are fewer but longer and coarser.
The relative height of flowers and young pods mentioned by
Koch and others seems scarcely distinctive. In this plant the
pods do not surpass the flowers in normally developed racemes,
but examples are frequent where they do so, apparently as a
result of arrested development.
ErucastrumM Po.iicait Schimp. & Spenn. Park. River,
Aug. 8, 1913; Williston, Aug. 15, 1915; Oakes, July 18, 1919;
Cooperstown, Aug. 25. 1919. This has now been found at
many places, chiefly along the railroad tracks. The seeds have
been identified in several samples of timothy and millet coming
from near Grand Forks or a short distance north or south of
this locality. The pods are not flattened, as stated in Bergman’s
key, but rounded, the valves with a rather prominent mid-
nerve; racemes leafy bracted. The plants grow vigorously in
late fall as shown by a flowering specimen collected at Fargo,
Oct. 26, 1914.
STEVENS: NortH DAKOTA PLANTS: 95
MALVA VERTICILLATA L. I have not seen the specimen
which Bergman referred to this species but am inclined to think
that it was M. borealis Wallm., which is the common plant at
Fargo and other places in the state. A specimen of what seems
to be M. crispa L. + was sent by A. G. Sorlie, Grand Forks,
Oct. 10, 1916.
EuPHORBIA Esuta L. Wahpeton, June 3, 1920, Geo. P.
Wolf; Hazelton, July 15, 1919, in field of Bromus inermis; New
Rockford, July, 1911, J. R. Campbell; Mona, June 5, 1914,
Albert Graves; Ray, July 2, 1917, H. A. Nelson. In one field
near Fargo patches are scattered over 20 acres. Apparently
a bad weed, spreading by the roots. Time and manner of in-
troduction unknown.
} SILENE DICHOTOMA Ehrh. Edgeley, July, 25, 1917,
Brenckle; Langdon, July 19, 1918; Regent, July 20, 1916, A.
C. Goldtrap. In timothy fields. Plants grown from seed
proved to be coarse biennials larger than S. noctiflora.
+ SILENE Faparia (L.) Sibth. & Sm. Specimens determined
by Paul C. Standley.
Biennial; glaucous, much branched, 6-9 dm. high. Basal
and lowest stem leaves narrowed at the base, tances stem leaves
ovate, sessile; flowering stems dichotomous wit flower
in the fork, each branch with several remote clusters of three
to seven flowers each. Flowers about I cm. wide, white, opening
at night; petals cleft to the throat, not appendaged, the lobes
oblanceolate, rounded at the apex; calyx indistinctly fifteen-
ribbed, closely investing the ovate horny capsule, but not
especially enlarged in fruit
Venturia, July 16, 1919; Oakes, July 18, 1919. Lower stems
and leaves with very much the aspect of Vaccaria Vaccaria.
A specimen in the herbarium of Dr. J. F. Brenckle was collected
at Kulm by him in 1916, and he states that he had found the
plant there previous to that date. Apparently well established
at Oakes and Venturia along roadsides. Dr. Standley writes
that this is probably the first time it has been collected in this
country.
+ LycuHnis ALBA Mill. Devil’s Lake, July 13, 1920.
CHENOPODIUM
For several years I have been interested in C. album and
related forms, especially in regard to seed characters. The
plants are rather difficult to collect as the seeds are slow in
maturing and the lower leaves are likely to be lost by the time
the fruiting branches are well matured. With respect to the
adherence of the pericarp it is to be noted that it rubs off more
easily in fresh material.
96 STEVENS: NorTH DAKOTA PLANTS
In the fall of 1916 I collected twenty-six specimens in the
vicinity of Fargo and submitted them to Dr. Standley, who
determined them as follows:
C. album (4);
+ C. paganum Reichenb. (11);
C. paganum, approaching C. album (8);
+ C. ferulatum Lunell (3).
Apparently C. paganum is the commonest form here. I
have not as yet been able to make much further progress in
clearing up the relations of these forms but wish to offer what
few notes I have.
C. ferulatum (which Standley notes is perhaps not sufficiently
distinct from C. album) is separated by the fruiting calyx being
open, exposing the fruit. While this does not seem to be quite
constantly true for C. ferulatum, I find it also ina rather variable
degree in the C. album specimens, being most prominent in
well-matured plants. No. 11, determined as C. paganum, has
calyces closed in specimen collected Aug. 25, but fruit from same
plant collected in September shows them quite open. As to C.
paganum, most of the plants were either so badly infested by an
insect or so late in maturing that very little mature seed was
obtained. From material at hand I would describe the seeds
as follows:
C. album—1.2-1.4 mm. wide, upper surface flattened and with
shallow, oblong pits (which show through the pericarp).
C. paganum—1.2-1.4 mm. wide, upper surface dull, neither
flattened nor pitted.
C. ferulatum—1.2-1.4 mm. wide, similar to C. PENH:
C. leptophlyllum—o.9 mm. wide, not flattened, hini
The size of the seed of C. paganum is given by Standley
(N. Am. Fl. 21: 21. 1916) as 1.3-2 mm. In one of the twenty-
six specimens, a plant collected near the woods along the Red
River, the seeds are variable in size, measuring I.4-2 mm., but
in all the others they are scarcely larger than those of C. album.
The three specimens of C. ferulatum differ among themselves.
One, collected in an old garden (No. 3), which I take
to be typical, is pale grayish green, simple below and with short
branches above, the leaves oblong-rhombic with four or five
short teeth on each side; another (No. 9, labelled by Standley
as apparently a form of C. ferulatum) is bushy branched with
narrow entire leaves. The third (No. 2) is in a rather advanced
condition with lower leaves mostly fallen; the flowering branches
STEVENS: NortH DAKOTA PLANTS 97
are unusually thick and dense and the seeds are pitted on the
upper side. I am inclined to think it is a plant of C. album with
an unusual amount of well-matured fruits and therefore with
the spreading calyces unusually prominent.
The seeds of these species are described by Standley as
“nearly smooth, black and shining”’ except in C. ferulatum,
where they are said to be “‘puncticulate,” but I find in our
material that C. album has pitted seeds and C. ferulatum
smooth (excepting as in No. 2 just noted; type material from
Lunell also examined). The seeds of C. leptophyllum seem to be
quite easily distinguished from those of the other common field
species by their smaller size.
{| CHENOPODIUM PRATERICOLA Rydb. Narrows, Aug. 12,
1913, Lunell.
} ATRIPLEX PATULA L. Wahpeton, Aug. 6, 1919. Alonga
street by buildings. Dr. Standley, who examined a specimen,
writes that this species is perhaps only a form of A. hastata L.,
but we seem to have no other specimens which would be referred
to this form.
} SUCKLEYA SUCKLEYANA (Torr.) Rydb. Belfield July, 4,
1914.
CORISPERMUM
Following Standley’s treatment of this genus (N. Am. FI.
21: 79-80. 1916) I would place our material as follows:
C. hyssopifolium L. Mandan, Wright 610.
+ C. nitidum Kit. Sandhills near Anselm, Aug. 21, 1918.
+ C. villosum Rydb. Sandhills near Anselm, Aug. 21,
1918; also all other specimens referred by Bergman to C. hyssopi-
folium, except Wright 610. The Fargo specimen of C. villosum
was collected along the railroad (doubtless introduced in gravel)
and no plants have been observed since. It was a well matured
plant with very little pubescence except on the bracts. The
Anselm plants are young, upright, rather slender, and densely
pubescent.
PotyGonumM HypropirpEr L. Hankinson, July 30, I9g19.
Lunell (Am. Mid. Nat. 5: 184. 1918) has referred the Ft. Ransom
specimen to P. punctatum leptostachyum (Meisn.) Small, but
it has dull akenes and flowers in the lower axils, as in P.
Hydropiper.
ASCLEPIAS SYRIACA L. Westfield, July 16, 1919. This does
not seem to be common so far west. The specimen reported
98 STEVENS: NortH DAKOTA PLANTS
by Bergman from Morton County (Bell 30) is undoubtedly A.
speciosa Torr.; it is sterile but has the characteristic pointed
leaves of that species. The specimen reported by Bergman as
possibly a hybrid of the two species seems scarcely different
from A. syriaca.
+ ASCLEPIAS PuMILA (A. Gray) Vail. Mr. W. W. Eggleston
has examined our material of A. verticillata L. and refers the
following to A. pumila: Esther, Bell 542; Fleak, Bell 1370.
} ACERATES LANUGINOSA (Nutt.) DC. Janesburg, Bell 3o,
of Bergman’s catalogue should be referred to this species instead
of to A. viridifiora Ell.
+ VERONICA MARITIMA L. Kongsberg, Sept. 1918, Fred
Schmidt, Jr. A few plants in a-grove of planted trees. Deter-
mined by F. W. Pennell.
CuscuTA ARVENSIS Beyr. The Logan County record is by
Brenckle, Aug. 18, 1912. We have also the following records:
McLeod, Bell 385: Valley City, July 26, 1913; Enderlin, Aug. 20,
1918. Dr. T. G. Yuncker, who has examined portions of the
Logan County and Valley City specimens, writes that he would
call them C. pentagona Engelm. The other specimens seem to
be the form which he calls C. calycina Engelm. The former
were growing upon plants of the dry prairie, while the latter
preferred those of riverbanks and similar localities.
Cuscuta Gronovit Willd. Fargo, Aug. 14, 1890, Waldron,
is the only specimen which I consider typical. When preparing
my paper on Cuscuta (Am. Jour. Bot. 3: 185-188. 1916) I was
unable to find fresh material, but all that I have found at Fargo
since that time agrees better with what I had called C. plattensis
A. Nels. The habitat of C. plattensis, as stated by Nelson,
hardly indicates this plant and I become doubtful of its identity.
Dr. Yuncker writes me that an examination of the type of C.
plattensis shows my plant to be quite different, although the
description of the species as given by Nelson agrees. He refers
my specimens to C. Gronovii var. curta Engelm. Ina flax field at
Ft. Ransom, where considerable damage was caused, this form
and C. Coryli Engelm. were both present, sometimes separate
and at others tangled together on the same host plant. The
field was a small one next to the woods of the river.
STEVENS: NortH DAKOTA PLANTS 99
PHLOX
Following the treatment of the western forms by E Nelson
(Ninth Rept. Wyoming Agr. College, 1899), our plants would
seem to be separated more satisfactorily as follows:
Leaves 5-10 mm. long; flowering stems one-flowered. P. Hoodii.
Leaves 1-2 cm. long; flowering stems two-to several- flowered: more up-
right, the bark peeling off in shreds. P. andicola.
PHLOX ANDICOLA (Britton) E. Nelson. P. Douglasii of
Bergman’s Flora, in part. This seems to be better separated
from P. Hoodii Rich. by the above characters than by the size
of the flowers. The Medora and Washburn specimens referred
by Bergman to P. Douglasii Hook. are evidently P. Hoodii.
+ LappuLA CENCHRUSOIDES A. Nels. Marmarth, July 4,
1918, in sandy soil of river valley. Habit of growth similar to
L. occidentalis (Wats.) Greene, nutlets similar to those of L.
Lappula (L.) Karst. but larger, the spines longer, and with a row
of elongated tubercles on the middle of the back of the nutlets.
The following, previously referred to L. Lappula, also belong
here: Medora, Bergman 1276; same locality, June 19, 1910;
Williston, Bell 25.
{+ LappuLA CUPULATA FOLIOSA (A. Nels.) Nels. & Macbr.
Marmarth, same as preceding. Often three of the nutlets of
a flower have the spines confluent, forming a spreading border,
the fourth nutlet with simple spines. In the specimens col-
lected, however, there seems a decided tendency for the spines
to be simple on all nutlets of the lower flowers. There are also
nutlets with spines of intermediate form, broadened but not
united. Counts on three plants show nutlets as follows:
Plant No. 1—8o united, 71 simple, 40 intermediate;
Plant No. 2—61 united, 106 simple, 34 intermediate;
Plant No. 3—47 united, 47 simple, 14 intermediate.
The color of the corolla is an uncertain character, white
flowers often appearing bluish in the dried specimens.
7+ LAPPULA TEXANA HOMOSPERMA (A. Nels.) Nels. & Macbr.
Marmarth, same as preceding, occasional plants in patches of
L. occidentalis, all plants dried up and only fruits collected.
From a planting of fruits the following season about twenty-five
plants of L. occidentalis were obtained from fruits of that species.
No plants were secured from the few fruits of the other two
species.
+ LAPPULA FLORIBUNDA (Lehm.) Greene. Lake Ibsen (Leeds)
July 4 and 17, 1914, Lunell; Pleasant Lake, June 29, 1920.
100 STEVENS: NoRTH DAKOTA PLANTS
+ CRYPTANTHA CALYCoSA (Torr.) Rydb. Bowman, June 23,
1918, in loose burned clay around large boulders of the same
material.
AMSINKIA MeEnzigEsiI (Lehm.) Nels. & Macbr. A. imter-
media F.& M. Rugby, July 7, 1917, and July 27, 1918,a number
of plants along the railroad track near the station; the first col-
lection in flower, the second in fruit. Bergman’s specimen from
Pembina is evidently this species also, instead of A. lycopsoides
Lehm. |
+ LAMIUM AMPLEXICAULE L. Langdon, July 18, 1918, a
quantity in a dooryard.
+ DRACOCEPHALUM THYMIFLORUM L. Belfield, July 4, 1914,
several plants in a field of Bromus inermis. Several other
introduced plants were also in the field: Potentilla argentea L.,
Campanula sibirica L., Chrysanthemum Leucanthemum L. and
Bromus tectorum L. To be consistent with Bergman’s nomen-
clature this plant should perhaps be referred to Moldavica but
I have not thought it desirable to make such a change at this
time. The plant is different in appearance from D. parviflorum
Nutt., being slender and having very small flowers.
+ Lycopus communis Bicknell. Pleasant Lake, July 26,
1912, Lunell; Anselm, Aug. 29, 1920 (common in alder swamps).
The Anselm plant has been verified by P. A. Rydberg.
} SALVIA LANCEOLATA Willd. In fields. Carrington, Aug.
27, 1919; Mandan, Sept. 13, 1920. :
CHAMAERHODOs ERECTA (L). Bunge. This was included in
the seventh edition of Gray’s Manual, the record being appar-
ently based on a specimen collected near Crookston, Minn.
(Minnesota Bot. Studies 2: 584. 1901). I had thought that
it might have been introduced in gravel as railroad ballast, but
so far as I have been able to learn the locality mentioned is a
gravel pit. It is probably one of the instances of a plains species
occuring on the eastern edge of the Red River Valley. I have
not seen the plant east of the hills along the Sheyenne River at
Sheyenne and Valley City. :
} CRATAEGUS MOLLIs T. & G. Fargo, May 26, 1917. One
tree is in a thicket east of the fair grounds, and quite a number
occur in a bend of the river two miles farther north.
} Mepicaco FaLcaTta L. Oakes, July 18, 1919, a plant
along the railroad tracks.
STEVENS: NorTH DAKOTA PLANTS IOI
+ ASTRAGALUS PARVIFLORUS (Pursh) MacM.. Marmarth,
July 3, 1918. Quite common on the higher parts of the hills.
} HEpysARumM sp. Sentinel Butte, Aug. 0916, Brenckle, a
single pod collected. Reported by Arthur (N. Am. FI. 7: 450.
1921) as H. cinerascens Rydb.
} EVONYMUS ATROPURPUREUS Jacq. Owego, Sept. 1916,
R. A. Shunk.
} Lomatium MAcROCARPUM (Nutt.) Cov. & Rose. Dunseith,
May 18, 1918, Lunell, and fruits collected at Minot in June, 1910,
seem properly referred to this species.
{ Stcyos ancuLatus L. I have seen a specimen collected
by A. H. Shunk along the Sheyenne River near Anselm.
{| CAMPANULA SIBERICA L. Belfield, July 4, 1914, a single
plant found in a field of Bromus inermis.
} XANTHIUM PENNSYI.VANICUM Wallr. A specimen in the
Gray Herbarium from Leeds, Aug. 21, 1902, Lunell, is referred
by Millspaugh and Sherff (Field Mus. Nat. Hist. Bot. Ser. 4:
33. I919) to this species. Some specimens of Xanthium
collected along the river at Fargo in 1919, to show variations in
burs, were determined by Sherff as X. italicum Mor.
canadense of the Flora) and X. acerosum Greene, t with the com-
ment that the latter was perhaps not distinct.
{| CHRYSOTHAMNUS NAUSEOsUS (Pall.) Britton. Williston,
Aug. 11, 1915, on the hills along the Missouri River about fifteen
miles southeast of the city. A plant of quite different appear-
ance from C. graveolens (Nutt.) Greene, which is common there
and on the buttes in the bad lands—the crown low, and with
gray branches only 1-2 dm. long. This is the plant referred by
Lunell (Am. Mid. Nat. 5: 41. 1918) at my suggestion to C.
formosus Greene, but it evidently is not that species. ;
HELIANTHUS GIGANTEUS L. The form referred to in Berg-
man’s Flora under this name is quite common in the central
(and western?) part of the state, but I am as yet in doubt as
to its status. It is evidently Lunell’s H. mitidus (Am. Mid.
Nat. 1: 235. 1914). Specimens from Valley City and New
Rockford were determined by Standley as H. tuberosus. The
Fargo specimens are H. Maximiliani Schrad.
HELIANTHUS GROSSE-SERRATUS Martens. The Fargo speci-
men of the Flora is certainly H. Maximiliani. The stem
is only slightly hispid above and the leaves are broader and more
nearly flat than usual. The Kenmare specimen (Bergman 2744)
belongs to the preceding form.
102 STEVENS: NortH DAKOTA PLANTS
+ MaApIA GLOMERATA Hook. Spring Brook, Aug. 17, 1915,
a quantity in a prairie slough near the town.
+ ARTEMISIA PABULARIS (A. Nels.) Rydb. Mandan, Sept. I1,
1020. Determined by Rydberg. There seems to be no other
specimen among our material which resembles this. Mr.
Thysell of the Northern Plains Station showed me the plants,
only two places where they were growing being known.
} SENECIO MANITOBENSIS Greenman. Bottineau, July 8,
1917. Determined by J. Lunell. In the meadow of a dried up
pond in the Turtle Mts.; Towner, July 13, 1911, Lunell (speci-
men in the Gray Herbarium).
7 CENTAUREA PIcRIS Pall. Spring Brook, Aug. 11, 1918.
Determined by Standley. Received from Jacob Widman who
referred to it as abundant in a field. Especially a few years
ago the akenes were often found in Turkestan alfalfa seed, but
this is apparently the first record of its having become established.
SONCHUS ARVENSIS L. Ellendale, July 24, 1919. Several
spots around buildings in the town.
{SONCHUS ULIGINOSUs Bieb. S. arvensis of Bergman’s Flora,
Fargo, Aug. 1, 1916; Oakes, July 18, 1919; Crosby, June 7, 1919,
J. H. Phelps; Williston, Aug. 1915. Specimens of this and of
the preceding were examined by Standley, who stated that no
North American specimens were in the U: S. National Museum
collections. A specimen sent to the New York Botanical -
Garden was reported by Rydberg as apparently new to this
country*. This seems rather strange in view of the fact that it
is the common form through the Red River Valley, and westward
in the northern part of North Dakota to somewhat beyond Devil's
Lake, southern Manitoba, and western Minnesota. It is said
to have first appeared at-Portage Plains, Manitoba, about 1900.
This form differs from S. arvensis in the absence of glandular
hairs on the upper stems and involucres. While I have not
been able to compare carefully fresh material, I believe the heads
are smaller, paler, and the rays more inclined to become recurved.
Vegetative reproduction is by long horizontal. roots and not
rootstocks as commonly stated. The development of the
terminal head is often stopped before flowering and several flower-
ing branches arise just below it.
*
mall has recently listed the species as an addition to the American
flora, from Pennsylvania, where it was collected in 1921 (see Torreya 21: 100.
1922).
STEVENS: NORTH DAKOTA PLANTS 103
Lactuca virosA L. Many European authors describe this
as having horizontal leaves and black, broadly margined akenes.
This does not apply to our plants. The pinnatifid-leaved form
is only occasional in the state (Kulm, Brenckle 782; Bottineau,
July 23, 1918; Fargo, Aug. 17,1920). Pammel’s notes (Rhodora
20: 180-181. 1918) on the replacement of the entire-leaved form
by the pinnatifid in Iowa suggest that it will be interesting to
watch for a similar case here.
7 CREPIS OCCIDENTALIS: Nutt. Bowman, June 23, 1918.
Quite common on Twin Buttes.
+ CREPIS CAPILLARIS Roth. Fargo, Aug. 26, 1920; Willow
City, F. M. Rich.
} SPIRODELA POLYRHIZA (L). Schleid. Fargo, Aug. 30, 1919,
a few plants among masses of Lemna.
+ CYPERUS DIANDRUS Torr. Anselm, Aug. 1916, R. A. Shunk.
{+ CYPERUS ESCULENTUS L. Fargo, Oct. 2, 1920.
ScIRPUS VALIDUS Vahl. I cannot support Bergman’s separa-
tion of most of our material as S. occidentalis (Wats.) Chase.
There do seem to be two forms present, one with spikelets as
figured in Gray’s Manual (ed. 7, f. 290), the other with more
rigid panicles and longer spikelets. The specimens, however, do
not match Bergman’s separation. Both of these forms were
found in a small patch at Oakes on July 18, 1919, the first
occupying a definite part of the patch. Specimens of each were
identified by Agnes Chase as S. validus.
CAREX OBTUSATA Liljebe. Verified by Standley. Fargo,
June 24, 1920. Quite abundant, at least in one place in low
prairie. Kensal, Bergman, 1744, is alsothis in stead of C.
_ stenophylla Wahlenb.
The following additional species of Carex were recorded by
Lunell (Am. Mid. Nat. 3: 234-237. 1916) from specimens deter-
mined by Mackenzie: C. Hookeriana Dewey, C. athrostachya
Olney, C. tenera Dewey, C. praticola Rydb., C. Emoryi esis
C. scirpiformis Mackenzie, C. Parryana Dewey, C. laevic
Dewey and C. atherodes Spreng.
+ SYNTHERISMA SANGUINALE (L.) Dulac. Fargo, Oct. 2,
1920.
+ PANICUM PERLONGUM Nash. Verified by Hitchcock.
Fargo, June 24, 1920. Very abundant in one place in low
prairie.
104 STEVENS: NoRTH DAKOTA PLANTS
+ SPOROBOLUS ASPER (Michx.) Kunth. Mayville, Aug. 21,
1919: Fargo, Aug. 16, 1920; Steele, Sept. 10, 1920.
FESTUCA VIRIDULA Vasey. Lunell (Am. Mid. Nat. 4: 224
1917) has reported this species from Dunseith. I have examined
carefully a specimen received from him (‘‘det. by U. 5. Dept.
Agr.’’) but can see no reason why it should not be referred to
F. Hallii (Vasey) Piper.
Bromus jJAPonicus Thunb. Fargo, Oct. 1918. Determined
by Agnes Chase. All of the specimens referred by Bergman to
B. commutatus Schrad., and several others not reported, seem
to belong here.
Fae Lo.ium ricIpUM Durutet Hook. Determined by Agnes
ase
Annual: culms 3 to 4 dm. high, rather a somewhat
scabrous above; leaves upright, 5 mm. wide, 1 dm more long,
glabrous or nearly so; glume about three-fourths ; as lide as the
well developed itkalate equalling the younger or shorter ones;
spikelets five- to ages habia the larger 2 cm. long; lemmas
o 10 mm. long bearing an awn 7 to 12 m ng.
Described from a specimen collected ie 7, 1919, grown
from seeds found in a sample of wheat from the northeastern
part of the state. An early maturing annual, not so large as
L. temulentum L. and quite different in the longer, lanceolate,
awned florets. The culms commonly bear a branch from the
first node, some of these in the specimen cited being only 1 dm.
long with spikes barely protruding. The Milton specimen cited
by Bergman under L. temulentum belongs here, both it and plants
from the 1919 culture having been examined by Mrs. Chase.
The plant seems well established in that vicinity as evidenced
by these cases, by material received at one or two other times
and by seeds found in several samples of wheat.
+ Lottum REMoTUM Schrank. In flax plots, Mandan, 1918.
AGROPYRON
Three species of this genus are of great economic importance
in North Dakota. Having had occasion to pay particular
attention to these, I find that as a rule, descriptions and figures
seem to have been made from immature material, causing certain
characters to be overlooked. The spikelets are well figured by
Hillman (Bur. Pl. Ind., U. S. Dept. Agr. Circ. 73. 1911). I
offer the following descriptions:
STEVENS: NoRTH DAKOTA PLANTS 105
the aie ee that ihe break away readily, with a rounded
base), and with a strong transverse apis rte on mm
above the base. Glumes from half as long to nearly equalling
the spikelet, strongly nerved, with about a dozen short teet
on the keel* near the apex x which i is acute or obtuse, often bearing
n awn 2-10 mm. long; margins of the glumes thin and translu-
cent except toward the base where they are indurated, rounded
usually slightly separated exteriorly and quite widely interiorly.
A. Smithit Rydb. Spikelets not articulated (breaking away
with a rough base) and only slightly impressed. lum
about one-half as long as the i indistinctly nerved and
without distinct teeth on the back; margins thin on lower third
sein meee, 6: abruptly narrowed to a long stiff, acumin-
ate poi
A. tenerum Vasey. Spikelets not articulated and only
slightly impressed, Glumes about equalling the spikelets, acute,
strongly nerved, the nerves all rough with small teeth, margins
nearly straight.
+ AGROPYRON DASysTACHUM (Hook.) Scribn. A specimen
from Bottineau, July 7, 1920, is referred here (“form which has
been called subvillosum”’) by Hitchcock. Bowman, June, 30
- 1918, and Kenmare, July 15, 1913, are the same, and I think
all the specimens previously referred to A. molle (excepting
perhaps Bell 385) belong here.
+ Dryopteris THELYPTERIS (L.) A. Gray. Anselm, R. A.
Shunk
+ DRYOPTERIS SPINULOSA (Retz.) Kuntze. Another speci-
men of the same collection as the preceding seems to belong here,
as does also Walhalla, Bergman 2000.
Since the above was written I have received from Mr. F. P.
Metcalf a reprint of an article (Jour. Washington Acad. Sci. 10:
188-198. 1920) in which he reports eighteen additional species for
the state. Several of the records suggest the probability of an
error of indentification or of a difference in interpretation. In
reply to my inquiry regarding a few of them Dr. Standley
wrote that he has been able to find but one of the specimens,
Rumex Britannica L. (McLean County), and that that seemed
to be correctly determined.
* This is not actually the central nerve, the broader part of the glume being
exterior and only one or two nerves on the side next the rachis.
Three new species of Cuscuta from Mexico
T. G. YUNCKER
(WITH THREE TEXT FIGURES)
In studying collections of Cuscuta from Mexico three species
have been discovered which, it is believed, have not been
hitherto described. Descriptions of these new species, together
with camera lucida sketches, are given below. With the addi-'
tion of these the number of species of Cuscuta now known to
occur in Mexico is thirty-six.
Cuscuta dentatasquamata sp. nov.
s medium to slender. Flowers reddish, 3-4 mm. long,
on Soirée shorter than or mostly about equal to the ieee
in rather compact, cymose clusters; calyx deep, lobes exceeding
the corolla tube, triangular, acute, slightly overlapping at the
acute, upright or spreading, shorter than the tube and with
sli ghtly uneven margins; stamens shorter than the corolla lobes, |
filaments stoutish and about equal to the oval or orbicular
anthers; scales about reaching the stamens, dentate, with
Fic. CUSCUTA DENTATASQUAMATA Yuncker
A. Flower, X 5. B. Opened corolla, X 5. C. Individual scale, X Io.
_ D. Opened calyx, X 5. E. Capsule, X 5.
few processes, bridged at about the middle, not firmly adherent
to the corolla; styles slender, about equal to, or longer than, the
depressed- globose ovary. Capsule large, diepressed plobose,
thin, somewhat irregularly circumscissile, carrying the ea
corolla about it; seeds about 1.5 mm. long, globose, two to fou
in each capsule, hilum oblong, transverse.
This species falls in section EUGRAMMICA, subsection LEPTI-
LOBAE, near Cuscuta Choisiana Yuncker. It differs from that
107
108 YuUNKER: NEW SPECIES OF CUSCUTA
species, however, in the color of the flowers, and in the shape of
the scales and of the calyx and corolla lobes. It superficially
resembles Cuscuta partita Choisy, but differs from that species
in possessing shorter corolla lobes, dentate scales, shorter fila-
ments and larger flowers.
The only specimen examined was from Los Pinitos, Sonora,
Mexico (Hartman 119), the type, in the Gray Herbarium.
Cuscuta cozumeliensis sp. nov.
Stems rather coarse. Flowers 4-5 mm. long, on pedicels
about as long as, or shorter than, the flowers, in cymose clusters;
calyx shorter than the corolla tube, lobes ovate-orbicular, over-
lapping, somewhat fleshy toward the base, margins very uneven;
corolla campanulate, lobes much shorter than the tube, erect
or spreading, overlapping, obtuse or rarely slightly acutish,
margins uneven; stamens shorter than the lobes, filaments
stoutish, about equal to, or slightly longer than, the oval anthers;
scales not reaching the stamens, oblong, moderately fringed
Fic. 2. CuscuTa COZUMELIENSIS Yuncker
, Flower, X 5. B. Opened corolla, X 5. C. Opened calyx, X 5: D.
Individual scale, X 10. E. Ovary, X 5.
fragmentary and not ripe enough to show capsules. The ovary,
owever, gives indications that the capsule would be definitely
circumscissile and probably carry the withered corolla about it.
This species falls in section EuGRAMMICA, subsection OB-
TUSILOBAE, near Cuscuta macrocephala Schaffner, from which it
differs in the stamens not being sessile as they are in that species,
and in having smaller stigmas and longer processes on the scales.
YUNCKER: NEW SPECIES OF CUSCUTA 109
It might be confused with Cuscuta floribunda H. B. K., from
which it differs in having shorter corolla lobes and shorter scales;
with Cuscuta corymbosa: grandiflora Engelm., from which it
differs in the corolla not bulging between the filament attach-
ments and with broader scales; and with Cuscuta linctoria
wartius, from which it differs in its larger flowers and pro-
portionately shorter corolla lobes and shorter scales.
€ specimen examined was from Cozumel Island, Yucatan,
Mexico (Gaumer go), the type, in the Gray Herbarium.
Cuscuta durangana sp. nov.
Stems slender. Flowers membranous, about 2 mm. long, on
pedicels mostly as long as, or longer than, the flowers, in cymos
ager calyx about as long as the corolla or sarcey reaching
the s; lobes ovate, obtuse or, rarely, acutish, not over-
ania corolla campanulate; lobes ovate, obtuse, spreading,
later becoming reflexed, about as long as the tube; stamens short-
er than the lobes, the large,
oval, versatile anthers about
equal to the stoutish filaments;
scales about reaching the sta-
bridged low; styles slender and
longer than the globose ovary.
Capsule papas globose,
circumscissile, carry ing the
withered corolla about it; ap-
parently mostly two-seeded ;
seeds ovate, slightly rostrate,
about 1.5 mm. long, hilum
oblong, oblique.
This species falsl in section EUGRAMMICA, subsection OBTUSI-
LOBAE, near Cuscuta applanata Engelm. It differs from this
species, however, in having longer pedicels and a looser inflores-
cence and in not exhibiting keels on the calyx. It superficially
resembles Cuscuta umbellata H. B. K. The obtuse corolla and
calyx lobes easily distinguish this species, however.
The only specimen examined was from Durango, Mexico
(Endlich 268), the type, in the herbaruim of the Botanical
Institute at Dahlem (a small bit in the writer’s herbarium). .
DEPauw UNIVERSITY
Fic. 3. CuscuTA DURANGANA Yuncker
A. Flower, X 5 __B. Opened corolla,
x C. Opened calyx, X 5. D. Cap-
sule, X 5. E. Individual scale, X 10.
INDEX TO AMERICAN BOTANICAL LITERATURE
1908-1921
The aim of this ee is to incluge all current botanical literature written by.
Americans, published in *- >rica," “or Sina upon American material; the word
America being used in te
Reviews, and papers chat relate ee to foresty, agriculture, horti-
culture, manufactured products of vegetable origin, ¢ or laboratory methods
t included, and tt f bacteriology.
An occasional exception is made in favor of some paper appearing in an Ameri-
can periodical which i 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 ie attention of the editor to errors or omissions, their
apne will be appreciated.
s Index is reprinted monthly on cards, and furnished in this form to
ii at the rate of three cents for each card. Selections of cards are
not permitted; each ec must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Acosta, C. Coleccién de maderas cubanas. Revista Agr. Com.
y Trab. 3: 55. F 1920.
Acosta, C. El tararaco [Hippeastrum]. Revista Agr. Com. y
Trab. 3: 56. F 1920. [Illust.]
Acosta, C. Sobre el cayeput [Melaleuca leucadendron L.]
Revista Agr. Com. y Trab. 2: 535-537. f. 1, 2. N. 1919
Addis, J. M. Estudio de los platanos y guineos Spices Re-
vista Agr. Com. y Trab. 3: 418-429. f. 1-28. DI
Andrews, A. L. Notes on North American ae ‘VIL.
Bryologist 18: 1-6. 16 Ja I9gI5.
Andrews, A. L. Notes on North American jeans Vil.
Bryologist 20: 84-89. 13 O 1917.
Andrews, A. L. Tortula ee oe new species. Bryologist
23: 72-76. $1.5... 2D 16a
From North Carolina.
Arber, A. Leaves of certain amaryllids. Bot. Gaz. 72: 102-
105. f. 7-8. 15 Au Ig2!.
Arber, A. The leaf structure of the Iridaceae, considered in
relation to the phyllode theory. Ann. Bot. 35: 301-336.
I-66. Jl 1921. ;
II2 INDEX TO AMERICAN BOTANICAL LITERATURE
Ball,C.R. The relation of crop-plant botany to human welfare.
Am. Jour. Bot. 8: 323-338. Jl 1921.
Ballou, C. H. Los mejores mangos para Cuba. Revista Agr.
Com. y Trab. 1: 187-191. Ap 1918. [Illust.]
Bennett, A. Sparganium angustifolium Michx. Jour. Bot. 59:.
235, 236. Au 1921
Bergman, H. F. Intra-ovarial fruits in Carica Papaya. Bot.
Gaz. 72: 97-101. f. 7-6. 15 Au 1921.
Berry, E. W. A fossil sea bean from Venezuela. Am. Jour.
Sci. 50: 310-313. f. z. O 1920.
Berry, E. W. Age of the Dakota flora. Am. Jour. Sci. 50:
387-390. N 1920.
Berry, E.W. A palm nut from the Miocene of the Canal Zone.
Proc. (iS. Nat. Mus:99: 21, 22.°f. I-3.. 1921.
Berry. E.W. Paleobotany: a sketch of the origin and evolution
of floras. Smithsonian Report 1918: 289-407. pl. 1-6
+ f. I-42. 1921.
Berry, E. W. Tertiary fossil plants from Costa Rica. Proc.
U.S. Nat. Mus. 59: 169-185. pl. 22-27. 1921.
Includes 8 new species.
Blakeslee, A. F., Welch, D. S., & Cartledge, J. L. Technique
in contrasting mucors. Bot. Gaz. 72: 162-172. f. I-2.
S 1921:
Blasquez, A. La vegetacion las lluvias. Revista Agricola 6:
333-345. Je 1920.
Bois, D., & Geréme, J. La chilacayote du Mexique (courge de
Siai): Cucurbita ficifolia Bouché (C. melanosperma Al.
Braun). Bull. Mus. Hist. Natur. 26: 675-678. 1920.
Borden, A. D. A biological study of the red date-palm scale.
Phoenicococcus Marlatti. Jour. Agr. Research 21: 659-
667. pl. 127-130. 1 Au 1921.
Botto, A. La sweet tussac, Phalaris bulbosa Cav. Revista
Facult. Agron. y Veter. Univ. La Plata II. 12: 226-242. f.
iq. 1916; 13: 1-76. f. 740.2: 1918.
Britton, N.L. Stewardson Brown. Jour. N. Y. Bot. Gard. 22:
110-THR Je; i921.
Bruner, S. C. Enfermedades de la vid [Vitis] en Cuba. Re-
vista Agr. Com. y Trab. 1: 406-409. Au 1918. [Illust.]
INDEX TO AMERICAN BOTANICAL LITERATURE tt3
Bruner, S. C. La enfermedad del “mosaico’”’ 0 de ‘“Rayas
amarillas’’ de la cafia de azucar en Cuba. Revista Agr.
Com, y Trab. 2: 437-441. f. 7, 2. S 1919.
Bruner, S. C. La ‘‘Phomopsis’’ de la berenjena. Revista
Agr. Com. y Trab. 1: 468, 469. S 1918.
Bruner, S. C. La pudricién negra del cacao. Revista Agr.
Com. y Trab. 2: 630-636. D 1919. [Illust.]
Calvino, M. El zacate prodigio (Tripsacum latifolium, Hitch-
cock). Revista Agr. Com. y Trab. 3: 62-67. F 1920.
[Ilust.]
Calvino, M. Estudio sobre el cultivo de la soya [Soja max]
en Cuba. Revista Agr. Com. y Trab. 3: 124-131. My
1920. [Illust.] |
Calvino, M.. La lechuga [Lactuca sativa], su cultivo y apro-
vechamiento. Revista Agr. Com. y Trab. 1: 411-418.
Au 1918. [Illust.]
Calvino, M. La luz y la produccién vegetal. Revista Agr.
Com. y Trab. 3: 532-536. Ap i921. [Illust.]
Calvino, M. La jicama de agua (Pachyrhizus tuberosus)
Revista Agr. Com. y Trab. 2: 84-87. Mr 1g1g._ [Illust.]
Calvino, M. Nuevos variedades de cafia de azticar. Rev.
Agr. Com. y Trab. 3: 436-440. Ja 1921. [Illust.]
Carpenter, C. W. Morphological studies of the Pythiwm-like
fungi associated with root rot in Hawaii. Bull. Exp.
Sta. Hawaiian Sugar Pl. Assoc. Bot. 3: 59-65. pl. 16-23.
Au 1921.
Child, C. M. Le probleme de l’integration physiologique.
Scientia 30: 115-126. 1 Au 1921.
Clute, W. N. Note and comment. Am. Bot. 27: 116-113.
Au 1921.
Clute, W. N. The Victoria water lily. Am. Bot. 27: 81-86.
Au 1921.
Collins, J. L. Reversion in composites. Jour. Hered. 12: 129-
133. f. 26-19 10 Au 1921.
Conard, H. S. Fossil plants and classification. Am. Bot. 27:
95-101; Atl 192):
Coulter, J. M., & Land, W. J. G. A homosporous American
Lepidostrobus. Bot. Gaz. 72: 106-108. 15 Au 1921.
114 INDEX TO AMERICAN BOTANICAL LITERATURE
Cremata, M. Una excursién botanica a Isla de Pinos. Revista
Agr. com. y Trab. 3: 47-49. F 1920. [Illust.]
Cruz, F. B. El cultivo del tabacoen Cuba. Revista Agr. Com.
y Trab. 1: 172-183. Ap 1918. [Illust.]
Cummings, C. E. Mushrooms. Hobbies 2: 3-17. S 192!.
Dachnowski, A. P. Peat deposits and their evidence of climatic
changes. Bot. Gaz. 72: 57-89. f. 1-22. 15 Au I9g2!I.
Davidson, A. New species from southern California. Bull. S.
California Acad. Sci. 20: 49-53. Au 1g2t [Illust.]
Dusen, P., & Neger, F. W. Uber Xylopodien. Beih. Bot.
Centralbl. 38: 258-317. f. r-r10. 25 My i921.
Studies of Brazilian plants.
Duursma, G. D. Pilocereussen. Floralia 42: 376-378. 17
Je 1921. [Illust.]
Duursma, G. D. Rhipsalideae. Floralia 42: 378, 379. 17
Je 1921. oe
Eames, E. A. An unusual. form of Habenaria clavellata.
Rhodora 23: 126-127. pl. 131. 26 Au Ig2t.
Edgerton, C. W., & Tiebout, G. L. The mosaic disease of the
Irish potato and the use of certified potato seed. Louisiana
Bull. 181: 1-15. f. 7-3. Au 1921.
Elorduy, S. T. Selecciédn del tabaco. Rev. Agricola 5: 80I-
810. f. 1-9. Ap 1921; 6: 14-18. f. 10-72. My 1921.
Emerson, R.A The genetic relations of plant colors in maize.
Cornell Univ. Agr. Exp. Sta. Mem. 39: 1-156. pl. I-11.
Mr 1921.
Felippone, F. Contribution a la flore bryologique de 1’ Uruguay.
Fasc. 3: 40-45. 1917. [Illust.]
Includes new species in Didymodon (1), Leptodontium (1), Fabronia (1).
Feustel,H. Anatomie und Biologie der Gymnospermenblatter.
Beih. Bot. Centralbl. 38: 177-257. 25 My 1921.
Fortun,G.M. Notas sobre una excursion a ‘“‘El Retiro’’ [Cuba].
Revista Agr. Com. y Trab. 3: 410-413. D 1920. [Illust.]
Residence of Jose Blain.
Fortun, G. M., & Bruner, S. we Investigaciones sobre la en-
fermedad del “mosaico” o “‘rayas amarillas’ dela cajfia
de azucar. Revista Agr. Com. y Trab. 3: 441-445. Ja
1921. [Illust.]
Fraser,G. The dwarf trees of Vancouver Island. Gard. Chron.
70: 102. f. go. 20 Au 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 115
Fred, E.B. The fixation of atmospheric nitrogen by inoculated
soybeans. Soil. Sci. 11: 469-472. pl. 1-3. Je 1921.
Fred, E. B., Wright, W. H., & Frazier, W. C. Field tests on the
inoculation of canning peas. Soil Sci. 11: 479-491. pl.
1-3. Je 1921.
Frye, T. C. Notes on useful and harmful mosses. Bryologist
zo: Fi. -2 1 19a.
Gagnepain, F. Classification des Eugenia. Bull. Soc. Bot.
France 64: 94-103. 1917.
Gile, P. L., & Carrero, J.O. Assimilation of nitrogen, phospho-
rus and potassium by corn when nutrient salts are confined
to different roots. Jour. Agr. Research 21: 545-573. f. I.
te i) 19020.
Girola, C. D. Contribucién al curso de cultivas industriales.
Revista Facult. Agron. y Veter. Univ. La Plata II. 10:
83-114. 2078:
Girola, C. D. El cultivo de la yerba-mate. (Jlex paraguari-
ensis St. Hil.) Revista Facult. Agron. y Veter. Univ. La
Plata II. 11: 163-184. f. r-g. 1915; 12: 196-225. f. 1-6.
1916.
Girola, C.D. Monografia sobre el cafiamo (Cannabis sativa L.).
Revista Facult. Agron. y Veter. Univ. La Plata II. 10: 9-
68. f. 1-23. 1913.
Girola, C. D. Observaciones sobre el cultivo del henequen.
Revista Facult. Agron. y Veter. Univ. La Plata II. 11:
IOI-151. f. 7-19... 1615.
Grant, M. Saving the redwoods. Nat. Geog. Mag. 37: 519-
536. Je 1920.
Griffiths, D. The production of the Easter lily in northern
climates. U.S. Dept. Agr. Bull. 962: 1-31. f. r-14. 12 Au
1921.
Grosovich, J. M. Las principales malezas de los sembrados.
Revista Zootech. 8: 186-192. 15 My 1921.
Grove, W. B. Species placed by Saccardo in the genus Phoma.
Kew Bull. Misc. Inform. 1921: 136-157. I92I.
Includes 2 new American combinations in Phomopsis.
Hardy, M. E. Earth stars. Am. Bot. 27: 86, 87 Au 1921.
Harris, J. A. Leaf-tissue production and water content in a
mutant race of Phaseolus vulgaris. Bot. Gaz. 72: I51-
161. 15 3: FOat.
116 INDEX TO AMERICAN BOTANICAL LITERATURE
Harter, L. L., & Weimer, J. L. Studies in the physiology of
parasitism with special reference to the secretion of pectinase
by Rhizopus Tritici. Jour. Agr. Research 21: 609-625.
1 Au 1921
Heim, A. Chiceernitniede der Halbinsel Niederkalifornien.
Vegetationsbilder XIII. 3, 4: pl. 13-24 + descriptive text.
1916.
Hoerner, G. R. Germination of aeciospores, urediniospores
and teliospores of Puccinia coronata. Bot. Gaz. 72: 173-
15703 Th S 2041.
Hohenkerk, L. S._ Botanical identifications of British Guiana
trees and plants. Jour. Board Agr. British Guiana 11:
98-106, 178-185. 1918.
Hohenkerk, L. S. British Guiana timbers. Jour. Board Agr.
British Guiana 12: 152-187. Jl 1919.
Hollick, A. Loco weeds. Nat. Hist. 21: 85-91. F 1921.
H[osseus], C. C. Dr. Federico Kurtz [1854-1920]. Bol. Acad.
ac. Cienc. Cordoba (Argentina) 24: LIv-LIx. 1921.
Howe, M. A. Some plants from tropical sea gardens. Nat.
Hist. 20: 561-568. D 1920.
Hutchinson, J. The family Winteraceae. Kew Bull. Misc.
Inform. 1921: 185-191. 1921. [With map.]
Hutchinson, J. The genus Peeks le Kew Bull. Misc.
Inform. 1921: 201-205. f. 7.
Illick, J. S. The hard pines of the ee Am. For. 27:
487-496. Au 1921.
Illick, J. S. The pines of the South. Am. For. 27: 551-559;
574. S 1921. [Illust.]
Johnston, J. R. El platano y sus enfermedades. Revista Agr.
Com. y Trab. 1: 419-421. Au 1918. [Illust.]
Jones, L. H., & Shive, J. W. Effect of ammonium sulphate
upon plants in nutrient solutions supplied with ferric
phosphate and ferrous sulphate as sources of iron. Jour.
Agr. Research 21: 701-728. pl. 136+ f. 1-9. 15 Au 1921.
Jones, S. R. Preliminary report on the flora and fauna of the
eastern part of the Rosebud Reservation, now known as
Gregory County [South Dakota]. South Dakota Geol.
Surv. Bull. 4: 123-142. 1908.
Jouvenaz, J. H. Cereus. Floralia 42: 371, 372. 17 Je 1921.
[Illust.]
INDEX TO AMERICAN BOTANICAL LITERATURE 117
Jouvenaz, J. H. Echinocactus. Floralia 42: 372. 17 Je
1921. [Illust.]
Jouvenaz, J. H. LEchinocereus. Floralia 42: 372. 17 Je 1921.
[Illust.]
Kearney, T. H. Pollination of Pima cotton in relation to the
yield of seed and fiber. Jour. Hered. 12: 99-101. 10 Au
1921.
Kermode, F. A preliminary catalogue of the flora of Vancouver
and Queen Charlotte Islands. Pp. 1-89. Victoria, British
Columbia. 1921.
Kirjasoff, A. B. ee the beautiful. Nat. Geog. Mag. 37:
247-292. Mr 19
Kobel, F. Einige Stat na zu den Astragalus- und a
bewohnenden Uvomyces-Arten. Ann. Mycol. 19: 1-1
1921.
Korstian, C. F. Grazing practice on the natural forests and its
effect on natural conditions. Sci. Mo. 13: 275-281. f. 1-7.
Kradusel, R. Die Bedeutung der Anatomie lebender und
fossiler Hélzer fiir die Phylogenie der Koniferen. Naturw.
Wochenschr, 32: 305-311. f. I-g. 10 Je 1917.
Krausel, F. Welche Ergebnisse liefert die Untersuchung tertiarer
Pflanzenreste? Naturw. Wochenschr. 33: 209-213.
14 Apig18.
Krieger, L. C. C. Common mushrooms of the United States.
Nat. Geog. Mag. 37: 387-439. pl. 1-16 My 1920.
Lantes, A. El alamo [Populus]. Revista Agr. Com. y Trab.
2: 612, 613. D 1ig1g. ({Illust.]
Lingelsheim, A. Oleaceae-Oleoideae-Fraxineae. Pflanzenreich
IV. 243: 1-66, 115-125. f. 1-11 + map. 29 Je 1920.
Includes 2 new North American species of Fraxinus.
Livingston, B. E., & Shreve, F. The distribution of vegetation
in the United States, as related to climatic conditions.
Carnegie Inst. Washington Publ. 284: r-xvi + 1-590. pl.
7} +f. Ie. Feet.
Locy, W.A. The earliest printed illustrations of natural history.
Sci. Mo. 13: 238-258. f. z-ro. S 1921.
MacDougal, D. T. Growth in trees. Carnegie Inst. Wash-
ington Publ. 307: 3-41. f. 1-16. 1921.
118 INDEX TO AMERICAN BOTANICAL LITERATURE
MacDougal, D. T. Growth in trees. Proc. Am. Philos. Soc.
OO: Pri4. pF +f. 3-5. 921.
MacDougal, D. T. The action of bases and salts on biocolloids
and cell-masses. Proc. Am. Philos. Soc. 60: 15-30. I92I.
MacDougal, D. T., & Working, E.B. Another high- Bas apie
record for srt and endurance. Science II. 54: 152,
153. I9 Au 1921.
McMurray, N. Freak dogwood. Am. Bot. 27: 113. Au 1Ig2l.
MacTaggart, A. The influence of certain fertilizer salts on the
growth and nitrogen-content of some legumes. Soil Sci.
11: 435, 454. pl. z. Je Iga.
Meier, H. F. A. Effect of direct current on cells of root tip of
Canada field pea. Bot. Gaz. 72: 113-138. pl. 2-3 + f. 1-3.
15 S 1921.
Moodie, R. L. Bacteria in the American Permian. Science II.
54: 194-196. 2S 1921.
Mottier,D.M. On certain plastids, with special reference to the
protein bodies of Zea, Ricinus, and Conopholis. Ann. Bot.
35: 349-364. pl. 15. Jl gat.
[Nicholls, H. A. A.] Useful timbers of Dominica. In Bell, H.
H., Notes on Dominica and hints to intending settlers.
34-56. 1919. [Illust.]
Nichols, G.E. The vegetation of Connecticut. VII. The asso-
ciations of depositing areas along the seacoast. Bull.
Torrey Club 47: 511-548. f. 1-ro. 24 N 1920.
Oakley, R.A. Effect of the length of day on seedlings of alfalfa
varieties and the possibilities of utilizing this as a practical
means of identification. Jour. Agr. Research 21: 599-607.
pl. 111-121. 15 Jl 1921.
Overton, J.B. The mechanism of root pressure and its relation
to sap flow. Am. Jour. Bot. 8: 369-374. Jl 1921.
Pack, D. A. Chemistry of after-ripening, germination and seed-
ling development of juniper seeds. Bot. Gaz. 72: 139-
150.- 1§ S 102i. i
Pellett, F. C. American honey plants. pps. 1-297. f. I-I§2.
1920.
Pennington, L. H., & others. Investigations of Cronartium
ribicola in 1920. Phytopathology 11: 170-172. Ap 192I.
INDEX TO AMERICAN BOTANICAL LITERATURE 119
Petrak, F. Mykologische Notizen. II. Ann. Mycol. 19: 17-
128. 1921
Pittier, H. Acerca del genero Gyranthera Pittier. Contrib.
Fl. Venezuela 14-19. 1921.
Includes G. caribensis sp. nov. from Venezuela.
Pittier,H. Arbdes y arbustos nuevos de Venezuela. I. Contrib.
Fl. Venezuela 3-14. 1921.
New species in Inga (1), Raputia (1), Esenbeckia (1), Trichilia (1),
Stillingia (1), Gustavia (1), Mouriria (1), Oxythece (1), Pouteria (1), Sider-
‘ylum (1).
Plemper van Balen, B. A. Cephalocereus senilis, de ‘‘Grijsaard.”
Floralia 42: 371. 17 Je 1921. [Illust.]
Pondal, M. L. Fabricacién del extracto de quebracho. Bol.
Acad. Nac. Cienc. Cordoba (Argentina) 24: 293-324. f. I-
7s. 3 3021.
Popenoe, W. La mora gigante de Colombia. Revista Agricola
6: 503-507. S 1920.
Popenoe, W. The pejibaye, a neglected food-plant of tropical
America. Jour. Hered. 12: 154-166 f. 4-9. Ap 1921.
Povah, A. H. W. An attack of poplar canker following fire
injury. Phytopathology 11: 157-165. f. 1-3. Ap I92!.
Prouty, W. F. A more phenomenal shoot. Science II 54:
170. 26 Au 1921.
Shoot of Paulownia tomentosa.
Ramirez, R. El chahiuztle rojo del frijol. Revista Agricola
5: 830. Ap rg2r. [Illust.]
Ramirez, R. La cenicilla del tomate, Physalis dubescens L.
Revista Agricola 5: 830. Ap i921. [lIllust.]
Reagan, A. B. Notes on the flora and fauna of the Rosebud
Reservation west of Gregory County [South Dakota].
South Dakota Geol. Surv. Bull. 4: 143-167. 1908.
Reiche, K. Die physiologische Bedeutung des anatomischen
Baues der Crassulaceen. Flora 114: 249-255. f. 1, 2.
1921.
Reiche, K. Zur Kenntnis von Sechium edule Sev. Flora 114:
232-248: Ff, 3-9. 3084.
Reiche, K. Zur Kenntnis von Senecio praecox DC. Flora 114:
255-261. fi 3, 6, 1928.
Renson, C. El Tripsacum latifolium, Hitchc. Revista Agri-
cola 5: 351-368. Je 1919.
Richards, V. F. The sassafras. Am. Bot. 27: 105. Au I9g21.
120 INDEX TO AMERICAN BOTANICAL LITERATURE
Roberts, H. F. Relation of hardness and other factors to
protein content of wheat. Jour. Agr. Research 21: 507-
522. pl. roo. 15 Jl 1921.
Rock, J. F. The Okala berry of Hawaii. Jour. Hered. 12:
147-150. f. 7, 2. Ap 1921.
Roig, J. F., Cremata, M., & Bruner, S.C. Exploracién botanica
en la Cienaga de Zapata [Cuba]. Revista Agr. Com. &
Trab. 3: 213-221. Jl. 1920. [With map.]
Safford, W. E. Datura, an inviting genus for the study of
heredity. Jour. Hered. 12: 178-190. f. ro-16. Ap 1921.
Shufeldt, R. W. Common American mushrooms. Am. For.
27: 579-587. f. 1-13. S 1921.
Smith, E. F. Effect of crowngall inoculations on Bryophyllum.
Jour. Agr. Research 21: 593-597. pl. ror—-r110. 15 Jl 192I.
Snell, W. H. Chlamydospores of Fomes officinalis in nature.
Phytopathology 11: 173-175. f.z. Ap 1921.
Sprague, T. A. A revision of the genus Capraria. Kew Bull.
Misc. Inform. 1921: 205-212. I92I.
Sprague, [T. A.]. Begonia (Gireoudia) trigonoptera, Sprague
[Begoniaceae]. Kew Bull. Misc. Inform. 1921: 218, 219.
1921
New species from Guatemala,
S[prague], T. A. The generic name Wikstroemia. Kew Bull.
isc. Inform. 1921: 175, 176. 1921.
Sprague, T. A., & Riley, L.A. M. Notes on Raimanniaand allied
genera. Kew Bull. Misc. Inform. 1921: 198-201. 1921.
Includes nine new combinations.
Taylor, N. The flora above the timber line of Mount Marcy.
Am. Scien. & Hist. Preserv. Soc. Ann. Rep. 25: 317, 318-
1920.
Thatcher, L. E. A fungus disease suppressing expression of
awns in a wheat-spelt hybrid. Jour. Agr. Research 21:
699, 700. pl. 135. 15 Au Ig2t.
Theye, C. Nutricién nitrogenada de los vegetales. Revista
Facult. Letr. y Cienc. Univ. Habana 31: 145-151. Je 1921.
Thurston, H. W., & Orton, C. R. A Phytophthora parasitic
on peony. Science II. 54: 170, 171. 26 Au 1921.
Tisdale, W. H. Two Sclerotium diseases of rice. Jour. Agr-
Research 21: 649-657. pl. 122-126. 1 Au 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE I2I
Torres, L. G. Plantas forrajeras: una leguminosa forrajera
indigena (Desmodium leiocarpum G. Don—Meibomia leiocar-
pa). Revista Agricola 6: 84-86. Je 1921. [Illust.]
Trelease, W. A natural group of unusual black oaks. Proc.
Am. Philos. Soc. 60: 31-33. pl. 2-4. 10921.
Includes 3 new species from Mexico.
Uphof, J. C. T. Vegetationsbilder aus dem Staate Michigan.
Vegetationsbilder 13: pl. 43-48. 1921
Valdivia Montanez, M. A. El mani, su cultivo, productos y
usos. Revista Agr. Com. y Trab. 1: 570-579. N 1918.
[Illust.]
Arachis hypogea L.
Valencia, F. V. Mechanical tests of some commercial Philippine
timbers. Philipp. Jour. Sci. 18: 485-535. pl. 1 +f. r-r0.
My 1921.
Vendrell, E. La seleccién de las semillas. Revista Agr. Com.
y Trab. 1: 260, 261. My 10918.
Verbeek Wolthuys, J. J. Pereskieae en Opuntieae. Floralia
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Visher, S.S. The geography of South Dakota. South Dakota
Geol. & Nat. Hist. Surv. Bull. 8: 1-189. f. 1-52. Jl 1918.
Contains chapter on biogeography. [Corrected reprint]
Waby, J. Z. Some interesting species of palms. Jour. Board
Agr. British Guiana 11: 172, 173. O 1918; 12: 188—-195.
Jl 1919.
Weimer, J. L., & Harter, L. L. Respiration and carbohydrate
changes produced in sweet potatoes by Rhizopus Tritici.
Jour. Agr. Research 21: 627-635. I Au 1921.
Weimer, J. L., & Harter, L. L. Wound-cork formation in the
sweet potato. Jour. Agr. Research 21: 637-647. 1 Au
1921.
Weir, J.R. Note on Cenangium abietis (Pers.) Rehm on Pinus
ponderosa. Phytopathology 11: 166-170. f.z. Ap 1921.
Weir, J. R., & Hubert, E. E. Forest disease surveys. U. S.
Dept. Agr. Bull. 658: 1-23. f. 1-23. 12 Je 1918.
Weiss, F., & Harvey,R.B. Catalose hydrogen-ion concentration
and growth in the potato wart disease. Jour. Agr. Re-
search 21: 589-592. 15 Jl 1921.
I22 INDEX TO AMERICAN BOTANICAL LITERATURE
Welton, F. A. Longevity of seeds. Monthly Bull. Ohio Agr.
Exp. Sta. 6: 18-24. 1921. [Illust.]
[Corrected reprint. |
Wercklé, C. Die natiirlichen Wachstumsbedingungen der
epiphytischen Orchideen in Costa Rica. Gartenflora 70:
90-94. 15 My 1921.
Wester, J.P. The mango. Philipp. Bur. Agr. Bull. 18: 1-70.
pl. I-17.
Second revised edition.
Wherry, E. T. The reactions of the soils supporting the growth
of certain native orchids. Jour. Washington Acad. Sci.
8: 591-598. 4 N 1918.
Wherry, E. T. Soil reactions of Spiranthes cernua and its
relatives. Rhodora 23: 127-129. 26 An 1921.
White, D. Value of floral evidence in marine strata as indi-
cative of nearness of shores. Bull. Geol. Soc. Amer. 22:
221~227. 10 My I09gII.
Williams, R.S. Hyophila subcucullata sp. nov. Bryologist 24:
22. pl. 2. 25 [) 1921,
From Cuba.
Williams, R. S. Sematophyllum Smallii sp. nov. Bryologist
23: 76-78. pl. 6. 2 D 1920.
From Florida,
Wilson, W. J. Palaeobotany. Summ. Rep. Canada Geol.
Surv. Dept. Mines 1916: 300-302. 1917. ,
Wright, C. H. Aechmea chromatica, C. H. Wright [Bromeli-
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291. 21.
New species from tropical! America.
Yamanouchi, S. Life history of Corallina officinalis vat.
mediterranea, Bot. Gaz. 72: 90-96. 15 Au 1921.
Vol. 49 : No. 5
BULLETIN
OF THE
TORREY BOTANICAL CLUB
MAY, 1922
New British and American species of Lobomonas:
a study in morphogenesis of motile algae
Tracy E. HAZEN
(WITH PLATES 5 AND 6)
The genus Lobomonas was established in 1899 by Dangeard
(3) on a single species, L. Francei, found somewhat frequently
in the vicinity of Poitiers, France. This species had been figured
twenty-one years previously by Stein (7, pl. 73, f. 17, 18) asa
form of Chlamydomonas pulvisculus Ehrenb. Golenkin also
appears to have had this species in Russia, and to have confused
it with another genus, for one of his figures (5, f. 19), described
as a reduced form of Pteromonas alata (Cohn) Seligo, can hardly
be anything else than a young cell of L. Francei. Apparently
the species has not been studied or scarcely even reported other-. —
wise, except by Dangeard. In 1902 Chodat (2) transferred to
the newly founded genus his species Chlamydomonas stellata,
briefly described six years .before (1); he even appears to have
been doubtful of the distinctness of his form from Dangeard’s
type, though both species have been accepted by West (8, p. 172)
and Wille (9, p. 19), who have reproduced the original illustra-
tions.
The genus presents a cell organization almost precisely like
that of Chlamydomonas, probably its nearest relative, with the
exception that its outer wall is furnished with variable irregulari-
ties or protuberances, which in the type species are frequently
more developed on the posterior part of the cell, while in Z.
stellata the more uniformly triangular lobes are figured as cover-
ing the wall nearly to the region of insertion of the cilia. It
[The BuLLetin for April (49: 75-122. pl. 3, 4) was issued May 18, 1922.]
123
124 Hazen: NEW SPECIES OF LOBOMONAS
is probable that Lobomonas has escaped the notice of collectors
to some extent because of the minute size of the cells, but doubt-
less it is actually one of the rarest genera of the Chlamydomonas
group. The two new species now presented serve to emphasize
the unity and distinctness of the genus, and also provide certain
features of division and conjugation not known hitherto, which
indicate a fundamental parallelism with Chlamydomonas.
Lobomonas pentagonia sp. nov.
This species, the first of the genus to be reported in England,
I believe, was discovered in considerable abundance, together
with two species of Pteromonas, at Ham Common, Surrey, near
Kew, in 1920. The habitat was the border of a small shallow
pond at the west end of the common, a place frequented by cows
and horses, and therefore supplied with water containing a
considerable amount of nitrogenous organic matter in solution:
in fact the general conditions were very like those of the much
smaller pool where Lobomonas rostrata, to be described presently,
was found with another species of Pteromonas in New Jersey.
Like all other species of the genus, L. peniagonia is very
minute, but it is more constant in form than L. Francei or L.
rostrata. When seen in side view the cell appears rather pent-
agonal in outline (Fics. 1-4), but careful focussing shows that
the angles do not all lie in the same plane. Polar views often
show five to eight protuberances (Fic. 14), likewise not all at
the same level. One of the most symmetrical specimens, when
resting with its ciliated end turned upward, shows four anterior
protuberances and, alternating with them, four others at the
posterior end of the zoospore (FIG. 5); the number, however, is
variable. The protoplast, including the chromatophore, in
young individuals usually extends into and fills the protuber-
ances, and ends anteriorly in a somewhat obtuse beak, to which
the two cilia are attached (Fic. 1). In older specimens the pro-
toplast usually retracts from one or more of the wall protuberances
and leaves them empty: they do not appear like dense gelatinous
structures. The chromatophore is more or less hollowed out
in the common Chlamydomonas fashion but thickened in the
region where the pyrenoid lies in a somewhat lateral position.
The two alternately pulsating vacuoles lie in a plane nearly
perpendicular to that passing through the two extended cilia
(Fic. 5), as in most species of Chlamydomonas, so that in the
ordinary face view of the cell only one is usually seen. The
HaAzEN: NEW SPECIES OF LOBOMONAS 125
narrow rod-shaped red eye-spot lies in or just beneath the plasma
membrane slightly in front of the middle of the cell. Its position
with relation to the cilia is more like that which it occupies in
Brachiomonas than that in Chlamydomonas, as brought out
recently by the writer (6), though the dorsiventral differentiation
here is perhaps less constant and definite than in the two related
genera.
ASEXUAL REPRODUCTION. Dangeard (3) reports that his
attempts to cultivate ZL. Francet in a moist chamber did
not succeed, and that it was difficult to obtain the multipli-
cation of the organism. He shows that the cells come to rest
and generally become rounded in shape, then divide into four
or eight daughter cells which escape as zoospores. He gives
no figure to support Wille’s (8) surmise that division is longitu-
dinal. In our two new species I found that very generally after
motile cells were mounted in a hanging drop they would for the
most part come to rest in a few hours and proceed to divide,
though in many cases the daughter cells failed to become motile
or escape. In both species pyrenoids were not seen in any
dividing specimens until the daughter cells had begun to take
on the typical form, so that it would appear necessary to con-
clude that in this genus, as in Brachiomonas and in some species
of Chlamydomonas, the pyrenoid disappears before the first cleav-
age, and that one is formed de novo in each daughter cell.
In L. pentagonia the first division plane usually appears at
first sight to be transverse to the longitudinal axis of the cell,
but several cases were observed which lead to the conviction
that there is regularly a rotation of the protoplast during or
before the beginning of cleavage. Fics. 6 and 7 show two stages
of division where the protoplast has revolved to an oblique
position when the cleavage begins; probably the rotation was
delayed more than usual in this case; here the original contrac-
tile vacuoles persist after the division of the nucleus. Fig. 8
shows a case where what I am sure the original vacuoles persisted
until the end of cleavage into two daughter cells. In case of the
formation of four zoospores the second plane of cleavage is per-
pendicular to the first, i. e. it lies in a longitudinal axis of the
mother cell: it may lie in a single plane through the two halves
(Fic. 11), or the second division plane in one half cell may be
perpendicular to that in the other (Fic. 12).
126 HazEN: NEW SPECIES OF LOBOMONAS
SEXUAL REPRODUCTION apparently has not been observed hith-
erto in this genus. I considered it good fortune, therefore, to dis-
cover three or four cases of conjugation taking place almost
simultaneously in one hanging drop. The gametes have the
general form and character of the vegetative cells, but are much
smaller, provided with an excessively delicate cell wall, and the
cilia are longer than the cell body. In the first case seen, one
gamete was rather broad and pillow-shaped, with a distinctly
visible cell wall, while the other was narrower and probably more
nearly cylindrical, and furnished with a wall so delicate that it
was detected only in its subsequent behavior. The two gametes
became engaged by. their cilia, but not otherwise in contact,
and remained in this position for at least a half hour, with slight
dancing movement (FIG. 15); the cilia, most if not all of this
time, were trailed backward along the sides of the narrower
gamete. The first movement toward joining was in a sudden
break of the anterior papilla of the broad gamete, whereby a
broader papilla of colorless cytoplasm surged forward and pres-
ently plastered itself on the beak of the unchanged gamete (FIG.
16), the remainder of the protoplast of the broader gamete then
distinctly withdrawing from the posterior part of its wall (FIc.
17). Now for a period of about a half hour the remainder of
the protoplast of the broader gamete was gradually oozing out
of its wall and into the narrower gamete: even yet the narrower
gamete hardly showed a wall, but it seemed evident that one
must be present, since the posterior part of this gamete rigidly
retained its original form (Fics. 19, 20). After this point more
active ciliary movement carried the zygote beyond possibility
of observation.
A second case showed a similar figure of the narrow gamete
remaining rigid for as long a time as it could be followed. In
all probability this wall of the narrow gamete is finally thrown
off separately, permitting the rounding up of the plasmatic mass
to form a spherical zygospore. A third case showed the gametes
more nearly equal, and it was clear that the walls of both were
practically alike in character and not easily abandoned(FIG. 21 )y
so that fusion was long delayed.
There is in this species apparently little differentiation be-
tween the gametes; indeed one might be inclined to regard the
difference in size as merely accidental. Nevertheless from the
behavior of the cilia, that is, both pairs for the most of the time
stretching back alongside the narrower gamete, one may assume
HAZEN: NEW SPECIES OF LOBOMONAS 127
a certain degree of differentiation. The conjugation of this species
presents a close parallel to that described by Goroschankin for
Chlamydomonas reticulata (5, p. 126, pl. 3, f. 6-8). He states
that the gametes show little difference in size, and that they
sometimes simultaneously slip out of their walls; but more
frequently, after the beginning of conjugation, one of the gametes
first throws off its wall and takes on a globose form, then the
second does Jikewise, and then the two rounded masses go on to
complete fusion.
The subjoined Latin diagnosis ceaiants the chief character-
istics oe this species:
monas pentagonia sp. nov. L. cellulis vegetativis minu-
tis, seer erag S latere aspectata forma aliquanto pentagonia
sed angulis rotundatis (verrucis) haud omnibus in eodem plano
a vertice aspectata rotundata cum 5-8 verrucis, aliis anterioribus,
chromatophoro excavato, pyrenoidem unum sublateralem port-
ante, et parte eacavata nucleum lateralem includente;
stigmate bacilliformi paululum ante mediam cellulam sito;
vacuolis contractilibus es in rostello Spoplemacn positis.
Propagatio fit 2 aut 4 zsooporis intra ce ellulae matricalis
membranam ortis, divisione priore visa quasi ee eps
propter ibe ng rotationem sed vero longitudina
o fit gametis parvulis, vel subaequalibus a aliquan-
to paiva membrana tenuissima vestitis, inter se binatim
copulantibus.
Longit. cell. veg. 10-13 uw; lat. 9-10 wu. Longit. gametarum
- ca. 8 ys; lat. 4-5 uw; longit. ciliorum ca. I
Hab. in stagni margine. Ham Consnen, Surrey, England,
1-7 Aug. 1920
Lobomonas rostrata sp. nov.
This form, the first representative of the genus to be reported
in America, at first sight appeared very similar to L. Francei
Dangeard (3, p. 115), but careful study disclosed differences as
important ds those which distinguish most species of Chlamy-
domonas, so that I feel obliged to regard it as a new species. I
first found a few individuals in examining a collection of Gonium
pectorale and an undetermined species of Chlamydomonas, ob-
tained the last of September, 1919, from a rain-water pool o
a highway in the southern part of Englewood, New Jersey. Lat-
er it was interesting to discover that the species had been collect-
128 Hazen: NEW SPECIES OF LOBOMONAS
ed about a week earlier in a much deeper pool, about half a mile
distant from that just mentioned; this discovery was due to the
fact that a fine colony of the Lobomonas developed on an agar
plate containing a sample of my first collection of Pteromonas
from this pool. The new species continued to appear sporadically
in later gatherings from the same pool, the last being made
November 12; it was also collected in October, together with
Chlamydomonas metastigma Stein, from another rain-water pool
in a wheel-rut, not far from the one first mentioned, though
separated from it by railroad tracks bordered by a deep ditch
on either side. The first mentioned wheel-rut yielded a few
individuals in the following season. A few specimens of this
species were also discovered during the past summer in a col-
lection from a similar wheel-rut in northern Vermont: here it
was accompanied by Gonium pectorale, Pandorina, Chlorogonium,
Chlamydomonas, and a very interesting new form of Polyble-
pharides, to be described in a forthcoming paper. The Lobomo-
nas never appeared to be abundant like its associates in the
same pools; usually not more than a dozen or two specimens
turned up in one hanging drop mount.
The vegetative cells or zoospores of this species most common-
ly have a somewhat obpyriform shape (Fic. 22-27) though they
are sometimes almost ellipsoid. In younger individuals the
cell wall is so delicate and close-lying as to be indistinguishable
for the most part, but in older cells it is well developed (FIGs.
28, 29); it is generally produced into a variable number of lobes, of
which from five to seven or sometimes as many as ten appear
in a face view; that these lobes are developed on al) sides of the
cell is clearly shown in a polar view (FIG. 30). At the anterior,
usually broader end, the wall is extended into a truncate, wedge-
shaped beak, or possibly more typically this takes the form of
a more or less double papilla (Fic. 33); on account of the minute
size of the organism it is often most difficult to see clearly the
exact structure of this protuberance, which is one of the most
characteristic features distinguishing this species from the two
hitherto described in Europe. Sometimes the wall appears to
be uniformly thin, sometimes thickened at the end of the lobes,
and sometimes considerably thickened throughout. The proto-
plast, indistinguishable outwardly from the bright green chro-
matophore, fills the lobes in young individuals; in older cells it
retracts more or less, so as to leave some or all of the lobes color-
less. In such cases it is not easy to determine whether the
HAZEN: NEW SPECIES OF LOBOMONAS 129
lobe is a dense, gelatinous structure, or simply membranous and
separated by a space from the protoplast. In certain cases the
latter interpretation is clearly indicated, for the tip of the lobe
is manifestly of thickened grayish wall substance, with a clear
space inside (FIG. 37). The cilia are attached at a single point
to the anterior end of the protoplast, which is usually obtuse,
though it may have a slight beak; they immediately diverge
at a wide angle to pass separately through the papilla of the wall,
and are often seen in quiescent individuals stretching out stiffly
in the form of av; their length is as variable as the cell outline,
often being less than the cell length, but perhaps more character-
istically distinctly greater than the cell length. At the base of
the cilia are the alternately pulsating vacuoles, lying regularly
in such a position that a line passing through the two is perpen-
dicular to the plane in which the quiescent cilia lie, so that only
one of the vacuoles is seen when both cilia are equally clear, but
both may appear at the same level hci one of ne cilia i < bebind
the other (FIG. 29).
in the deeply hollowed out chromatophore (Fr IGS. 24, 26), contrast-
ing sharply with the axial pyrenoid in a massive chromatophore
described and figured by Dangeard (3) in Lobomonas Francei.
The small rod-shaped red eye-spot lies in front of the middle
of the cell, but apparently not in a constant position with ref-
erence to planes passing through the cilia and contractile vacu-
oles, as is the rule in Pteromonas and in many species of Chlamy-
domonas.
REPRODUCTION. The earliest stages of division found pre-
sented the appearance of a cleavage transverse to the longi-
tudinal axis of the cell. More careful consideration, however,
here, as in L. pentagonia, indicates that an early rotation of the
protoplast has eluded observation, for in Fic. 36 two contractile
vacuoles, lying in what appears to be the original colorless an-
terior cytoplasm, now appear on the side of the cell. Even in
such a case as that shown in Fic. 37, there is a colorless central
region which can only be explained on the supposition that the
anterior end of the protoplast had revolved ninety degrees from
its original position, here not clockwise as in Fic. 36, but in a
vertical plane with reference to the observer. The mother cell
retains very much of its original form throughout the process of
division, instead of rounding up, which Dangeard describes as
being the general rule for L. Francei. Here also there may be
four or eight daughter cells formed, and they regularly show the
130 Hazen: NEW SPECIES OF LOBOMONAS
typical obpyriform and lobed shape, and sometimes even show
the protoplast somewhat separated from the new cell wall, before
escaping from the mother cell. The escape appears to be accom-
plished by a gradual softening and disintegration of the wall of the
mother cell, rather than by rupture at a single point (FIG. 40).
The eight daughter cells of this figure are so small as to occasion
the surmise that they might be gametes, but conjugation was
not seen in this species.
The chief characteristics distinguishing this species from
L. Francei Dang. are: the general obpyriform shape, the well
developed anterior beak or papilla, the lateral pyrenoid, and the
persistence of the form of the mother cell as division. The
description may be summarized as follows
Lobomonas rostrata sp. nov. L. caltulis vepeintves plus
minusve obpiriformibus, rarius Secor. membran cellulae
cta, per quo
longitudine corpus scilie sia sitin vel longiora, vel bre-
viora: proto plasto primum membranae verrucas complente,
deinde plus minus contracto et ellipsoideo: Boia top nak valde
excavato, pyrenoidem unum lateralem portante: stigmate bacil-
nunc
contractilibus binis juxta papillam anteriorem suppositis.
esopoentto | fit protoplasto cellulae vegetativae jam immobilis
diviso in 4 a zoosporas, quae formam typicam priusquam
evaderunt ex seme matricalis membrana adipiscuntur.
Copulatio haud hier
ongit, cell. veg. Sra i: lat. 4-8 u.; long. ciliorum Ca. 5-14 u.
Hab. in aqua pluviali quae colligitur in viis terrenis, et in
lacuna quadem lutulenta in pascuo pecuario. Englewood, New
Jersey, Sept—Nov. 1919, Sept. 1920:Shelburne, Vermont, 3
Aug. 1921
It has been already pointed out elsewhere (6) that Lobomonas
is to be regarded as a special offshoot from Chlamydomonas,
not leading to any higher group so far as we know at present.
It might be thought simple to derive the genus directly from
the Polyblepharidaceae, even from the genus Dunaliella, which
clearly appears to be the immediate ancestor of Chlamydomonas,
since it has all the features of cell organization of the latter
genus except for the lack of a firm cell wall. When, however,
it is recalled that the gametes of Lobomonas, reported above for
the first time, possess cell walls, it will be more natural to look for
HAZEN: NEW SPECIES OF LOBOMONAS 131
its ancestry among those species of Chlamydomonas which posses
walled gametes, and are therefore ranked as the primitive mem-
bers of the genus, since their vegetative cells and gametes
differ (visibly) only in point of size. Our two new species of
Lobomonas, moreover, also resemble a number of the relatively
primitive members of Chlamydomonas in their method of cell
division, namely through a cleavage which is fundamentally
longitudinal but early shifts to a transverse position. The
question then arises, what influences led to the divergence of
cell form, which is practically the sole basis of separation between
the species forming the small genus Lobomonas and the much
larger number comprised in Chlamydomonas.
A CONSIDERATION OF MORPHOGENESIS IN PRIMITIVE ALGAE
This whole problem of the origin and inheritance of cell
form in primitive organisms is one of very great interest which
has received comparatively little attention. Perhaps the most
extended discussion of the question has been furnished by D’Arcy
Thompson (22), who regards surface tension as the paramount
factor in the determination of cell form. Though at one point
he admits that ‘the physical cause of the localized inequalities
of surface tension remains unknown’, and at other times hints
that an internal chemical heterogeneity may have some influence
in connection with such differences in surface tension, never-
theless again and again he reiterates his main thesis in regard
to unicellular organisms, ‘that not only their general config-
uration but also their departures from symmetry may be correlated
with the molecular forces manifested in their fluid or semi-fluid
surfaces’. This explanation seems to us entirely inadequate
and not in harmony with the general weight of evidence. For
the particular organisms considered here and in the previous
paper on Brachiomonas (6) we can offer little direct evidence;
but there are certain facts which suggest that the conception of
the non-homogeneity of the protoplasmic structure of the cell,
as developed by Rhumbler (19) and Harper (12) supplies a
much more workable hypothesis than the idea of mere surface
tension forces.
Passing over Rhumbler’s work on protozoa, the most
thorough treatment of morphogenesis within a small group
of primitive plants is found in the studies of Harper (11, 12)
on Pediastrum. He believes that the general four-lobed form
132 Hazen: NEW SPECIES OF LOBOMONAS
of the cells familiar in most species of Pediastrum may well
have arisen in evolution as a consequence of the pressure and
contact relations of the young cells in the sixteen-celled colony,
regarding them merely as surface tension globules: nevertheless
he has shown repeatedly that this four-lobed form does not de-
pend in ontogeny upon the forces which may have been respon-
sible for it originally, but that it is inherited and may reach
full expression when there is the least possible contact with other
cells of the colony. Repeatedly Harper calls attention to the
view that though surface tension is commonly acting as a
morphogenetic factor, nevertheless ‘it is the inherited anomo-
genous consistency of the cells which is of most significance in
determining their form.’ In strictly unicellular organisms like
Lobomonas and Brachiomonas, there is an absence of the inter-
action of contact and pressure stimuli which are important
influences in the variation of the Pediastrum colony; never-
theless in the fundamental organization of the cell the factors
must be parallel in the main. In both cases the lobed form may
be regarded as adaptive for the general metabolism of the cell.
_In Pediastrum the development of spines is usually Jooked upon
as a case of primitive differentiation for protection, and in
Lobomonas and Brachiomonas the lobes might easily be conceived
of as subserving a similar function: in point of fact, I have ob-
served that when these forms are found in the same pools with
smooth-walled ovoid Chlamydomonas cells, it is the latter that
are devoured by protozoa, while they rarely or never prey upon
the lobed forms, even though the latter are smaller. However,
when two species of Chlamydomonas are present together, some-
times one is rapidly consumed by protozoa while the other is
ignored. The anti-selectionist, moreover, might fairly inquire
why it is that these genera of bizarre form have produced very
few species, in comparison with the extraordinarily successful
genus Chlamydomonas, which has attained well nigh three score
species, so far as taxonomy goes, in recent years.
Wille (9, p. 19) ascribed to the zoospores of Lobomonas the
characteristic of ‘deutliche Metabolie.’ I cannot find that
Dangeard uses this term in connection with this genus; but as
defined by him elsewhere* as amoeboid movement, or used
*On désigne sous le nom de métabolie une sorte de mouvement par con-
traction du corps particulier 4 quelques Euglénes, Amibes, Monades, etc.
C’est ce que nous appelons mouvement amiboide. [Dangeard: aoe
sur les algues inférieures.. Ann. Sci. Nat. Bot. VII. 7:144. 1888.]
HaAzEN: NEW SPECIES OF LOBOMONAS 133
more exactly by other recent writers (especially in connection
with the Polyblepharidaceae) in the sense of euglenoid or amoe-
boid change of form—and this was the usage of Perty*, by whom
the word appears to have been coined in 1852—the term is mis-
leading when applied to Lobomonas: or at least it can be used only
in a very restricted sense in connection with this genus. The
young cells of Lobomonas of course do undergo a certain change
of outline during the formative process within the wall of the
mother cell: ordinarily essentially the mature form is acquired
and fixed before their escape, and I have found no evidence that
it is appreciably altered afterward during activity, though, in
appearance only, the mature cell may be distinctly reminiscent
of Amoeba. Nevertheless, this idea of amoeboid change of
form is most suggestive in a discussion of the formative period
of such genera as we have under consideration, and for this re-
stricted period I believe we are entirely justified in drawing a
parallel with the results of certain recent researches on the
production of pseudopodia.
A brief survey of this work may be useful in this connection.
McClendon (16) has attempted to explain amoeboid movement
as due to local increase in permeability, the Amoeba simply re-
ceding from the side on which the permeability has been in-
creased. Its author himself admits the difficulty of explaining
positive reactions by this theory, and it certainly does not offer
any sufficient explanation for the assumption of the character-
istic cell-form shown by our developing chlamydomonads. The
closely related hypothesis that production of pseudopodia may
be accounted for by local variation in surface tension has been
widely invoked. The re-statement of this theory lately made
by Thompson (22) may here be passed over, even as this stim-
ulating writer has all but ignored the evidence which has been
accumulating against the surface tension explanation. Both
the permeability idea and the surface tension explanation as
ordinarily employed are objectionable, in that they depend too
tabolie. Hirunter verstehe ich die durch innere Vorgdénge bedingte
wechselnde Gestaltanderung. Seit langem bei den Amiben bekannt ist sie
bei den Infusorien so viel als nicht beachtet worden. [Perty: Zur Kenntniss
kleinster Lebensformen, p. 127. Berne. 1852.
Since this term Metabolie seems to be unfamiliar to American botanists
because of its very restricted usage, and since it has even been r endered as
equivalent to metabolism in a recent inane dictionary for chemists,
it appears to be worth while to call is original definition.
134 Hazen: NEW SPECIES OF LOBOMONAS
much upon external environmental influences to account for
the requisite local variability, though this element is not neces-
sarily involved in either theory.
The beginning of the more recent undermining of the surface
tension explanation of form change is found in the work of
Jennings (14), who from painstaking direct observation reported
that the currents in a moving Amoeba as a whole ‘are not similar
to those of a drop of inorganic fluid that is moving or elongating
as a result of a local increase or decrease in surface tension’, and
in particular that ‘the movements of material in a forming
pseudopodium are not like those in a projection which is pro-
duced in a drop of inorganic fluid as a result of a local decrease in
surface tension’. From these observations Jennings was forced
to the conclusion ‘that changes in the surface tension of the
body are not the primary factors in the movements and reac-
tions of Amoeba’. Similar conclusions from direct observation
were announced two years Jater by Dellinger (10), whose clever
photographs of Amoeba viewed from the side reveal the locomo-
tion of the organism as a sort of ‘walking,’ rather than a flowing of
a fluid substance upon the substratum. These negatory con-
clusions have been reinforced and extended by a series of recent
researches in the field of microdissection (15, 21, 13). From
these there is general agreement that the structure of Amoeba
(and probably of many other rhizopods and cells of primitive
organization) is not of the nature of a simple fluid mass governed
chiefly by surface tension forces, but rather that it is a highly
non-homogeneous system, consisting of comparatively fluid
endoplasm surrounded by an ectoplasm which has often the
character of a semi-rigid gel, possessing a considerable elasticity,
though the two regions may very probably grade into one an-
sage imperceptibly.
Finally, the experiments of Hyman (13) reveal in each
pseudopodium of Amoeba a gradient in susceptibility to potas-
sium cyanide, the susceptibility being greatest at the distal end
and decreasing proximally. This susceptibility gradient is re-
garded as being a metabolic gradient which arises before the
pseudopodium appears, ‘and hence the metabolic change which
produces increased susceptibility is the primary cause of pseud-
opodium formation.’ Liquefaction or solation is regarded as
the cause of the extension of a pseudopodium, and coagulation
or gelation as the cause of its retraction: the liquefaction is be-
lieved to be brought about by the metabolic change just re-
HAZEN: NEW SPECIES OF LOBOMONAS 135
ferred to. This theory that amoeboid movemnt is due to alter-
ations of the colloidal state is only in the nature of confirm-
ation and extension of the view advanced more than forty
years ago by Montgomery (17, 18) that protoplasmic movement
of amoeboid organisms consists in ‘an alternate expansion and
contraction of organic substance’: Montgomery even anticipated
Hyman in expressing the idea that the liquefaction which occas-
ions pseudopodium advancement is itself due to metabolic changes,
while he anticipated Rhumbler (19) in the idea of a non-homo-
geneity of the primitive protoplasmic mass which permits various
functions to be carried on in different regions at the same
time.
This somewhat lengthy excursus (which is yet only the
briefest possible summary of a voluminous literature) has been
introduced here only to bring before botanists, to whom the field
may be unfamiliar, facts which it is believed may be directly
applied in the case of the chlamydomonads we are considering.
The form development of Lobomonas, Brachiomonas, and Ptero-
monas must be essentially amoeboid for-a brief period during
the organization of the daughter cells, and we are justified
in assuming that their lobes and excrescences are the expression
of the same non-homogeneous organization of the protoplast
as is characteristic of Amoeba.
For this view, furthermore, we may draw an additional
parallel from the results of microdissection. In the developing
oogonium of Fucus, Seifriz reports (21) that the viscosity of the
protoplasm changes from liquid consistency in the young uni-
nucleate stage to slightly viscous consistency when the division
into eight eggs is just complete, and to decidedly viscous consis-
tency (just under the viscosity of glycerine) in the mature nor-
mally discharged egg: that is, in Seifriz’s scale of ten grades of
viscosity—the first attempt on the part of microdissectionists
at standardization in this matter—the variation is from grade
3 to grade 6. Yet further, from the behavior of disintegrating
eggs of Fucus, Seifriz (20) finds that the process may be localized
in such a manner as to indicate ‘a gross structure of the egg
plasm, i. ¢., the protoplasm is composed of many centers of activity
in which different chemical reactions take place.’ [have recently
found a condition almost precisely similar in the case of a newly
discovered polyblepharid genus which it is hoped may soon be
published. The cells of this species, though surrounded only
by an exceedingly delicate protoplasmic membrane, are never-
136 Hazen: NEW SPECIES OF LOBOMONAS
theless able to maintain for extended periods an elaborately
eight-ridged or winged prismatic form, and in disintegration
often break only at one or two points, thus permitting the greater
body of the protoplasm to remain practically intact, while only
small streams ooze out. From these observations we are justified
in concluding that such cells as those of Brachiomonas and Lo-
bomonas, could we apply methods of microdissection, would be
found to have protoplasm of a comparatively fluid consistency
during division, but that local increase in viscosity gradually
permits the fixation of the characteristic lobed form of the cell.
When it is recalled that in these two genera the pyrenoids regular-
ly disappear in preparation for cell division, and are reorganized
with the maturing of the daughter cells, it will readily be seen
that in this reorganization, combined with the ordinary processes
involved in division of cells inheriting differentiation of polarity
in at least three axes or planes, there is abundant room for the
play of such metabolic changes as might well account for con-
siderable differences in viscosity in different parts of the develop-
ing daughter protoplasts. It is this non-homogeneity of struct-
ure, involving very likely chemical as well as physical differences,
which may be regarded as the dominant factor in the determina-
tion of form in such organisms. From this standpoint, the prob-
lem as to how this characteristic form may be transmitted in
heredity does not seem so insoluble as it does on the assumption
of form determining unit factors in a germ plasm.
One further point may be emphasized. It is stated by
Thompson (22) that when ‘owing to some heterogeneity of the
substance’ the operation of uniform surface tension forces is
modified so as to result in the production of the ellipsoid cell
characteristic of yeast, for example, ‘this or any other asym-
metrical form, once acquired, may be retained by virtue of the
solidification and consequent rigidity of the membranous wall
of the cell.’ In the case of the organisms with which we are
here concerned at least, I am confident, the development of the
cell wall is not a necessary condition of the maintenance of
specific form; for the new polyblepharid mentioned above, and
others of the same group, as well as the gametes of Brachiomonas,
are able to preserve essentially the same form for long periods,
in spite of the fact that they are clothed only with a protoplasmic
membrane of such excessive thinness that it is practically unde-
monstrable. It is then, owing to relative viscosity in their
HAZEN: NEW SPECIES OF LOBOMONAS 137
protoplasmic substance that these motile organisms are able —
to attain and maintain their specific form.
In both Brachiomonas and Lobomonas we have noted that
not infrequently mature cells exhibit a more or less rounded form
of the protoplast inside of the cell wall, while the latter main-
tains the typical lobed form as fixed in the formative period.
Such conditions are not improbably produced by shrinkage due
to loss of a certan amount of water from the protoplast, accom-
panied by a tendency to decrease in viscosity of the outer layer,
in which case, surface tension would tend to bring about a more
rounded form. In other words, the maturing protoplast might
be said to show a tendency to revert to what might be considered
the ancestral form in preparation for reproduction.
It is a pleasure to acknowledge my debt to Professor R. A.
Harper for numerous stimulating discussions of problems con-
nected with morphogensis, and for reading this paper in manu-
script.
BARNARD COLLEG
COLUMBIA UNIVERSITY
138 HazEN: NEW SPECIES OF LOBOMONAS
LITERATURE CITED
A. In connection with Lobomonas
.Chodat, R. Matériaux pour servir 4 l’histoire des Protococ-
coidées V. Bull. Herb. Boiss. 4: 273-280. 1896.
Algues vertes de la Suisse. Berne. 1902.
.Dangeard, P. A. Mémoire sur les Chlamydomonadinées ou
histoire d’une cellule. Le Botaniste 6: 65-292. f. I-20.
1899.
Golenkin, M. Pteromonas alata Cohn. Bull. Soc. Imp.
Nat. Moscou 5: 417-430. pl. 12. 1891.
. Goroschankin, I. N. Beitrage zur Kenntniss der Morpho-
logie und Systematik der Chlamydomonaden. Bull.
Soc. Imp. Nat. Moscou 5: 101-142. pl. 1-3. 1891.
. Hazen, T. E. The phylogeny of the genus Brachiomonas.
Bull. Torrey Club 49: 75-92. pl. 3,4. 1922.
. Stein, F. Der Organismus der Infusionsthiere 3'. Leipzig.
1878.
_
wo d-
tn >
an
sy
8. West, G. S. Cambridge Botanical esuthiOoks. Algae 1.
Cambridge. 1916.
. Wille, N. Chlorophyceae. Engler & Prantl, Nat. Pflanz-
enfam 12, Nachtr. IgII.
‘Oo
B. On the problem of morphogenesis
. Dellinger, O. P. Locomotion of Amoebae and allied forms.
Jour. Exp. Zoél. 3: 337-358. pl. 1, 2+ f. 1-27. 1906.
. Harper, R. A. The evolution of cell types and contact and
pressure responses in Pediastrum. Mem. Torrey Club
17: 210-240. f. 1-27. 1918.
Organization, reproduction, and inheritance in Pedia-
strum. Proc. Am. Phil. Soc. 57: (375--439- bh. 5, OF 7.
I-35. 1918.
13. Hyman, L. H. Metabolic gradients in Amoeba and their
relation to the mechanism of amoeboid movement.
Jour. Exp. Zéol. 24: 55-99. f. I-14. 1917.
14. Jennings, H. S: Contributions to the study of the behavior
of lower organisms. The movements and reactions of
Amoeba. Carnegie Inst. Washington Publ. 16: 1317
234. f. 30-78... 1904.
15. Kite, G. L. Studies on the physical properties of proto-
plasm. I. The physical properties of protoplasm of
ie)
—
ol
Hazen: NEW SPECIES OF LOBOMONAS 139
certain animal and plant cells. Am. Jour. Physiol. 32:
146-164. I913
- McClendon, J. F. Ameboid motion and tropisms. Physi-
cal chemistry of vital phenomena, pp. 148-156. Prince-
TON. 1917:
. Montogomery, E. The elementary functions and _ the
primitive organization of protoplasm. St. Thom. Hosp.
Rep. 9: 75-100. 1879.
a
On
Lea!
“J
18. Zur Lehre von der Muskalooneeactiar II. Die amé6-
boide Bewegung. Arch. Gesam. Physiol. 25: 499-509.
pl. g. 1881
19. Rhumbler, L. Der Aggregatzustand und die physikalischen
Besonderheiten des lebenden Zellinhaltes. Zeits. Allg.
Physiol. 1: 279-388. 1902.
20. Seifriz, W. Observations on the structure of protoplasm
by aid of microdissection. Biol. Bull. 34: 307-324. f. r-3.
1918.
Viscosity values of protoplasm as determined by micro-
dissection. Bot. Gaz. 70: 360-386. 1920.
. Thompson, D’Arcy W. On growth and form. Cambridge.
1917.
iS)
Ny
Explanation of plates 5 and 6
Figures drawn with camera lucida from living material kept in hanging
drops (VanTieghem cells of standard height of 5 mm.): Leitz compens. oc.
I2 was used in combination with oil immers. obj. I-12 inch, or Spencer 2 mm.
The drawings have been reduced one half in reproduction, making the present
magnification approximately 1150 diameters for Fics. 1-21 and 41, and 1375
diameters for F1Gs. 22-40.
PLATE 5
LoBOMONAS PENTAGONIA Hazen
Fics. 1-5. Typical vegetative cells or zoospores: in Fics. 1-3 the
deca lies on the under side of the ce
G.5. Anterior 5 view: four unterige lobes emtpy, chromatophore
filling fae posterior lo
rotoplast yoreling clockwise in preparation for division; 7 P. M.
e.
si IG. 8. Division into two zoospores completed at 6:30 P. M.
.9. Another individual: pyrenoid and cilia beginning to appear in
‘ienhene cells; nucleus in most advanced lobe: 1
Fic. 10. Pentagonal form attained by Uaiighter cells.
Fic. 11. Four sich cells in one plane.
Fic. 12. Two daughter cells with axes perpendicular to those of the
other pair:
140 HazeEN: NEW SPECIES OF LOBOMONAS
: An unusually simple syst cilia contracted into a ball.
Fic. 14. Cell in posterior polar v
Fics. 15-20. Stages in coninsation of a pair of slightly unequal gametes:
Fic. 15 at 1:45 P. M.; i IG. 16 at 2:15; Fic. 17 at 2:16; Fic. 18 at 2:25; FIG.
19 at 2:40; Fig. 2 2:45.
FIG. 21. . Noha by conjugation of equal walled gametes.
PLATE 6
LOBOMONAS ROSTRATA Hazen
Fics. 22-27. Typical young vegetative cells.
Fics. 28, 29. Mature cells of less common form, but with well developed
wall, : IG. 29 showing edge view beak.
G. 30. Anterior polar v
cane 31, 32. Cells evel Si itmnce of form some time after loss of
motility
Fic. 33. Relatively large mature cell, pyrenoid and stigma underneath.
Fic. 34. The same ry 24 hours later: four daughter cells completely
organized except for cilia, 1
1G. 35. One of the oa daughter cells, one day later, not very well
formed, probably because of unfavorable conditions.
Beginning of cleavage, after clockwise rotation of protoplast;
9:30 P. M.
Fic. 37. Similar division, after rotation of protoplast in vertical plane;
10:30 P. M
Fic. 38 cleavage just completed; 10:30 P. M.
Secon
Fic. 39. Eight deenysins cells (perhaps gametes) 6 P.
Fic. 40. The same at 9:30 P. M.: daughter cells held gelatinized wall
of mother cell: cell ¢ Saud out sluggishly at 8:30; at 10 Pp. M. cell b backed
out presently rested on end, cilia downward as at 5}.
G. 41. Mature cell from Shelburne, Vermont, August 3, 1921. All other
ae from New Jersey material, in 1919.
nes
LOBOMONAS PENTAGONIA Hazex
LUB
Buti. Torrey
Buti. Torrey Crus
The development of the flower and embryogeny of
Martynia louisiana
FLORA ANDERSON.
(WITH PLATES 7 AND 8 AND TWENTY-FIVE TEXT FIGURES)
INTRODUCTION
The most common and widely distributed representative
of the Martyniaceae in temperate North America is Martynia -
lowisiana Mill. Although the literature on the family is chiefly
taxonomic, the descriptions are meagre and confusing. All the
species have, at some time, been placed in the genus Martynia.
The ten or more known species comprise three genera, native
to the warmer parts of the western hemisphere.
Efforts were made to secure seeds and to grow plants of the
different species for comparison, in order, if possible, to work
out the taxonomy of the family. Seeds were obtained from
seed-houses both in the United States and in England and also
from the Bureau of Plant Industry, the University of Texas
and the Mississippi Agricultural College.
From seeds secured through the Bureau of Plant Industry,
quite a number of plants of Martynia louisiana were grown—
all the plants producing viable seeds—and one specimen of
Martynia lutea Lindl. which, although maturing aire
normal fruit from selfed flowers, produced no viable s
Reciprocal crosses of Martynia lutea and Martynia plan
and of Martynia lutea and Martynia fragrans Lindl. failed.
From seeds furnished by seed-houses, the University of
Texas and the Mississippi Agricultural College, quite a number
of plants of Martynia louisiana and Martynia fragrans were
grown in Montgomery County, Indiana, during the summers
1915-1919. The two species mentioned above cross very readily,
producing in all cases viable seeds. These plants, grown in the
open in rich soil and under practically the same conditions,
showed very few differences except in the color of the corolla,
which is more or less a variable character. The similarity of
the plants seems to indicate that they belong to the same species,
but more cultures must be studied before this point can be def-
initely determined.
141
142 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
In as muchas no morphological work seemed to have been
done on this family, a study of Martynia louisiana and Martynia
fragrans was undertaken, the results of which are given in the
' following pages. While the study was based mainly on Martynia
louisiana no structural differences between the two species were
observed.
MATERIAL AND METHODS
The material used for this study was collected from plants
of Martynia louisiana and Martynia fragrans grown in the field,
in Montgomery County, Indiana. Collections were usually made
during the month of August, when the plants were most vigorous.
The apices of racemes, small flowers and stamens, pistils and
young fruits in various stages of development were killed and
fixed in a I per cent chromo-acetic acid or in chrom-osmic-acetic
acid one to two hours and then in chromo-acetic acid twenty-four
to thirty-six hours, washed, dehydrated, and embedded through
chloroform. To determine the time necessary for the pollen
tube to reach the egg, pollination was controlled, and, beginning
at one hour after the application of the pollen, collections were
made at intervals of one hour for fourteen consecutive hours.
These collections were of whole pistils which were killed and
fixed in chrom-osmic-acetic acid one to two hours and then in
chromo-acetic acid thirty-six hours. Beginning thirty hours
after pollination, collections of pistils and young fruits were
made at intervals of six hours for two hundred and four hours.
Sections were cut 4-10 wu, most of them 5 u thick; those of whole
pistils were longitudina] and 4-5 y. in thickness. Both modified
triple and Heidenhain’s iron-alum-haematoxylin were used for
staining, the former being more satisfactory.
THE DEVELOPMENT OF THE FLOWER
As in most inflorescences, the flower of Martynia louisiana
arises as a lateral branch (Fic. 1, X) in the axil of a
bract (Fic. 1, B).. The first bract arises as a lateral papilla
near the apex of the main axis of the raceme (Fic. 2, B’).
This bract grows rapidly and about the time its tip reaches
‘the apex of the raceme, a rather broad protuberance appears
in the axil (Fic. 1, X). With further growth of the axis of the
raceme other bracts appear, and in the axils of these the primordia
of other flowers develop (Fic. 4, X). As the rudiment of
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 143
the flower grows it becomes broader and somewhat oblique
at the apex (Fics. 2 and 3, X). On the adaxial side of
the primordium of the flower there appears a protuberance
or ridge, the upper lobe of the calyx, extending about one third
the distance around it (Fic. 3, S). Very soon the other Icbes
of the calyx appear as small protuberances similar to the ad-
axial lobe. The individual lobes appear as separate protuber-
ances, but later grow up as a tube except on the abaxial side of
Fics. 1-7. Development of the flower. Fics. 1-3, X 45; Fics. 3-7,
x 30. B, bract; X, flower primordium; S, S’, calyx lobe; P, corolla lobe; A,
stamen; O, pistil; b, bracteole.
the flower. At this point, the two lower lobes of the calyx are
united only at the base and in the mature flower overlap. The
adaxial lobe of the calyx is the largest, and in the bud covers the
other lobes (Fics. 4 and 7, S$). The calyx lobe, shown atS’ in
Fic. 4, is the edge of one of the lateral lobes and appears smaller
than it really is. Just within the calyx lobes are seen small
144 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
protuberances (Fic. 4, P), the primordia of the corolla lobes, .
which are alternate with those of the calyx. Arising simul-
taneously with the corolla lobes (FIG. 5, P) and alternate with
them are the epipetalous stamens (Fic. 5, A). Fics. 6 and 7
are other sections of the flower shown in Fic. 5. Fic. 6 is a
median section through the primordia of two lobes (P) of the »
corolla, while Fic. 7 is a median section through the primordia
of two stamens (A). A comparsion of these two figures will give
the relative size of the parts at this stage. There are primordia
Fics. 8-13. Development of the flower (con.), X 30. B, bract; b,
bracteole; S, calyx lobe; P, P1, corolla lobe; A, A?, stamen; O, pistil; L, pla-
centa; C, ovarian cavity.
of five stamens, but very early the one between the posterior
pair becomes aborted (Fics. 10 and 14,A’). The pistil is the last
cycle to appear, and in Fic. 5 is seen as a broad protuberance (O).
In Fics. 5-9, at B, are seen cross sections of the bract which
curves over the flower. These figures are of the flower as viewed
from the adaxial side. And in the same figures at b, are seen
sections of the two bracteoles. The two bracteoles arise as
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 145
protuberances similar to those of the lobes of the calyx, but
earlier than the calyx lobes. The bracteoles develop inferior
to the calyx, opposite each other and at right angles to the
bract (Fic. 10, b, B). Since Fic. 9 is a median section through
the flower, only the edge of each bracteole is shown. At this
stage, they are large enough to cover the young flower.
Fics. 8 and 9 show the further development of the parts
of the flower. The calyx (S) is large enough to cover the other
floral parts. An almost median section of two lateral lobes (P)
of the corolla is shown in Fic. 9, while a cross section of the tip
of the lower lobe is shown at P! in Fic. 8. The lower lobe of
of the corolla is the largest and in
the bud is covered by the others.
The manner in which the corolla lobes
overlap is shown in Fic.
median section through two of the
stamens (FIG. 8, A) of the same flower
shows the further development of J5
the stamens seen at A, Fic..7._ Asin
most cases, the development of the
anther precedes that of the filament
(Fic. 8, A). However, at this time
the tissue of the anther is undifferen- Fic. 14. Tees stamen.
tiated. At the time the pollen A, aborted stam
mother-cells are formed the filament oe or Matare ie
is still shorter than the anther. The ee Seiwa sige a
filaments remain short (Fic. 14) until — Fy¢. 16. Mature pistil and
almost the time of anthesis, when they stamensin position. _H, stigma
elongate rapidly, bringing the anthers lobes.
into the throat of the corolla, just below the lips of the stigma
(F.1G 16). The stamens are didynamous, the posterior pair
being the longer. Just before anthesis, the lobes of the anther
diverge, forming a straight line transverse to the filament. At
the same time, the filaments of the corresponding pairs of
stamens converge in such a way as to bring the diverging
anthers into contact (Fic. 16) along the line of their linear
dehiscence. At time of shedding, the rather large pollen grains
are characterized by an exine that is thickened in the form
of hexagonal areas.
As was stated previously, the pistil appears as a rather broad
protuberance (FIG. 5, O), but soon a depression is formed in
the center (Fic. 9, O). The primordium of the pistil continues
146 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
to grow up as a ring enclosing a central ovarian cavity as is
seen in cross section (Fic. 10, C). Ata very early stage in the
growth of the pistil, two parietal placentae appear. These
are seen as outgrowths from the walls of the cavity (Fics. 10
and 11, L). These outgrowths extend toward the center of the
cavity and soon each forms two lobes (Fic. 12, L) which develop
into the two lamellae of each placenta, resembling in cross
section the letter T (Fic. 13, L). The placentae grow together
at the base, thus forming a short columella. A cross section
through the base of the ovary would appear as two-celled, but
no ovules are developed from this region of the placentae. The
upper part of the pistil is tubular and more narrow, forming
a long, slender, hollow style which terminates in an unequally
two-lipped stigma (Fic. 15, H).
DEVELOPMENT OF MEGASPORE AND EMBRYO-SAC
The young ovule first appears as a small, nipple-like protub-
erance upon the placenta. Very early in its development, a
hypodermal cell near the tip becomes differentiated and may be
distinguished from the surrounding cells by its large nucleus and
dense cytoplasm (PLATE 7, FIG. 17). Thiscell, the megaspore
mother-cell, grows rapidly, keeping pace with the lengthening
nucellus. At this stage, the nucellus consists of merely the en-
larged megaspore mother-cell, covered by the epidermis (FIG. 18).
At the base of the nucellus the single integument appears as a fold,
at first more pronounced on the side next to the cavity of the
ovary (Fic. 18, 1). By the more rapid growth of this side the
ovule soon becomes sharply curved, so that the long axis of the
nucellus is now parallel with that of the funiculus. At this
stage, the integument extends around the nucellus, which has
become more elongated (Fic. 19). With further growth the
integument soon equals and finally exceeds the nucellus (FIGs.
20.and 21). The megaspore mother-cell has also grown rapidly
and when fully developed is still enclosed by only a single layer
of nucellus (Fic. 21). The nucleus, which has increased in
size preparatory to the first division and which has had a posi-
tion near the center of the cell (Fic. 20), comes to lie nearer the
chalazal end, where it undergoes the first or heterotypic division
(Fic. 21). All the spindles of this stage found were rather small
as compared with the size of the cell. Whether cell formation.
follows this first nuclear division was not ascertained, but the:
ANDERSO:: EMBRYOGENY OF MARTYNIA LOUISIANA 147
typical axial row of four megaspores is finally formed (Fic. 22),
and, as is usually the case, the innermost megaspore, the one
next the chalazal end, persists to form the embryo-sac (Fics.
22 and 23). The upper three potential megaspores are soon
destroyed by the rapid growth of the functional one (Fic. 23.)
By the time the tetrad is formed, the nucellus has reached
its maximum growth, and the cells of the integument adjacent
to the nucellus have become dense in cytoplasm and now func-
tion as a nutritive jacket (Fic. 22). In the meantime the
integument has grown and exceeds considerably the length of -
the nucellus (Fic. 23).
The functional megaspore does not increase much in size
before the first nuclear division but, after this first division, it
elongates rapidly and occupies a space almost the length of
the nucellus (Fics. 23 and 24). As the megaspore elongates
the two nuclei keep their relative positions—one in each end
of the cell. Most of the cytoplasm of the cell is in the ends,
thus leaving a large vacuole in the center. A few strands of
cytoplasm penetrate the vacuole and form a connection between
the masses at each end of the cell. The rapid growth of this
cell has consumed the three upper potential megaspores and the
nucellus around the center of the spore (FIG. 24). Before the
second nuclear division of the megaspore takes place, the upper
three megaspores have practically disappeared and the functional
one now extends nearly the length of the nucellus, which is rapid-
ly being used as food (Fic. 25). At this time the central vacuole
is not so pronounced—at least in some cases. The spindles of
the second nuclear division may be seen—one in each end of the
cell. With the exception of a few cells at the chalazal end, the
nucellus is consumed by the growing megaspore, which comes
to occupy the space left by the disorganizing tissue. There
is no evidence of a nucellar cap being formed as Merrell reported
for Silphium (3) and other plants that have a similar develop-
ment. The nuclei now undergo another division which results
in eight nuclei—four in each end of the cell. Next follows the
formation of the three cells of the egg-apparatus and the three
antipodals. One nucleus from each end of the embryo-sac, the
polar nuclei, remains free in the cytoplasm. At this stage, the
embryo-sac is considerably elongated, being about seven times as
long as wide. The micropylar end, containing the egg-apparatus,
is enlarged to about one and a half times the diameter of the
Other part of thesac. This enlarged portion comprises nearly one
148 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
third the length of the embryo-sac. The egg-apparatus consists
of the large pear-shaped egg-cell, which extends far down into
the sac cavity, and the two large synergids (PLATE 8, FIG. 26,
GandSn). The lower part of the egg in which the nucleus lies is
well filled with cytoplasm, but the upper part is vacuolate. The
synergids, one on either side of the egg, extend down about two
thirds the length of the egg-cell. Each synergid has a diameter
nearly equaling that of the egg and a centrally located nucleus,
which is a little smaller than the egg nucleus. As is frequently
found in other angiosperms, a fairly large vacuole lies below the
nucleus in the synergid (Fic. 26, Sn).
In the chalazal end of the embryo-sac are the three, rather
long, slender, well-developed antipodals (Fic. 26, N). In most
instances, the nuclei of the antipodals are fairly large and the
cytoplasm has the character of that of the body of the embryo-
After the formation of the cells at either end of the embryo-sac
the polar nuclei move to the center and lie side by side for some
time (Fic. 26, Pn), but before the pollen tube reaches the embryo-
sac, the two polar nuclei fuse (FIG. 27). Sections were made of
ovaries from flowers in which pollination had not occurred and
it was found that the polar nuclei had fused (Fic. 28). So the
fusion of the polar nuclei is in no way due to a stimulus from
the pollen tube. The fusion nucleus does not approach the egg-
apparataus, as Kanda reported for Verbena (2), but remains in
the center of the embryo-sac. Each polar nucleus has a very
prominent nucleolus and after fusion the two nucleoli are very
noticeable for some time (FIG. 27), but later only one large
nucleolus is usually seen (Fic. 28).
POLLINATION
The polar nuclei seem to unite about the time of anthesis,
for conditions like those represented in F1G. 26 are observed in
ovaries fixed just before the expansion of the corolla lobes, while
the fusion nucleus is seen in flowers that have been open one or
two hours. Most of the flowers open early in the morning, and
shortly after daylight the bees, which are the chief pollinizing
agents, are busy gathering nectar from the lower portion of the
corolla tube.
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 149
The two lobes of the sensitive stigma stand open just above
the open anthers (FIG. 16), in the upper portion of the throat
of the corolla, and the bee, entering the corolla tube, brushes
the stigma, thus depositing the pollen that may be present on
the upper part of its body and securing a new supply from the
open anthers just below as it walks down the tube. The stigma
is very sensitive when the flower first expands and the slightest
touch will cause the lobes to close. If pollen be present
they usually remain closed, but, in the absence of pollen, they re-
open in from five to ten minutes. If pollen is witheld the stigma
lobes may be made to close and open many times, but after one
to three days the stigmas tend to lose their sensitiveness.
However, this sensitiveness is not an index to the receptiveness
of the stigmatic surface, for flowers hand pollinated after the
stigma ceased to be sensitive and even after the corolla dropped,
produced viable seeds. Viable seeds were also secured from
flowers hand pollinated in the bud before the stigma lobes nor-
mally opened. This was accomplished by gently forcing pollen
between the closed lobes by means of a needle. In most flowers
thus treated the stigma lobes never opened. At this time the
anthers are still closed, but pollen from these as well as from
open anthers was shown to be effective.
FERTILIZATION
Usually the pollen grains germinate immediately on the stig-
matic surface, as was readily seen in material fixed one hour
after the pollen was applied. At this time, many of the tubes
were well into the tissue of the style. The growth of the pollen
tube through the tissue, usually two or three cells from the
stylar canal, is very rapid, for fertilization frequently occurs
within six hours after pollination. The pollen tube is filled
with plastids and granules that take such a dense stain that it
is impossible to demonstrate the presence of the sperm nuclei,
but, since the long, slender crescentic generative cell in the mature
pollen grain shows only one nucleus, it is thought that the for-
mation of the two sperms takes place in the tube. The pollen
tube, as it grows nearly straight down through the style to the
placenta, is comparatively broad and seems to retain its densely
staining quality throughout the greater part of its length.
150 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
On reaching the ovule, the pollen tube
grows through the micropyle and enters
the embryo-sac, sometimes a little to one
side, crushing one or both the synergids.
The tip of the pollen tube usually covers
the egg-cell, thus obscuring the nucleus of
the latter (Fic. 32). In some instances it
seems that the tube enters the egg-cell at
the upper end and transverses the entire
length of it (Fic. 31). In all cases ob-
served, there was evidence that the tube —
had discharged the sperm nuclei in such
a way that one of them came in direct
contact with the egg nucleus (FIG. 30).
Only the sperms and a little of the con-
tents from the tip leave the pollen tube
(Fics. 30 and 31). The fate of the
second sperm nucleus was not ascertained,
but it seems evident that it plays no part
in the formation of the endosperm. The
second sperm nucleus may not leave the
Pes
2 ae somes ts Oe —
Ce ae ae
ae
ae
=
S Deg r~
= wr
=<
>
RP aan tS
geet,
oe
i Sas as 2 oO BF tr
f 3.
Hie tube, but this point can not be demon-
et strated until some method of fixing and
tana staining can be found that will reveal the
=
ee
ies: :
root presence of the nuclei among the granules
{]
A | 5 on Oy 8
ineeren in the cytoplasm. The pollen tube in the
micropyle and in the upper part of the
ip
eke
embryo-sac retains its form and densely
staining quality long after the embryo-sac
is filled with endosperm (FIG. 35).
The actual union of sperm and egg nu-
cleus is similar to that in other angio-
sperms. The instances of fertilization
observed were similar to that of Lilium
Martagon, figured by Mottier (4). For
Verbena, Kanda (2) figures the sperm
nucleus in contact with the egg-cell and
speaks of it as fertilization. In all the
fertilization stages observed in Martynia,
Fic. 35. Embryo-sac filled with endosperm,
xX 140. T, pollen tube; N, antipodali; E, embryo;
U, suspensor.
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 165i
the sperm has been in direct contact withthe egg nucleus.
This would seem to indicate that the pollen tube carries the
sperm into the egg-cell.
THE ENDOSPERM
The primary endosperm nucleus, resulting from the union
of the two polar nuclei, occupies a position near the center of
the embryo-sac. Usually before the pollen tube enters the sac
this endosperm nucleus divides and a transverse wall is formed
between the two resulting nuclei, thus dividing the embry-sac
into two chambers (Fic. 29). At this time, the cytoplasm in
the embryo-sac has a frothy appearance due to the many small
vacuoles present. This frothy character seems to appear after
the fusion of the polar nuclei (Fic. 28). Very’ conspicuous,
densely staining plastid primordia and granules are present in
the cytoplasm. The synergids seem to have increased in length
and to have become more vacuolate (FIG. 29). The egg-cell
remains apparently unchanged. The antipodals are more
elongated and sometimes contain a granular, densely staining
cytoplasm (Fic. 29). After the wall dividing the embryo-sac
into two chambers is formed, the nuclei move to a nearly central
position in their respective cells. And, as one would infer, the
endosperm cell at the micropylar end of the sac grows more
rapidly than the one at the chalazal end (Fic. 31). The pollen
tube usually reaches the embryo-sac when the endosperm is in
a two-celled stage. In Fic. 31, it may be seen that fertilization
has taken place. The outline of the tip of the pollen tube is shown
by a dotted line in the lower part of the egg-cell, and the shading
represents the relative density of the contents in different parts
of the tube. Only the tip seems to have discharged its contents.
A remnant of one synergid still shows. The synergids, if
not crushed out by the entrance of the pollen tube, are soon ab-
sorbed. The endosperm cell in the micropylar end of the sac
is very large and contains a large nucleus with irregular outline
and a prominent nucleolus. Many plastids are still seen in the
frothy cytoplasm of both endosperm cells, and some are present
in the antipodals, which continue to grow, especially in length
(Fic. 31). The increase in size of the antipodals is better illus-
trated in Fic. 32. Their cytoplasm is somewhat vacuolate at
this stage and the large nuclei contain very prominent nucleoli.
Usually the first division of the endosperm cell is transverse
152 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
and the second longitudinal, the one in the micropylar end divid-
ing before the one in the chalazal end; but, in FIG. 32, it appears
as if the first division had been longitudinal and the second
transverse. The cytoplasm in the large cells around the fertilized
egg contains many large as well as small vacuoles, while
that of the lower cells contains only small ones. Very few
plastids are present at this stage. They are seldom noticeable
in later stages of endosperm development. Fic. 32 shows also
the old densely staining pollen tube in the micropyle and upper
end, of the embryo-sac, its tip covering the fertilized egg. By
focusing down on the tip of the tube, the
nucleus of the fertilized egg is seen and is
represented in the drawing by a dotted line
(Fic. 32). In the meantime, the whole em-
bryo-sac has been enlarging at the expense
of the integument which also has grown.
The enlargement is more pronounced at first
in the micropylar end where the fertilized
egg is elongating. The layer of cells of the
integument adjacent to the embryo-sac is
always dense and rich in cytoplasm and func-
tions as a nutritive jacket to the growing ~
endosperm. Fic. 33 shows the further de-
velopment of the endosperm, and the
elongation of the fertilized egg. The old
pollen tube is still very prominent in the
G56.” Outils micropyle and upper end of the sac, obscur-
of Pie x 44. I, integ- ing the upper portion of the fertilized egg-cell.
ument; M,embryo-sac The cytoplasm in the large endosperm cells
E, embryo; U, suspen- of the micropylar end is very vacuolate, due
sor; T, old pollen tube.
probably to the rapid growth of the cells and
to the loss of food to the developing embryo. Only a few
endosperm cells develop in this end of the sac, for very soon
a long suspensor is formed which brings the embryo into the
chalazal end. The endosperm cells in the middle of the sac
are smaller and richer in cytoplasm than the upper ones. The
antipodals have nearly reached their maximum growth. They
seem to function as elaborators of food and persist long after
the embryo-sac is well filled with endosperm (FIG. 35). In
several cases, four antipodals, all well developed, were found
(Fic. 33). The extra one had probably been formed by the
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 153
division of one of the three normal ones. FIG. 34, a cross section
of the chalazal end of an embryo-sac in the same stage as the
above, shows the arrangement of the four antipodals. At this
stage, the nuclei of the antipodals appear as densely staining
homogeneous masses.
As the embryo with its very long suspensor grows through
the center of the elongating embryo-sac, large endosperm cells
rich in cytoplasm are formed, but when it reaches the lower
part of the sac where the first division of the embryo takes
place, the endosperm growth practically ceases in the middle
and upper part of the sac. The cells of this part of the sac are
rather large and similar in shape to those first formed, while
those in the lower part are smaller and not so rich in cytoplasm
(FIG. 35). It is also seen from Fic. 35 that the embryo-sac has
enlarged considerably, especially the lower part which contains
the developing embryo. The nutritive jacket is still very active
in the elaboration of food for the growing endosperm. FIG. 36
shows in outline the relative size of the embryo, endosperm and
integument at the time the embryo is in the two-cell stage as
seen in FIG. 35.
THE EMBRYO
After fertilization, the egg with its nucleus occupying a
nearly central position in the lower part of the cell grows rapidly
in length (Fic. 33). Its manner of growth down through the
endosperm is similar to that of a pollen tube, for it becomes a
long narrow cell with the nucleus retaining a position near the
growing tip. No division seems to take place until the distal
end has transversed nearly two thirds the length of the embryo-
sac. The lower portion of the cell is rather dense in cytoplasm
and stains deeply, but the upper portion has very little and
stains so faintly that the course of its growth is hard to trace.
After the distal end has reached the lower portion of the
embryo-sac, a transverse division occurs, which forms a very long,
slender suspensor cell and a short, rounded distal one, the em-
bryonal cell (Fic. 37). In most instances the cytoplasm of the
embyonal cell contains many plastids and other densely staining
granules, while that of the suspensor is very poor in plastids.
After the first division two others occur almost simultaneously.
The suspensor cell divides, forming a short cell next the embryo-
nal cell, which has enlarged preparatory to division, and, at the
154 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
same time, the enlarged embryonal cell divides longitudinally
forming a two-celled embryo (Fics. 38 and 35). The second
division of the embryo is usually transverse and the third longi-
tudinal, thus forming the octant stage The further growth of
the embryo is illustrated in Fics. 39-43 and is so like the well
known development of Capsella that details are unnecessary.
The very long suspensor, shown in Fig. 35 in dotted
lines, is almost indentical with that of Trapella sinensis Oliv. as —
figured by Oliver (5, pl. 7, f. 36). The suspensor of Myoporum
ii
2
Cy
¢,
we
ang Oe,
‘
‘
,
t
'
'
4
4
1
‘
1
1
‘
Fics. 37-43. Development of the embryo. Fics. 37-38, X 500; FIGs.
39-42, X 175; Fic. 43, X 6. E, embryo; U, suspensor; C, cotyledon; R,
radicle; K, chalazal end of embryo-sac; Q, micropylar end of embryo-sac;
N, remains of antipodals; V, soft, inner portion of seed coat; W, hard, warty
portion of seed coat.
serratum as figured by Billings (1, f. sz) is also similar in struct-
ure but does not push the embryo as far down into the embryo-sac
as that of Martynia, although the shape of the embryo-sac and
the development of the endosperm are much the same in both
species. Usually a second and sometimes a third division
occurs in the suspensor, resulting in two or three short cells
next the embryo, the innermost of which seems to contribute to
its formation.
Fic. 43 shows a section of a young seed containing a nearly
mature embryo. The radicle (R) is short and thick while the
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 155
cotyledons (C) are very large and fleshy. This figure shows
also a little endosperm which, in the mature seed, comprises
a thin, transparent covering for the embryo. The embryo has
developed in such a way that it practically fills the lower part
of the sac, while the upper and middle portions (Q) form in the
mature seed a little brown tube on the micropylar end of the
remaining endosperm. There is also a small brown knob at
the chalazal end (K) marking that part of the embryo-sac for-
merly occupied by the large antipodals, the remains of which
appear at N. The broken line in this figure represents the
demarcation in the tissue forming the seed coat. That region
of tissue next the endosperm (V) matures into the soft inner
portion of the seed coat, while the walls of the cells forming the
outer portion (W) become thick and lignified, thus maturing
into a hard warty exterior which becomes very dark brown or
black in color.
SUMMARY
The flowers develop in the axils of protective bracts. The
cycles of the flower appear in the following order: calyx, corolla
and epipetalous stamens, and pistil.
Of the four potential megaspores formed, only one is function-
al.
The embryo-sac is long and narrow, with the micropylar end
considerably enlarged. The large egg-cell extends far down
into the sac cavity, with the two synergids on either side. The
three antipodals are long, slender cells that increase in size and
persist as elaborators of food.
he polar nuclei move to a position near the center of the
sac where they unite before the pollen tube reaches the ovule.
Fertilization usually occurs within six to eight hours after
pollination.
The rather large pollen tube stains densely throughout the
greater part of its length and persists in the micropyle and upper
end of the embryo-sac long after the endosperm is formed.
The first division of the endosperm cell, which is usually
transverse, occurs before. fertilization. At first, a few cells
are formed in the upper and middle portions of the embryo-sac,
but the bulk of the endosperm is in the lower part of the sac,
where the embryo develops.
The suspensor is very long and narrow and pushes the embryo
156 ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA
into the lower portion of the embryo-sac, where the first division
takes place. The development of the embryo proper is like
that of Capsella.
I take opportunity here to express my obligations to Pro-
fessor H. W. Anderson for the use of the Wabash College Bot-
anical Laboratory during the summers of 1915 and 1916 and to
Professor D. M. Mottier for valuable assistance and criticism
in connection with this study.
INDIANA UNIVERSITY
LITERATURE CITED
1. Billings, Frederick H. Beitrage zur Kenntniss der Samenent-
wickelung. Flora 88: 253-318.
2.Kanda, M. Field and laboratory studies of Verbena. Bot.
Gaz. 69: 54-71. pl. 6-9 + f. 10-17, 27, 54-63, 09-75. 1920.
3. Merrell, W. D. A contribution to the life history of
Silphium. Bot. Gaz. 29: 99-133. pl. 3-10. 1900
4. Mottier,D.M. Fecundation in plants. Carnegie Inst. Wash-
ington Publ. 15. rg04.
5. Oliver, F. W. On the structure, development, and affinities
of Trapella, Oliv., a new genus of Pedalineae. Ann.
2: 75-115. pl. 5-9. 1888
— of plates 7 and 8
PLATE 7
All figures, X 400
Fic. 17. Vertical section through the nucellus; megaspore mother-cell
shaded.
Fic. 18. Elongated nucellus with megaspore mother-cell; beginning of
integument, I.
Fi rtical section of young ovule turned on its axis; the integu-
ment reaching half the length of the nucellus.
Fic.
20. Nearly mature megaspore mother-cell enclosed in a single
layer of nucellus.
FIG Heterotypic soit in megaspore mother-cell.
1G. 22. Tetrad, the upper three potential megaspores disorganizing.
Nutritive jacket formed from fis layer of integument adjacent the nucellus.
F Section of young ovule showing the single integument and
first division of the megaspore; the upper three potential megaspores dis-
organizing.
Fics. 24 and 25. Further development of the megaspore; the central
portion of wate disorganizing.
iS
Loa)
.
VOLUME 49, PLATE 7
Bul. Torrey Cus
AME eS nan ee weet e
ee STE eco a
5 ER AAU No
y Naat INNS
OATS TTR TERS
Tate A
Aa
FLOWER AND EMBRYOGENY OF MARTYNIA
ANDERSON:
VCLUME 49, PLATE 8
Buti. TorrEY CLus
ANDERSON: FLOWER AND EMBRYOGENY OF MARTYNIA
ANDERSON: EMBRYOGENY OF MARTYNIA LOUISIANA 157
PLATE 8
FiGs. 26-33, longitudinal sections of embryo-sacs. F1G. 34, cross section
of antipodals.
Fic. 26. Mature embryo-sac; polar nuclei side by side near the center
of the sac; G, egg-cell; 5 synergid; N, antipodals; Pn, polar nuclei, * 300.
Fic. 27. Mature embryo-sac; polar nuclei fusing, x 300.
Fic. 28. Mature saheeaie: polar nuclei fused, xX
Endo eae:
.30. Fertilization. Sperm and egg nuclei in contact: old _
tube nt prominent; G, egg-cell; T, pollen tube; Sn, synergid,
Fics. 31 and 32. Development of endosperm and elongation oF ‘nbs.
podals, X 350.
Fic. 33. Development of endosperm and embryo; four antipodals, N;
E, elongating embryo; T, old pollen tube, X 350.
FIG Cross section of chalazal end embryo-sac showing four anti-
G. 34
podals, N, X 350.
INDEX TO AMERICAN BOTANICAL LITERATURE
1921
The aim of this Index is to include all c t botanical literature written by
Americans, published in America, or rt upon American material; the word
America being used in the broadest sen:
Reviews, and papers that relate ak a to foresty, agriculture, horti-
Seaebic ee saree! of vegetable origin, or labora atory methods
f bacteriology.
An occasional pene is made i in favor of some paper appearing in an Ameri-
can 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 satis attention of the editor to errors or omissions, their
kindness will be a ed.
This Index is fuk wt monthly on cards, and furnished in this form to
subscribers at the rate of three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Adams, J. F. Observations on the spores of Sichzophyllum
commune. Torreya 21: 98-100. D Ig2i._ [Illust.]
Allen, W. E. Some work on marine So kgecpeceeioe Trans.
Am. Micr. Soc. 40: 177-181.
Allen, W. E. The investigation of ocean pasturage. Ecology
2: 215-219... 4 De 1ga1.
Ames, O. Additions to the orchid flora of Panama. Proc. Biol.
Soc. Washington 34: 149-154. 21 D 1921.
Includes 7 new species. nee
Ames, O. Eria gigantea. Gigantic eria. Addisonia 6: 41,
42, pl. 273. 240 Wet.
Anderson, E. G. The inheritance of salmon silk color in maize.
Cornell Univ. Agr. Exp. Sta. Mem. 48: 539-554. N 1921.
Anderson, P. J., & Ickis, M. G. scree aaa pe of
Helvella. Mycologia 13: 201-229. pl. 11,
Arthur, J. C. Memoranda and index of cultures of Uredineae,
1899-1917. Mycologia 13: 230-262. 1921.
Ashe, W. W. Notes on Rhododendron. Rhodora 23: 177-179.
24 O 1921.
159
160 INDEX TO AMERICAN BOTANICAL LITERATURE
Bailey, L. ae sapiens stellata. Addisonia 6: 37, 38. pl. 2zr.
24 O1
Ball, C. R. ae on willows of sections Peniandreae and Nigrae.
Bot. Gaz. 72: 220-236. f. 1-4. 15 O 1921
Includes 2 new varieties of Salix from California.
Barker, E. E. Bud variation in the sugar cane. Jour. Hered.
LP BG 1374. $6.27 = 25.9 19021.
Bechtel, A. R. The floral anatomy of the ls sor Am.
Jour. Bot. 8: 386-410. pl. 15-22. 14 N 19
Bird, si Soil acidity in relation to insects and Sie Ecology
2: 193-197. ‘
Birge, ‘ A., & Juday, C. Further Gracia observations
on the Finger Lakes of New York. Bull. Bur. Fisheries
a4 211252, J. 2-5, 8 O 1071.
Includes notes on plankton botany.
Blakeslee, A. F., Cartledge, J. L., & Welch, D. S. Sexual
dimorphism in Cunninghamella. Bot. Gaz. 72: 185-219.
Jets I§ OC ie.
Boynton, K. R. Bergenia crassifolia. Thick-leaved saxifrage.
Addisonia 6: 43. pl. 274. 24 O 1921.
Boynton, K. R. Helichrysum bracteatum. Strawflower.
Addisonia 6: 46. pl. 275. 24 O 1921.
Bradshaw, R. V. Cynosurus echinatus in Oregon. Torreya
> 81-83. O 1921:
Braun, E. L. Composition and source of the flora of the Cin-
cinnati (Ohio) region. Ecology 2: 161-180. f. z. 3 D 1921.
Briggs, G. Para and Paspalum: two introduced grasses of
Guam. Guam Agr. Exp. Sta. Bull. 1: 1-44. pl. 1-6. 2
D 1921.
Britton, E. G. Clerodendron Thompsonae. Mrs. Thompson's
clerodendron. Addisonia 6: 39, 40. pl. 212. 24 O 1921.
nes = L. Lathyrus latifotius. Everlasting pea. Addiso-
6:35. pl. 210. 24 O 1921.
pain. N. L. Plant nomenclature: more suggestions. Jour.
Bot. 59: 296, 297. O 1921.
Brotherston, R. P. The pentstemon. Gard. Chron. 70: 208.
2°Q. I92t.
§
INDEX TO AMERICAN BOTANICAL LITERATURE 161
Burlingame, L. L. Variation and heredity in Lupinus. Am.
Nat. 55: 427-448. f. 1-3. O 1921.
Cayla, V. A propos de la qualité du cauotchouc d’Hevea
brasiliensis, ses variations, leurs causes. Importance
economique pour la. production de Il’Amazone. Arch.
Escol. Super. Agr. e Med. Veter. 5: 81-96. S 1921.
Chace, E. M., Wilson, C. P., & Church, C.G. The composition
of California lemons. U. S. Dept. Agr. Bull. 993: 1-18.
f. 1-4. 15 O 1921.
Chamberlain, C. J. Growth rings in a monocotyl. Bot. Gaz.
72: 293-304. f. 7-16. 15 N 1921.
Clark, A. H. A new classification of animals. Bull. Instit.
Oceanogr. 400: I-24. 20S Iga2l.
Includes a summary of plant classification.
Clements, F. E. Drouth periods and climatic cycles. Ecology
2: 181-188. 3 DD 162i
Coker, W. C., & Beardslee, H. C. The collybias of North
Carolina. Jour. Mitchell Sci. Soc. 37: 83-107. pl. 1 +f. 4-23.
D 1921
Includes Collybia lilacina, sp. nov.
Cook, F. C. Absorption of copper from the soil by potato
plants. Jour. Agr. Research 22: 281-287. 29 O Ig21.
Cook, M.T. The blossom blight of the peach. Phytopathology
11: 290-294. pl. 12. 15 N 1921
Cook, M. T. Falling foliage. Phytopathology 11: 337-339.
a LY toatl,
Cook. M. T. Peach yellows oa ae peach. Bot. Gaz. 72:
250-255. pl. 6, 7. 15 OI
Cook, M. T. Wilting caused by site trees. Phytopathology
11: 346. 9
Cook, O. F. Causes of shedding in cotton. Jour. Hered. 12:
199-204. f. 3-6. 23S 1921.
Coville, F. V. Directions for blueberry culture, 1921. U. S.
Dept. Agr. Bull. 974: 1-24. pl. 1-29. 15 O 1921.
Creswell, C. F., & Bidwell, G. L. ates of cotton seeds.
U. S. Dept. Agr. Bull. 948: 1-221. pl. 1+ maps I-11. 105
1921.
162 INDEX TO AMERICAN BOTANICAL LITERATURE
Dana, B. F. Two new Sclerotinia diseases. Phytopathology
11: 225-228. pl. 8. 6 O 1921.
Doolittle, S. P. Overwintering of the bacterial wilt of cucur-
bits. Phytopathology 11: 299, 300. 15 N 1921.
Dupler, A. W. The air chambers of Reboulia hemisphaerica.
Bull. Torrey Club 48: 241-252. f. 1-22. 13 D 1921.
Ehlers, J. H. Panicum virgatum var. cubense in Michigan.
Rhodora 23: 200. 15 D 1921.
Eldridge, A. E. Woodland flowers of the prairie region. Am.
For. 27: 714-716. N 1921. [Illust.]
Emerson, F. W. Subterranean organs of bog plants. Bot.
Gaz. 72: 359-374. f. I-1r. 15 D 1921.
Emerson, R. A. Genetic evidence of aberrant chromosome
behavior in maize endosperum. Am. Jour. Bot. 8: 411-424.
Jor 14: N 1931.
Ensign, M. R. Area of vein-islets in leaves of certain plants
as an age determinant. Am. Jour. Bot. 8: 433-441. pl.
azo 9 I
Evans, A. W. The genus Riccardia in Chile. Trans. Con-
necticut Acad. Sci. 25: 93-209. f. 1-13. D 1921.
Includes 3 new species and 17 new combinations. -
Fairman, C.E. The fungi of our common nuts and pits. Proc.
Rochester — Sci. 6: 73-115. pl. 15-20. S 1921.
Includes Sch. 14.
, gen. nov ,and 27 new species in various genera.
Fehlinger, H. Beatieninde Faserpflanzen. Tropenpflanzer
24: 173-176. D 1921.
Fujioka, M., & Takahaski, K. On the cause of the darkening
of the heartwood of ec caunies japonica Don. Jour. For.
19: 844-866. D 1921
Gardner, G. A. onenaiees sur les articles du Reverend Pére
Fontanel, S.J. Nat. Canad. 48: 49—-56.
Gardner, M. W., & Gilbert, W. W. Field tests with cucumber
angular leafspot and anthracnose. Phytopathology 11:
298, 299. 15 N 1921.
Gardner, M. W., & Kendrick, J. B. Soybean mosaic. Jour.
Agr. Research 22: 111-113. pl. 18 19. 8 O 1921.
Vol. 49 / No. 6
BULLETIN
OF THE
TORREY BOTANICAL CLUB
JUNE, 1922
Mycological notes for 1920*
L. O. OVERHOLTS
. (WITH PLATE 9 AND FOURTEEN TEXT FIGURES)
1. ZYTHIA RESINAE (Ehrenb.) Karst.
For several years there has been noted in the mountains
near State College a small inconspicuous fungus always confined
to resin exudate on species of two- and three-needle pines. It
has been found especially abundant in the vicinity of Charter
Oak and Greenwood Furnace, Pennsylvania. Its habitat is
the resin mass inhabited by the larvae of the ‘pitch midge,”
Cecidomyia resinicola Osten-Sacken. The origin of the resin
exudate has not been studied by the writer, but E. P. Feltt
states (on the authority of Miss L.S. Eckel) that the resin flow
from the tree is not the result of an injury caused by the
midges, but that they ‘‘take advantage of normal exudations
of resin on which to oviposit,” although ‘‘the flow of this
substance may be increased to some extent by the larvae
rasping the living tissues.” At any rate the final resin masses
are subglobose or depressed globose and 1.5-2.5 cm. in diameter
(PLATE 9, FIG. 2). They may occur on one side of the limb only,
as stated and illustrated by Felt, or the exudate may entirely
encircle the limb, though I have seen no instances of the saccate
type of resin mass illustrated by. Felt. These masses are in
*Contribution from the Department of Botany, The Pennsylvania
State College, No. 38. ‘‘Some mycological notes for 1919’’ was published in
Mycologia 12: 135-142. 1920.
TN. Y. State Mus. Memoir 8: 410-412. 1906.
[The BuLLETIN for May (49: 123-162. pl. 5-8) was issued June 15, 1922.]
163
164 MYcOLOGICAL NOTES FOR 1920
all cases found near the ends of the branches and always at the
forks of the twig
Eventually ee fanuti produces its pycnidia on this hardened
and blackened resin mass. These are minute elongate-
ovoid or almost conical bodies (PLATE, 9, FIG. I, at left), yellow
or amber-colored, and large enough to be easily visible, on close
inspection, to the unaided eye. The ostiole is usually quite
prominent, and may often be seen with a hand lens. In texture
the pycnidia are waxy and easily crushed when mounted in
water. The spores are globose, smooth, hyaline, and very small,
measuring only about 2 y. in diameter.
The nature of the pycnidium places the fungus in the Nec-
trioidaceae of the Fungi Imperfecti. After several unsuccessful
attempts at identification, the species was determined for me
as Zythia resinae by Mrs. F. W. Patterson, who wrote that my
specimens represented the only American material she had seen.
Later Dr. Seaver identified my plants as being the conidial
stage of Nectriella resinae (Fr.) Sacc., as listed by Saccardo.
Nectriella is a genus of the true Nectriaceae, order Hypocreales,
and consequently with an ascus in the life history. Examination
of material collected at various times over a period of several
years has failed to show on the same substratum any ascospore-
producing structure comparable to the perithecium of a Nectriella.
There is produced, however, on the same resin masses that bear
the pycnidia an apothecial fructification that has been referred
to Biatorella resinae (Fr.) Mudd., as listed below.
2. BIATORELLA RESINAE (Fr.) Mudd.
In company with Zythia resinae, as described above, there
frequently grows an apothecial fruiting body that was determined
as Biatorella resinae by Dr. L. W. Riddle, to whom my specimeris
were sent by Dr. Seaver. The very small apothecia are about
I mm. in diameter and yellowish or amber-colored (PLATE 9, FIG.
I, at right). They have been collected on resin exudate in-
habited by the larvae of the “pitch midge” on Pinus virginiana
at Charter Oak and on P. ponderosa at Greenwood Furnace.
The curious habitat and the frequent association of these apo-
thecia with the pycnidia of the Zythia suggest a connection
between the two, but if the Zythia is the pycnidial stage of the
Nectriella such could hardly be the case.
MYCOLOGICAL NOTES FOR 1920 165
3. PmLacrE Petersit B. & Br.
This rare species was found for the first time in the writer’s
experience on a dead snag of Acer rubrum, July 1, 1920, on Stone
reek, Huntingdon County, Pennsylvania. Hundreds of the
small brown stipitate fruiting heads were present on a small
area (PLATE 9, FIG. 4), and their abundance suggested that
this species is probably not so rare as has been supposed but,
because of its small size and inconspicuous coloration, is usually
@ ©
Fics. 1-4. PILAcre Perersi B. & Br.
1. Hyphae with clamp connections. 2 and 3. Basidia with spores.
4. Spores.
overlooked. Cross-septate basidia characteristic of the Auricu-
lariales are always difficult to demonstrate, but the writer has
located them with more ease in this species than in any other,
though not readily demonstrable at best (TEXT FIGS. 2 and 3).
The fungus departs widely in consistency and habit from other
members of the order. The following descriptive notes are
appended:
Sporophores gregarious on the daecst sae cancun Seow
ing from a slight superficial gray mycelium, 2-5
consisting of a slender stalk bearing a tits Siobhan or
hemisphaeric head; head 0.5-2.5 mm. in diameter, at first gray-
pruinose, then deep brown, covered by a thin membrane that
disappears at maturity, filled with a ‘perl spore mass mixed
either side of the basidium if — but on the convex surface
if curved; hyphae re then brown, somewhat bra bey
with cross walls and clamp connections, 2-3 yw in premix
166 MYCOLOGICAL NOTES FOR 1920
stem central, grayish white, 0.2-0.35 mm. in diameter. [TEXT
FIGS. I-4.]
On dead wood of deciduous trees.
4. TULASNELLA VIOLAE (Quél.) Boud. & Gal.
The genus Tuwlasnella, recently monographed for this
country by Burt, is an anomalous genus, Corticiwm-like in
appearance, but hardly so in structure. It is characterized by
the presence of large sterigmate-like processes, four in number
(usually), that are produced on a basidium-like organ, and that
in turn bear each a hyaline spore. Only three species are list-
ed by Burt, all of narrow geographical range and none hitherto
reported from Pennsylvania.
Asmall collection of T. Violae was made in the mountains of
Center County, Pennsylvania, on January 29, 1921, on the inside
of a gum (Nyssa sylvatica) log by the writer’s mycology class while
on a collecting trip. The discovery of the specimens is credited
to Mr. Kuntz
The color of these specimens was flesh pink or livid pink
(Ridgway) when collected, but it is not probable that they will
long retain these colors in the herbarium. In aspect the plants
resemble very much a thin species of Corticium, and casual ob-
servations of free hand sections show nothing very different
from sections of species of that genus. Spores were abundant
in the collection, however, and it was only when the usual
attempt was made to substantiate their connections to basidia
that the real nature of the hymenium became apparent. With
some difficulty the large clavate or elongate sterigmata were
made out, sometimes four in number and sometimes apparently
but three. After the development of the sterigmata the basidia
appear to collapse, or at least in a spore-bearing condition the
group of sterigmata is so large as to give this impression. The
spores are broadly ellipsoid to globose, and in these specimens
have two or three large conspicuous guttulae The spores
measure 5.5-7.5 X 5 u. No cystidia are present.
5. DACRYOMYCES HYALINA Quél.
The species of the genus Dacryomyces are not well known in
this country, partly because, in spite of their usually bright
colors, they dry down and are rather inconspicuous in dry
weather, when collectors are most likely to be in the field. D.
hyalina, as determined for me by C. G. Lloyd in 1919, was
MYCOLOGICAL NOTES FOR 1920 167
found for the first time in Pennsylvania on January 29, 1921,
on a fallen log of Pinus Strobus. The dried plants appear as
small scaly growths, dark in color, but when wet they swell
up and take on a more waxy appearance and gelatinous texture.
Fresh specimens measure I-2 mm. in diameter, or by confluence
as much as 3 mm. The color is a waxy yellow in young speci-
mens, verging to olivaceous or brownish when dried.. The
fructification is almost disk-shaped, being considerably flattened,
and the upper surface may be smooth or thrown into serpentine
folds. The hymenium is composed of the usual bifurcate
basidia (TEXT FIG. 7), each with two spores. The spores (TEXT
FIG. 5) are short-cylindric though often narrowed at one end,
hyaline, one-celled, and measure 10-12 x 3-4 p. The hyphae in
Fics. 5-8. DACRYOMYCES HYALINA Quél.
5. Mature spores. 6. Hypha with clamp connection. 7. Mature
basidia. 8. Hypha with scabrous wall and clamp connection
the trama vary from 4 v. to 6 w in diameter and the walls of the
larger ones appear scabrous. Branching and anastomoses are
common, but cross walls are not abundant. Contrary to the
usual condition in the-Tremellales and the Dacryomycetales,
clamp connections are found on the hyphae, though not abundant.
On the older scabrous hyphae they are rather inconspicuous
(Text FIG. 8) but on the younger hyphae they are not difficult
to locate (TEXT FIG. 6).
Good figures of this plant were recently published by Lloyd
(Myc. Notes, No. 58, p. 828. 1919), who reports the species
from North Carolina. The name employed to designate the
species is open to criticism. The plants seem to be specifically
distinct from D. deliquescens, which they resemble microscopical-
ly as to spores and basidia, since the latter seems to lack the large
168 MYCOLOGICAL NOTES FOR 1920
scabrous-walled hyphae and the clamp connections of D. hyalinus.
I have not made, for D. deliquescens, an exhaustive search for
clamp connections, but have failed to find them in the mounts
I have made. Neither have I ever seen D. deliquescens assume
the dark colors on drying, characteristic of the present species.
6. STEREUM RADIATUM Peck
The fourth Pennsylvania collection of this species was made
in October 27, 1920, on coniferous boards in the greenhouse at
State College. A fifth collection, made in the Ottawa Valley,
Canada, by R. J. Blair in 1917, is in my herbarium. Three of
these collections were taken from structural coniferous timbers,
four of them being from 7suga canadensis. The specific iden-
tity of the substratum of the fifth collection was not determined,
except that it was recognized as a coniferous host. The first
collection, made here in 1915, was determined by Dr. Burt.
The species is one of the easiest of the stereums to recognize.
The plants are almost entirely resupinate and the hymenium
is in all collections a uniform cinnamon brown or Sudan brown,
except that where the fresh specimens are wounded on the
hymenium they immediately turn black and remain so on dry-
ing. The hymenial surface is radiately lined with obtuse ribs
in all cases. The pileus where developed is black on top in
dried specimens and usually decidedly rugose.
Spores have not been certainly seen; in one specimen, how-
ever, small rounded hyaline spore-like bodies, 4—5 » in diameter,
were present on the hymenium but were not seen attached to
basidia. No cystidia are present. Sections of the hymenium
sometimes become pea green in KOH solution.
7. MERULIUS FUGAX Fr.
This rare and interesting species was collected on a conif-
erous board in the greenhouse at the Pennsylvania State College,
on October 26, 1920. It is a bright colored species, my field
notes recording a salmon buff color, varying to pinkish cinnamon
for the hymenial surface, and with the margin considerably
lighter. The specimen was rather young and the gyrose-
plicate hymenium was developed only in the center of the fruc-
tification. The plant is separable from the substratum when
fresh but as soon as it becomes dry it adheres tightly, although
the fructification is quite membranous in that condition. Micro-
scopic examination of sections through it shows the character-
MYCOLOGICAL NOTES FOR 1920 169
istic features of the species, which are the oblong-ellipsoidal to
subglobose spores and the large conspicuously incrusted hyphae
of the subhymenium next the substratum. These hyphae are
4—6 vy. in diameter and possess both cross walls and clamps. The
spores under the microscope are all but hyaline, although the
species is included among the Leptospori by Dr. Burt, who
states that they sometimes appear slightly yellowish. No
cystidia are present.
Merulius subaurantiacus Peck is recorded as a synonym by
Dr. Burt, and my sections of the type of that species show struc-
tures identical with those of M. fugax, except that no incrusted
hyphae are present next the substratum. Apparently no other
species described from America has similar spores.
8. SOLENIA FASCICULATA (Pers.) Fr.
Plants rather crowded on eigen wood, Spoppen
of isolated cylinders not more
thick, whitish or slightly mens externally compactly tomen-
tose; hymenium lining the hollow ae of clavate basidia
without cystidia; spores not seen attached to basidia but free
floating, hyaline, thin-walled, and 5-7 wu in diameter.
On coniferous wood. Reitz Gap, ee County, Pennsyl-
vania, September 20, 1920, J. F.
According to C. G. Lloyd, who pete the determination,
S. villosa Fr. is probably the same species. These specimens
depart from the available descriptions of S. fasciculata, which
call for a plant 2-7 mm. high. The plants under the hand lens
(PLATE 9, FIG. 3), resemble a simple Mucronella in which genus,
however, the basidia are external.
g. POLYPORUS CAERULOPORUS Peck
Collected on the banks of Stone Creek, Huntingdon County,
Pennsylvania, July 9, 1920.
A rare species, listed by Murrill as occurring in New England
and New York. The uniform pale indigv-blue color of cap,
tubes, and stem is unique among the pore fungi. A single
“| sapeeaalan collected, from which the following notes were made:
Plant stipitate, more or less uniformly Bees indigo- blue;
pileus wir fe orbicular, 4.5 X 3.5 X 0.3-0.7 cm., smoky indigo,
glabrous, dry, slightly rugose; margin lobed, " deflene: context
white, I-5 mm. thick, taste mild; tubes about 2 mm. lon ng,
‘pale blue wit thin, the mouths pale neeetine angular, thin-
walled, entire, averaging two per mm., decurrent on the stem,
170 MYCOLOGICAL NOTES FOR 1920
not separable from the context; stem excentric, nearly lateral,
deep indigo-blue, pallid within, glabrous, 2.5 cm. long, 0.5 cm.
ain spores snares. hyaline, 4-5 uw in diameter; cystidia
ene dried plant is dark bluish black on the pileus and stem,
and the hymenium has become brown.
10. Polyporus compactus sp. nov.
Plants resupinate to narrowly reflexed or cushion-shaped,
the pileus not more than I cm. broad, and 1.5 cm. thick, in re-
supinate condition 3-8 mm. thick, white or gray in color, cor
watery when fresh, firm and compact when dry, glabrous or
practically so, the margin rounded and obtuse; context light
wood color, friable but compact when dry, taste mild; tubes
oblique, less than 2 mm. long, often lacking or poorly erckiged.
the mouths subcircular, averaging three or four per mm., white;
dissepiments rather thick and rigid; basidiospores ellipsoid or
ovoid, often with a ee truncate aare ome 7.5-9 X 4.5-6
ae hlamydospores (?) abundant on the hymenium and the
hyphae of the context, subglobose to ‘rasta slitpeoid, colorless
or paar gree nish or yellowish under the microscope, 7.5-9
X 6-7.5 u 6-7.5 uw in diameter; basidia large, pyriform, 9-10
yin panies cystidia mied with the basidia as ee pia narrow,
flask-shaped organs 3-4 & in diameter; hyphae of two kinds:
(a) large hyaline, unstaining hyphae, often Oe piled, nes
deeply staining: and much branched hyphae ks at least
some of the chisniydoaporce with a few cross walls and clamps,
diameter 2-3.5 u. [PLATE 9, FIGS. 5 and 6; TEXT FIGS. 9-14.
_ On dead standing trunks or on stumps of Quercus, especially
Q. alba.
Specimens are at hand as follows, the first mentioned being
designated as the type:
New Brunswick, New Jersey, on dead standing Quercus alba,
August 25, 1920, Overholts & West, No. 7873; Lamar, Clinton
County, Pennsylvania, on dead standing Quercus alba, July 26,
1920, Overholts Herb. No. 7525; State College, Pennsylvania, on
oak stump, October, 1920, No. 7173; New Brunswick, New
Jersey, on dead Quercus alba, August 21, 1918, E. West 343,
No. 7584; New Brunswick, New Jersey, on dead standing oak
trunks, June 16, 1919, E. West 414, No. 7526.
After this manuscript was submitted for publication, the
specimens that had been previously designated as the types and
from which the photographs for this article were made, were
Fetes ee eae ee
ies 6b 5 ee aN
MYCOLOGICAL NOTES FOR 1920 ‘I7I
destroyed by a fice that burned the photographic studio where
the collection was kept at the time. This has necessitated
designating my herbarium No. 7873, as the type of the species
rather than No. 7525, from which the photographs were made,
and which is therefore no longer in existence. This will explain
any discrepancy that might appear in the failure, in later years,
to find a collection in my herbarium with the exact aspect
‘presented in the photographs.
Fics. 9-14. PoLyporus COMPACTUS Overholts
9. Small portion of the hymenium showing basidia, cystidia, and attached
spores. 10. Mature basidiospores. 11. Branched hypha. 12. Chlamydo-
spores. 13. Production of chlamydospores. 14. Hypha with clamp con-
Of the collections here cited, only the first and the last named
show basidia and basidiospores, while all bear the chlamydospore
condition.
The real status of the so-called chlamydospore condition has
not been determined, as no cultures have been attempted. That
the knob-like growths as shown in PLATE 9 involve this species
can scarcely be questioned. It has been suggested that these
growths may represent a diseased condition and that the bodies
here designated as chlamydospores may be in reality the spores
172 ; MYCOLOGICAL NOTES FOR 1920
of the parasite. They are not produced in chains but formed
singly as the termini of small hyphae deeply staining with eosin.
This fungus has been under careful observation for sev-
eral years, but only the chlamydospore condition was known
until 1920. A well-developed pileus is never present. In some
respects the species shows resemblance to P. Ellisianus Murrill,
which however is found on coniferous wood and has entirely
different spores. The cushion-shaped or nodular type of sporo-
phore, bearing only the chlamydospore condition, has the context
more or less zoned and with a disagreeable odor as of strong lard.
11. Fomes BAKERI Murrill
I found this species to be quite common, by far the most
common species of Fomes, in the vicinity of Kingston, New Jersey,
where I observed it in company with Mr. E. West in August, 1920.
Its only host in that locality is the river birch, Betula nigra,
which is its host also in the Mississippi Valley. The species
was described and illustrated in my notes for 1919, where
reference was made to a collection said to have been taken from
Betula lutea near Philadelphia in 1915 by Rhoads. In all prob-
ability the host for this collection was B. nigra, as B. lutea is
not found in that locality, according to Dr. Rhoads.
In connection with this species it was of interest to find
recently in the Schweinitz Herbarium at Philadelphia a specimen
of Fomes under the name of Polyporus annosus Fr. (Syn. N. Am.
Fungi No. 401) that is undoubtedly F. Bakeri, although it seems
not to have been recognized by those who have worked over
the Schweinitzian material in recent years. Lloyd once referred
this specimen to F. rimosus, noting, however, its similarity to
F. robustus Karst., which he has subsequently regarded as identical
with F. Bakeri. This Schweinitzian collection bears no date but
was made certainly prior to 1831 and was listed by Schweinitz
as occurring on Betula.
12. PHALLOGASTER SACCATUS Morgan
A rare plant, placed in the Phallaceae by some largely on the
foetid nature of the internal deliquescent gleba, but in appearance
more allied to the Lycoperdaceae. Two specimens were collect-
ed at Westport, Pennsylvania, August 11, 1920. Excellent
illustrations, by which the plant may readily be known, have
been published by Lloyd.
MYCOLOGICAL NOTES FOR 1920 173
Explanation of plate 9
Fic. 1, Enlarged Lig ahs 3 of a small area of the resin exudate on which
Zythia resinae (Ehrenb.) Karst. and Biatorella resinae (Fr.) Mudd. grow side
by side. The elongate bodies in the left hand corner are the pycnidia of the
Zythia, while the larger disk-shaped bodies to the right are the apothecia
of the Biatorella. Photographed with Micro-Tessar 10 X lens, by E. T. Kirk.
1G. 2, Resin exudate inhabited by the pitch midge and bearing the
fruiting Godin of Zythia resinae and Biatorella resinae, X 1. Photo by the
writer
Fic. 3, Solenia fasciculata (Pers.) Fr., showing the isolated white-tipped ~
cylinders in which the hymenium is produced. Photographed with Micro-
Tessar 10 X lens, by C. B. Neblette.
Fic. 4, Pilacre Petersii B. & Br., natural size, Photo by the writer.
Fic. 5, Rounded abnormal growth of Polyporus campactus Overholts,
as usually found on vertical yauare and bearing only the chlamydospore
condition, X 1. Photo by the wr
Fic. 6, Poroid form of Rania compactus, apa the largely resupi-
nate manner of growth, X I. Photo by C. B. Neblet
i}
BULL. 7
ORREY CLUB VOLUME 49, PLATE 9
1,2. ZYTHIA RESINAE (EHRENB.) ere anp BIATORELLA RESINAE (FR.)
x cared Rhos oRave FR.
4. PILACRE PETERSII :
5, 6. baie es ait OVERHOLTS
Unreported plants from Glacier National Park
PAuL W. GRAFF
Our national parks are each season attracting a greater
number of vacationists drawn to them by the beauties of nature,
some only by the great attractions of mountain scenery but
many by all the varying phases of nature to be enjoyed in these
national playgrounds. To the person of botanical or zoological
interests Glacier Park is one of the most attractive. Its high
rugged peaks, deep valleys, numerous glaciers and lakes, streams
and cascades offer a great amount of variation. The Continen-
tal Divide passes through the Park with the result that the
drainage of the western portion is into the Columbia River,
while the northeastern part slopes toward the Hudson Bay, and
the eastern into the Missouri River drainage. A greater amount
of variation will be found here and with less sophistication than
in any other of our park areas.
In the herbarium of the University of Montana Biological
Station at Yellow Bay are a number of plants which have been
collected at the Park. Among these are a number which have
not as yet been reported from that region even in Standley’s
recent publication.* The present list is offered as an addition
and slight contribution to the ] ledge of the flora of this region.
Of the thirty-two species here listed twelve have not before been
reported from the state of Montana. The names of these
twelve are preceded by a dagger (f). Two of these, Carex Hele-
onastes Ehrh., and Salix reticulata L., have not, so far as I am
aware, been reported from the United States. They are Canadian
species which have entered this country in the high mountains
of this region, both being extreme northern forms.
{7 1. DANTHONIA PINETORUM Piper, Fl. Northw. ne IQI5.
D. spicata var pinetorum Piper, Erythea 7: 1899. D.
thermalis Scribn. U. S. Dept. Agr. Div. Aue Circ. 30:
1901.
Mineral Park, August 10, 1910, M. E. Jones, at 1800 meters
elevation.
This is a northwest coast plant which has been reported as
*Flora of Glacier National Park. Contr. U. S. Nat. Herb. 22: 235-438.
pl. 33-52. 1921.
175
176 UNREPORTED PLANTS FROM GLACIER NATIONAL PARK
being found to the eastward in Wyoming and Idaho, but not as
yet in the state of Montana.
2. Poa Cusick Vasey, Contr. U. S. Nat. Herb. 1: 271. 1893.
Mineral Park, August 8, 1910, M. E. Jones, at 2400 meters
elevation.
Reported from British Columbia, south to Oregon and east-
ward through Idaho and western Montana
3. PoA NEVADENSIS Vasey, Bull. Torrey Club 10:66. 1883.
Greenwood’s Camp, August 15; 1910, M. E. Jones, at an
elevation of 1350 meters
Reported from British Columbia southward to hci
and eastward to North Dakota and Colorado.
4. Bromus Eximius (Shear) Piper, Contr. U.S. Nat. Herb. 11:
143. 1906. B. vulgaris var. eximius Shear, U. S. Dept.
Agr. Div. Agrost. Bull. 23: 44. 1900
Greenwood’s Camp, August 15, ie: M. E. Jones, at 1350
meters elevation.
This is primarily a Pacific coast plant, being found from
British Columbia southward to California. To the eastward
of this range it is found in the states of Montana and Wyoming.
5. ERIOPHORUM POLYSTACHYON L. Sp. Pl. 52. 1753.
Mineral Park, August 12, 1910, M. E. Jones, at 1800 meters
elevation. :
This is primarily a Canadian species which has been found
to the southward in the Rockiesand in the North AtlanticStates.
16. CAREX AQuATILIS Wahl. Kongl. Vet. Akad. Handl. II. 24:
165.
Bowman Tate, August 19, 1910, M. E. Jones, at 1200 meters
elevation; Grinnell Glacier, August, 1911, M. J. Elrod, at 2100
meters elevation.
A Pacific coast and Canadian species, also reported southward
in the Atlantic States.
7- CAREX DEWEYANA Schwein. Ann. Lyc. New York 1:65. 1824.
Waterton Lake, August 17, 1910, M. E. Jones, at 1200
meters elevation.
This is a northern species with a range extending southward
in both the Appalachians and Rockies. It has been reported from
Montana, but not from the Park region.
UNREPORTED PLANTS FROM GLACIER NATIONAL PARK 177
8. CAREX FESTIVA Dewey, Am. Jour. Sci. 29: 246. 1835.
Sperry Glacier, August 27, 1909, M. E. Jones, at 2250
meters elevation; Grinnell Lake, August 10, 1911, M. J. Elrod,
at 2100 meters elevation.
Found from California to the Arctic regions on the coast,
and in the Rocky Mountains
79. CAREX FOENEA Willd. Enum. 957. 1809.
Waterton Lake, August 17, 1910, M. E. Jones, at an eleva-
tion of 1200 meters
This is primarily a Canadian species which has crept down
into our range. Its range is from New Brunswick to Penn-
sylvania, thence northwestward to Minnesota, British Columbia,
and the Yukon
10. CAREX ee, Gay, Ann. Sci. Nat. II,11:191. 1839.
C. caespitosa Gooden., Trans. Linn. Soc. 2: 195. pl. 22, 1794.
Not Linn. 1753.
Lake Mc Donald, August 2, 1910. M.E. Jones, at 900 meters
elevation. Mineral Park, August 10, 1910, M. E. Jones, at
an altitude of 1800 meters.
Found from Newfoundland and Pennsylvania westward to
Alaska and Colorado.
7 11. CarEx HeteonastEs Ehrh.; L. f. Suppl. 414. 1781.
Sperry Glacier, August 27, 1909, M. E. Jones, at an altitude
of 2250 meters, growing in a damp locality among moss.
While reported from the Upper Temperate and Alpine
Life Zones in Canada, from Ontario and the Hudson Bay region
westward to British Columbia, this species does not seem to
have been reported from this side the “Border.” It is essentially
a northern plant which has found its way into the United States
in the high altitudes of the Park.
f12. CAREX HORMATHODES Fernald, Rhodora 8: 165. 1906.
Blackfoot Glacier, August 30, 1909, M. E. Jones, at 2100
meters elevation.
Reported previously from British Columbia, but not from
the northwestern United States.
713. CAREX MONILE Tuckerm. Enum. Meth. 20. 1843.
Grinnell Lake, August 20, IQII, M. J. Elrod, at 1500 meters
elevation.
This differs from C. vesicaria L., with which it is sometimes
confused, in the possession of three stigmas rather than two
and in the presence of more than one staminate spike. The
variety colorata Bailey has been reported from Yellowstone
178 UNREPORTED PLANTS FROM GLACIER NATIONAL PARK
Park, but the Gace form of the species has not been reported
from Montan
Reported ain of our range in Wyoming, aie across the
United States. It is also found along the Pacific coast from
California to Alaska.
+14. CAREX POLYGAMA Schkuhr, Riedgr. 1: 84. 1801.
Waterton Lake, August 17, 1910, M. E. Jones, at an elevation
of 1200 meters.
Reported well to the northward on both coasts, and across
the United States from California through Utah, Colorado, and
Missouri to the Atlantic coast. It is a southern species which
has found a congenial habitat to the northward on both sea-
boards.
115. CAREX SAxXATILIS Le Sp: Pl-o76.. 1753.
’ Bowman Lake, August I9, 1910, M. E. Jones, at 1200 meters
elevation.
C. saxatilis is anarctic species found across North America
from Labrador to Alaska. It has been reported as entering
the United States in the mountainous portion of Maine, but
has not been reported from the western states except in the
high mountains of Colorado. It is essentially an arctic plant,
and is found also in the extreme northern portion of Europe and
Asia.
16. SALIX ORBICULARIS Anders.; De Candolle, Prodromus 16?:
1868.
Flat-top Mountain, August, 1911, M. J. Elrod, at 1g00 meters
elevation; Iceberg Lake, August 18, 1911, M. J. Elrod, at 1800
meters elevation.
Reported previously from the Sperry Glacier region under
the name S. reticulata
This is an extreme northwestern species found from Alaska
to Hudson Bay, and southward into the United States in the
Rocky Mountains.
t17. SALIX RETICULATA L. Sp. Pl. 1018. 1753. Not Porter &
Coulter.
Blackfoot Glacier, Nee a: a0 1909, M. E. Jones, at an ele-.
vation of 2100 meter
This species has’ Bach asic previously as purely Cana-
dian. It has been reported from the Canadian Rockies and
northward to Alaska and Hudson Bay. Salix saximontana
Rydb. has been reported from this region with S. reticulata L.
UNREPORTED PLANTS FROM GLACIER NATIONAL PARK 179
as a synonym, an error for S. reticulata Porter & Coulter, a
quite distinct species.
18. ERIOGONUM OVALIFOLIUM Nutt. Jour. Philad. Acad. 7:
50. pl. 8. 1834.
Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580 meters
elevation.
A western species found from New Mexico northward through
California and Colorado to British Columbia.
19. POLYGONUM BISTORTOIDES Pursh, var. LINEARIFOLIUM
(Wats.) Small, Bull. Torrey Club 19: 252. 1892.
Iceberg Lake, August 18, 1911, M. J. Elrod, at an elevation
of 1800 meters
Found iar New Mexico northward within the Rocky
Mountains through Canada to the Arctic regions.
20. SILENE Dovuciasi Hook. Fl. Bor. Am. 1: 88. 1840.
Sperry Glacier, August 27, 1909, M. E. Jones, at 2250 meters
elevation.
Rather a common plant in the Rockies, and on the coast
from Washington to California.
21. ARENARIA SUBCONGESTA Wats.) Rydb., Bull. Torrey Club
24: 244. 1897.
Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580 meters
elevation.
This species is common in the Rocky Mountain region. It
seems to be the same as A. lithophila Rydb. and A. Burket
Howell and should possibly be considered only a variety of
A. congesta Nutt.
22. ARAGALLUS MOoNTICOLA (Gray) Greene, Pittonia 3: 212.
1897.
Mineral Park, August 8, 1910, M. E. Jones, at 2100 meters
elevation; Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580
meters elevation.
Not uncommon in the mountains from Myoung northward.
23. POLEMONIUM CONFERTUM Gray, Proc. Acad. Philadelphia
1863:'73. 1864.
Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580 meters
elevation.
A high elevation plant found in the mountains of Utah,
Colitade, Wyoming, and Montana; ‘occasionally in northern
New Mexico and California.
180 UNREPORTED PLANTS FROM GLACIER NATIONAL PARK
24. PHACELIA FRANKLINII (R. Br.) Gray, Manual, Ed. 2, 329.
1856.
Mineral Park, August 8, 1910, M. E. Jones, at 2400 meters
elevation; Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580
meters elevation.
From southern Idaho, Wyoming, and Michigan northward
to Hudson Bay and Alaska.
7 25. LAppuLA CILIATA (Dougl.) Greene, Pittonia 2: 182. 1891.
Mineral Park, August 12, 1910, M. E. Jones, at 1800 meters
elevation.
Reported from the states of Washington and Idaho.
26. PENTSTEMON Menziestt Hook. FI. Bor. Amer. 2:98. 1840.
P. fruticosus (Pursh) Greene, Pittonia 2: 239. 1892.
Sperry Glacier, August 27, 1909, M. E. Jones, at 2250 meters
elevation; Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580
meters elevation.
Found from California and Colorado northward to British
Columbia and Alberta. :
} 27. VERonIcA Cusick Gray, Syn. Flor. 2: 288. 1878.
Mineral Park, August 9, 1910, M. E. Jones, at 1800 meters
elevation.
Previously reported only from the high mountain regions of
the three states, Washington, Oregon, and Idaho.
28. VALERIANA OCCIDENTALIS Heller, Bull. Torrey Club 25:
269. 1898. V. sylvatica Banks; Richards, Frankl. Journ.
Ed. 2, Appendix 2. 1823. NotSchmidt, 1795. V. wyoming-
ensis E. Nels. Erythea 7: 167. 1899.
Swift-Current Pass, August 11, 1910, M. E. Jones, at 2100
meters elevation.
Found from Labrador and Quebec across Canada to British
Columbia, and southward through Washington and Montana
to Colorado and Utah.
29. SOLIDAGO Pursuti Porter, Bull. Torrey Club 21: 311. 1894.
S. humilis Pursh, Fl. Am. Sept. 543. 1814. Not Mill.
Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580 meters
elevation; Iceberg Lake, August 18, 1911, M. J. Elrod, at 1800
meters elevation.
Found in the Rocky Mountains from New Mexico to British
Columbia and northward.
UNREPORTED PLANTS FROM GLACIER NATIONAL PARK I81
30. ERIGERON GRANDIFLORUS Hook. Fl. Bor. Am. 2: 18. pl. 123.
1840.
Sperry Glacier, August 27, 1909, M. E. Jones, at an eleva-
tion of 2250 meters
Reported from Colorado to the extreme north; more es-
pecially in the high elevations of the Canadian Rockies.
7 31. ANTENNARIA ALPINA (L.) Gaertn. Fr. et Sem.2:410. 1791.
Gnaphalium alpinum L. Sp. Pl. 856. 1753. ;
This is essentially a Canadian species found across the con-
tinent from Labrador to Alaska, but extending southward into
the United States in the Rockies and Sierras.
33. SENECIO. AUREUS. 1... Sp. Fi. 870... 4763.
Brown’s Pass, August 13, 1911, M. J. Elrod, at 2580 meters
elevation; Iceberg Lake, August 18, 1911,.M. J. Elrod, at 1800
meters elevation.
This species is close to Rydberg’s S. crocatus of Colorado,
Wyoming, and the northern Rockies. Its range is from Alaska
to California, and across the continent.
UNIVERSITY OF MONTANA BIOLOGICAL STATION,
YELLOw Bay, MONTANA
peter
eyuhave
ne
.
Oe Sa
Two new plants from western Colorado
_ Gro. E. OSTERHOUT
’ Nuttallia marginata sp. nov.
Seemingly a biennial, the stem 3 dm. or more high, white,
smooth below, roughish pubescent above: ‘is branches alternate,
the lower ones elongate; the lower leaves oblanceolate, sinuate
dentate, 6-8 c . wide, ow base,
the upper leaves smaller, coarsely dentate, sessile, all of them
roughish with a barbed pubescence; the flowe llow, numerous in
a cyme of alternate branches, the whole flower—capsule and petals
—2 cm. long, py subtended by a narrow entire bract, five
outer petals 13 mm. Jong, the claws short and broad, the upper
portion ovate, point me the outside oe nve inner ra
smaller, spatulate; the capsule 8-11 mm. long, rounded a
oe
pubescent like the capsule; the style mm. long, =o
at the tip; seeds ey slightly over 2 mm. long, I.5 mm. wide,
the margin very narrow.
The only ae where I have collected this Nuitallia is
De Beque, in Mesa County, on the hills north of the town.
The type specimens (No. 5842) were collected August 22, 1918.
The species begins to blossom near the first of June. It is peculiar
in its narrowly margined seeds and pubescent petals.
‘Acrolasia humilis sp. nov.
An 1-1.5 dm. high, the stem white, smooth below,
debian: with slender barbed hairs above, the first branches
from near the base, but the branches few an d distant; the low-
est ‘eee spatulate, on short petioles, 3-5 cm. long, 5 mm. or
glabrous above or nearly so; the first flowers sessile in the axil
of the branches, the later ones clustered and sessile at the ends
of the branches, the petals five, yellow, broadly obovate, veined,
2.5-3 mm. long, the calyx lobes linear, 2 mm. gad the stamens
ten to fifteen, the Maras slender, the anthers ior the
capsule becoming about I cm. long, 3 mm. wide, rough wit
small barbed pubescence, ‘andl longer slender hairs not barbed;
the seeds prismatic and minutely muricate
The type specimens were collected at and Junction, on
the north side, near the coal mines, May 30, 1921, No. 6086.
I also collected it in the vicinity of Eckert, in Delta County.
On account of the entire leaves it rather resembles Acrolasia
183
184 Two NEW PLANTS FROM WESTERN COLORADO
latifolia Rydb. or A. dispersa (Wats.) Rydb., but it really be-
longs with A. albicaulis (Dougl.) Rydb. The seeds are similar
to the latter, and are in the capsule in three rows, not in a single
row and end to end as they are in A. latifolia and A. dispersa.
WINDSOR, COLORADO
INDEX TO AMERICAN BOTANICAL LITERATURE
1921
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or ae upon American material; the word
America being used in the broadest
Reviews, and papers that relate etnctinfiel to foresty, agriculture, horti-
oulene manufactured products of vegetable origin, or ls boratory methods
f bacteriology.
An occasional sista is tiade i in favor of some paper appearing in an Ameri-
can 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 three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Gardner, M. W., & Kendrick, J. B. Turnip mosaic. Jour.
Agr. Research: 22: 123. pl 20 15 O 1921.
Garrett, A. O. Some introduced plants of Utah. Torreya 21:
76-79. O 1921.
Gatin, C. L. (Premiére contribution a l’étude) De l’embryon
et de la germination des Aracées. Ann. Sci. Nat. Bot. X.3:
145-180. pl. 1-10. -N Ig2t.
Gericke, W. F. Root development of wheat seedlings. Bot.
Gaz. 32: 404-406. f. |. 15 D 1921.
Gleason, H. A. Botanizing in British “Galina: Jour. New
ork Bot. Gard. 22: 161-168. S 1921
Gleason, H. A. A rearrangement of the Bolivian species of
Centropogon and ees mpylus. Bull. Torrey Club 48:
189-201. 26S 19
Includes 2 new species 2 peace
Goldring, W. Annual rings of growth in Carboniferous wood.
ot. Gaz. 72: 326-330. pl. 14. 15 N. 1921.
Goss, R. W. Temperature and humidity studies of some
Fusaria rots of the Irish potato (Solanum tuberosum). Jour.
Agr. Research 22: 65-79. pl. 10. 11. 8 O 1921
185
186 INDEX TO AMERICAN BOTANICAL LITERATURE
Griffee, F. Comparative vigor of F; wheat crosses and their
parents. Jour. Azr. Research 22: 53-63. 8 O 1921.
Grimes, E. J. A new station for Pogonia affinis. Rhodora
23: 195-197. 15 D 1921.
Guppy, H.B. America’s contribution to the story of the plant-
world. Jour. Ecol. 9: go-94. S Ig2!.
Harlan, H. V., & Pope, M.N. Ash content of the awn, rachis,
palea, and kernel of barley during growth and maturation.
Jour. Agr. Research 22: 433-449. f.-15. 19 N 1921.
Harms, H. Drei neue Leguminosen aus Venezuela. Notizbl.
Bot. Gart Berlin-Dahlem 8: 51, 52. 1S 1921.
Species of Calliandra (1), Pithecolobium (1), and Piptadenia (1).
Harms, H. Kakteen als Uric kee Monatschr. Kakteenk.
31: 184-186. D 19
Harms, H. Kakteen und Sigillarien. Monatschr. Kakteenk.
31: 145-148. O I921.
Harper, R. M. Cape Cod vegetation. Torreya 21: 91-098.
D 1921. [Illust.]
Harrington, G. T. Optimum temperatures for flower seed ger- .
mination. Bot. Gaz. 72: 337-358. f. 1-10. 15. D Ig2l.
ete J. A., Sinnott, E. W., Pennypacker, J. Y., & Durham,
The vascular anatomy of hemitrimerous seedlings of
este vulgaris. Am. Jour. Bot. 8:375-381. 14 N 1921.
Harris, J. A., Sinnott, E. W., Pennypacker, J. Y., & Durham,
G. The interrelationship of the number of the two
types of vascular bundles in the transition zone of the
axis of Phaseolus vulgaris. Am. Jour. Bot. 8: 425-432.
19 D 1921.
Harshberger, J. W. The artistic anatomy of trees. Nat. Hist.
21: 387-397.. O 1921: [Blust.]
Harter, L. L., & Weimer, J. L. A comparison of the pectinase
produced by different species of Rhizopus. Jour. Agr. Re-
search 22: 371-377. f:4, 2. 12 W 192r.
Harter, L. L., Weimer, J. L., & Lauritzen, J. I. The decay of
sweet potatoes (Ipomoea batatas) produced by different
species of Rhizopus. Phytopathology 11: 279-284. 15 N
1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 187
Hastings, G. T. The forest flora of Grassy Sprain Ridge [New
York). Torreya 21: 73-76. O 1921.
Hastings, G. T. The boy scouts and conservation of wild
flowers. Torreya 21: 83, 84. O 1921
Hauman, L. Deux graminées géantes de la flore argentine.
Physis 5: 52-56. 31 O 1921.
Includes Sporobolus maximus, sp. nov.
Hauman, L., & Parodi, L. R. Los parasitos vegetales de las
plantas cultivadas en la Republica Argentina. Revista
Facult. Agron. y Veter. 3: 227-274. f. 1-4. D 1921.
Hawkins, L. A. A physiological study of grapefruit ripening
and storage. Jour. Agr. Research 22: 263-279. f. 1. 29 O
1921.
Heald, F.D. The relation of spore load to the per cent of stink-
ing smut appearing in the crop. Phytopathology 11: 269-
278. 15 N 1921.
Heller, H. H. Phylogenetic position of the bacteria. Bot.
Gaz. 32: 390-396. 15 D Ig21
Heller, H. H. Suggestions prere a rational basis for the
classification of the anaerobic bacteria. Jour. Bact. 6:
521-553. °‘N 1921:
Studies in pathogenic anaerobes IV.
Hoerner, G. R. Miscellaneous studies on the crown rust of
oats. Am. Jour. Bot. 8: 452-457. pl. 24. 19 D Ig2t.
Holm, T. Morphological study oi Carya alba and Juglans
nigra. Bot. Gaz. 32: 375—380. D/. 75, 70 + Ff. i. 15 -D
1921.
Holm, T. Recent botanical publications from the United
States National Museum. Am. Midl. Nat. 7: 165-180.
N 1921.
Considerable original matter contained in a review.
Hopkins, E. F. Studies on the Cercospora leaf spot of bur
clover. Phytopathology 11: 311-318. pl. 3, 14 + f. 8, 9.
9 D 1921.
Howard, G. E. Extraction and separation of the pigments of
Nereocystis Luetkeana. Publ. Puget Sound Biol. Sta.
79-91. pl. 14. 1 O 1921.
188 INDEX TO AMERICAN BOTANICAL LITERATURE
Howe, C. G. Pectic material in root hairs. Bot. Gaz. 72:
313-320. 15 N I92I.
Hubert, E. E. Notes on sap stain fungi. Phytopathology 11:
214-224. pl. 7 + f. 1-4. 6 O I92I.
Huhnholz, P. Echinocactus Anisitsii KK. Sch. Monatschr.
Kakteenk. 31: 150, 151. .O 1921.
Mick, J. S. The American walnuts. Am. For. 27: 699-704.
N 1921. [Illust.]
Inman, O. L. Comparative studies on respiration XX. The
cause of partial recovery. peer Gen. Physiol. 4: 171-175.
Fock, 25 -2O WN 192k.
Jackson, H. S., & Mains, E. B. Aecial stage of the orange
leafrust of wheat, Puccinia triticina Eriks. Jour. Agr.
Research 22: 151-172. pl. 21 +f. 2. 15 O 1921.
Jeffrey, E. C. The ar distribution of hybrids.
Science II. 54: 517. 25 N1
Johnson, D. S. Invasion of o soil in the tropics. Bot.
sag, Fe: 405-312; 7. 7, 2.°..15 1921.
Johnson, D. S. Polypodium vulgare as an epiphyte. Bot.
Gaz. 72: 237-244. f. 1-3. 15 O 1921.
Jones, F. R., & Tisdale, W.B. Effect of soil temperature upon
the development of nodules on the roots of certain legumes.
Jour. Agr. Research 22: 17-31. pl. 1-3 + f. 1-4. 1O 1921.
Jones, L. R., & Doolittle, S.P. Angular leaf-spot of cucumber.
Phytopathology 11: 297, 298. 15 N 1921.
Jones, L. R., & Keitt,G.W. Eugene Washburn Roark. Phyto-
pathology 11: 327. 9 D 1gat.
Kempton, J. H. Heritable characters of maize. III. Bra-
chytic culms. V. Adherence. Jour. Heredity 11: IfI-
115. f. 12. 6 My 1920; 11: 317-322. f. 16-19. 23 Mr 1921.
Kempton, J. H. Heritable characters of maize. VIII. White
sheathes. Jour. Heredity 12: 224-226. f. 18. 23 S 1921.
Kendrick, J. B., & Gardner, M. W. Seed transmission of soy-
bean hariacal blight. Phytopathology 11: 340-342- pl.
17; 9 D i921.
Vol. 49 7 No. 7
BULLETIN
OF THE
TORREY BOTANICAL CLUB
JULY, 1922
New species of Uredineae—XIV*
JosEPH CHARLES ARTHUR
In the tenth article in this series of new species an error
was made in the determination of the host for Uromyces fuscatus
(Bull. Torrey Club 45: 142. 1918), which should be Rumex
paucifolius Nutt., and not ‘Polygonum alpinum All.”’ as
printed. It was due to the persistent efforts of Mr, A. O. Garrett,
who first thought the host of the collection which he made was
Pentstemon (see Fungi Utahenses 244), that the facts in the case
were finally established.. In July, 1920, Mr. Garrett visited
the Utah locality at Gogorza, where he first collected the rust
in June, 1915, and secured a flowering stalk of the host, showing
it to be Rumex paucifolius. Upon my presenting the situation to
Professor Aven Nelson, he looked up the field notes and the
herbarium specimens collected in Idaho at the same place and
time as the type of U. fuscatus and came to the conclusion that
the host of this collecticn also was R. paucifolius. Only these two
localities for the species are known up to the present time.
The correction of these errors was complicated by a similar
uncertainty and error in the host names for Puccinia Polygoni-
alpini Cruch. & Mayor (Dicaeoma Polygoni-alpini Arth. N. Am.
Flora 7: 385. 1920), for which only two collections were known
at that time, a third one having been added from Greenville,
California, since the printing of the Flora. Rumex paucifolius,
host of the Uromyces, can not be told from Polygonum alpinum,
host of the Puccinia, when the fruiting parts are absent, hence
the difficulties that have arisen.
* Contribution from the ou Department of Purdue University
Agricultural Experiment Stat
[The BuLLetin for June (49: 565-190: pl. 9) was issued July 8, 1922 ]
189
190 Artuur: NEw sPECIES oF UrEDINEAE—XIV
In the twelfth article of this series, under Puccinia offuscata
(Bull. Torrey Club 47: 469. 1920), the name Uredo Zorniae
Dietel (Hedwigia 38: 257. 1899) should have been given as a
synonym. This name was applied to Ule’s 2296 from Brazil.
Although pyenia have not been seen yet there is circumstantial
evidence to indicate that no aecia are produced, and therefore
the species is probably a Bullaria, and should be called B.
Zorniae (Dietel) comb. nov. It is a species of the New World,
to which the name Puccinia Zorniae McAlpine does not apply.
In the same article, under Pucciniastrum americanum(Bull.
Torrey Club 47: 468. 1920), Rubus occidentalis was erroneously
given as a host for this species. The error was called to my
attention by Dr. J. J. Davis of the University of Wisconsin,
and was confirmed by subsequent examination under the
microscope. The type of P. arcticum americanum Farl.is on Rubus
neglectus Peck, collected at Bussey, near Cambridge, Massachu-
setts. This collection is first mentioned in the original article
by Farlow (Rhodora 10: 16. 1908) as on R. occidentalis near
Cambridge, but later in the article is referred to as on R. neglectus,
Bussey, October, 1875. The correction in the name of the host
is credited to M. L. Fernald. I have been able to ascertain these
facts through the assistance of Dr. R. Thaxter, who has carefully
examined the original material in the Farlow Herbarium.
Melampsoropsis roanensis sp. nov.
redinia hypophyllous, in crowded groups of 2-8 2
somewhat discolored spots, roundish or ovoid, 0.3-0.8 mm.lon
prominent, early dehiscent bi a central or lateral slit, eee
a coh or : Sh soot covering of epi idermis, pulveru rent;
peridium delica
ellipsoid, 20-26 fer 32-40 p; ml colariess, 1.5-2. 5 yu. thick, nn
a ely verrucose on one side grading to coarsely verrucose
on opposite side with iobiewhiat Se hie cylindric tubercles,
2p.
Telia iii in irregular and a confluent groups
on reddish spots, roundish or ovoid, 0.5-1 . across, elevated,
prominent, waxy ’ ‘raed: tardily naked ; teliospores aang
16-20 p. broad, almost or quite - long, in a series 65-100
long; wall colorless, thin, I yu, smooth; contents eiesed,
On Rhododendron catawbiense Michs, Weacaoeas): summit of
Roan Mountain, Carter County, Tennessee, altitude 6400 feet,
July, 1887, II, R. Thaster.
On Rhododendron punctatum Andr., LeConte Mountain,
Sevier County, Tennessee, altitude 6600 feet, June 8, 1921,
ARTHUR: NEW SPECIES OF UREDINEAE—XIV IgI
ii, ITI, H. F. Bain, communicated by John A. Stevenson 5812
(type).
The urediniospores of this interesting species are much
larger and more prominently verrucose than those of the common
form in Europe, M. (Chrysomyxa) Rhododendri. This is the
first rust on Rhododendron seen from the eastern United States,
although the European species has been ascribed to this country
by a number of authors. Duggar says (Fungous Diseases 432.
1909), ‘‘In the United States this fungus is particularly common
in the mountains of the east, and southward as far as the southern
limits of the Appalachians.’’ The statement does not apply to
any true rust on Rhododendron. Roan Mountain and LeConte
Mountain are about 100 miles apart. The rust was reported
abundant on the summit of LeConte Mountain. Mycologists
who visit the summits of these mountains and those nearby
should be on the lookout for the aecia, which may be expected
to occur on species of Picea.
Cronartium stalactiforme Arthur & Kern, comb. nov.
Peridermium stalactiforme Arth. & Kern, Ball. Torrey Club
33: 419. 1906
This Cronartium belongs to a group of three species which
have been much confused, both in nomenclature and in structural
characters. All three species have uredinia and telia on Castilleja
and related genera of scrophulariaceous hosts, which are difficult
to distinguish except by means of cultures. The aecia of the
three species, however, occuring on pine trunks and branches,
can be identified with considerable certainty, even by their gross
appearance. The aecia of C. stalactiforme are confined to the bark
and do not produce woody galls. They often extend over con-
siderable areas, and cause little or no swelling. The individual
aecia are low and flattened, with a circumscissile dehiscence.
Cronartium filamentosum (Peck) Hedge. (Peridermium fila-
mentosum Peck, 1882) is similar to C. stalactiforme in the effect
upon the host, but the individual aecia are more elongated,
often becoming cylindric, and have noticeable filaments extend-
ing from top to bottom through the individual aecia. There are
only rudiments of such filaments in C. stalactiforme
Cronartium coleosporioides (Diet. & Holw.) Arth. (Perider-
mium Harknessit Moore, 1876) differs from the two preceding
species by producing a woody gall, often of considerable size and
192 Artuur: NEW SPECIES OF UREDINEAE—XIV
abruptly rising from the healthy tissue. The individual aecia are
low and coalesce into a more or less continuous mass, and are
entirely without internal filaments.
There has been much confusion regarding the application of
the name Peridermium Harknessti, owing in the first place to
the obscure way in which it was first published*, and in the
second place to the later inclusion of the Pacific Coast forms of
the very similar P. Cerebrum, whose uredinia and telia occur
on leaves of oak.
The name was announced at a meeting of the San Francisco
Microscopical Society in July, 1876, by J. P. Moore, who
and who read a letter from H. W. Harkness describing the
species in an informal but accurate maaner. The collection to
which the name was applied was made by Harkness and Moore on
May 26, 1876, at Colfax, California, which is in Placer county,
* Our attention was directed to the early history of this name by Mr. E.
Bethel in a letter dated January 22, 1922. Through the assistance of Mr.
Hepburn, of the Purdue University Library, and Professor W. C.
Blasdale, of the University of California, the following facts have been estab-
lished.
In the minute book of the San Francisco Microscopical Society this re-
Pe occurs: “July 20, 1876. Mr. J. P. Moore read a paper written by Dr.
There is also in the archives of the society a newspaper clipping
giving a more extended account of the matter as presented to the society.
This clipping, Professor Blasdale has ascertained, came from the issue of the
Daily Alta California of San Francisco, of Saturday, July 22, 1876, volume
28. It consists of the Harkness paper in full, and a paragraph stating how
Moore proposed the name
The society did not fee any printed account of its proceedings during
this period of its existence, but occasionally, as the subject matter warranted,
he secretary transmitted a more or less formal minute to the London Monthly
Microscopical Journal. In the issue of that journal for September 1, 1876,
an account of the meeting of July 20 is given in the same words used in the
newspaper clipping, with three additional paragraphs
at is referred to as a paper by Dr. Harkness is in the form of a letter
to the secretary of the society. It begins with the sentence: ‘I have today
forwarded for the Society’s Cabinet a specimen of Peridermium,” etc. This
type material was destroyed in the great San Francisco fire, being bulky and
in a box separate from other type collections, as Mr. Bethel writes, who has
consulted the herbarium of the society both before and since the fire, and as
stated by Meinecke (Phytopathology 10: 281. 1920), who also calls atten-
tion to the duplicate specimen at the New York Botanical Garden
ARTHUR: NEW SPECIES OF UREDINEAE—XIV 193
about fifty miles west of Lake Tahoe. Harkness says the fungus
“appears both on the limbs and trunks of young trees of the
variety Pinus ponderosa, generally forming a complete circle
around the trees, its sporidia appearing as a zone of bright
orange yellow. The spores first germinate beneath the cuticle,
which it destroys. Owing to the irritation of this presence an
abnormal thickening of the cambium is produced, which in
turn gives place to an excessive growth of woody’ fibre. This
process being repeated from time to time a large bulbous ex-
pansion is soon formed, so that as often occurs a stem of but an
inch in diameter is enlarged to that of four or five. Above this
bulb the further development of the stem is retarded, or arrested
altogether, its place being supplied by a dense tuft of minute
branches.’”’ Among other matters it is also stated that “‘in the
vicinity of Colfax the fungus appears to be limited to an area
of but a few acres in extent. Within that area, however, it is
destroying the young growth.”
The quotation is taken from the report by the secretary of
the San Francisco society sent to the Monthly Microscopical
Journal of London, and printed on page 164 in volume 16,
bearing the date of September 1, 1876. This periodical was
maintained by the Royal Microscopical Society.
The name, host and locality are cited in Harkness & Moore’s
Pacific Coast Fungi in 1880, still without confusion with any
other form. But when Harkness published technical descriptions
of “New species of Californian fungi” in the Bulletin of the
California Academy of Science in 1884, other forms and hosts
were incorporated, and from this time on the application of the
name has been various.
The chief source of confusion has been in failing to rec-
ognize the Pacific Coast form of Peridermium Cerebrum Peck,
which occurs along the coast of California, and is especially
abundant at Monterey on Pinus radiata (P. insignis). The
present writers feel especially culpable for this confusion, for
they placed P. Harknessii under P. Cerebrum as a synonym
in their paper on North American species of Peridermium on
pine (Mycologia 6: 133. 1914), and have been followed by
Meinecke, in his paper on Peridermium Harknessii and Cro-
nartium Quercuum (Phytopathology 6: 225-240. 1916), and
by others.
The present knowledge of the Castilleja rusts indicates that
the correct application of the names should be as follows:
194 Artuur: NEW SPECIES OF UREDINEAE—XIV
Cronartium filamentosum (Peck) Hedge. (Peridermium fila-
mentosum Peck, 1882);
Cronartium coleosporioides (Diet. & Holw.) Arth. (Uredo
coleosporioides Diet. & Holw. 1893);
ronartium stalactiforme Arth. & Kern (Peridermium stalactt-
forme Arth. & Kern, 1906).
Should it be deemed advisable to combine these three species
as subspecie’s, races, or forms, the name to be employed for the
aggregate species, using the oldest specific name, would be
Cronartium Harknessii (Moore) Meinecke, Phytopathology
10: 282. 1920
DIABOLE gen. nov.
Cycle of development includes only subcuticular telia, with
a possibility of p
Telia som peat indefinite in faites without paraphyses.
Teliospores free, usuall ired onac on icel, one-celled,
more or less adhering laterally ; wall adated, usually verrucose,
at least above; pore one, in upper part of ce
Diabole cubensis comb. nov.
Uromycladium (?) cubense Arth. Mem. Torrey Club 17: 119.
18.
The species occurs in Cuba on Mimosa pigra L. (M. asperata
L. ) and was at first assigned with many misgivings to the genus
Uromycladium. That connection has become more and more
uncertain, and it is now believed to constitute a new and very
distinctive genus.
Puccinia Plucheae (Sydow) comb. nov.
Uredo Plucheae Sydow, Ann. Myc. 1: 1903
Uredo biocellata Arth. Bull. Torrey Club 33: ayy. 1906.
Puccinia biocellata Vestergr. Micr. Rar. Sel. 1267. 1908.
The teliospores of this rust have been collected on two hosts,
Pluchea fastigiata Griseb. and P. Quitoc DC., in Argentina,
and issued in Vestergren’s exsiccati, Nos. 1267 and 1368. The
rust occurs on a number of species of Pluchea in Florida, Guat-
emala and the West Indes, but no teliospores have yet been
found in North America.
Uredo nominata sp. nov.
Il. Uredinia amphigcnous, grouped upon discolored areas,
5-10 mm. across, oblong or linear, 0.4-5 mm. long, somewhat
ARTHUR: NEW SPECIES OF UREDINEAE—XIV 195 .
tardily naked, pulverulent, light-brown, ruptured the Oe
evident; Ratan Wee Seti ellipsoid or obovoid, 18-23 23-32 wu;
all light cinnamon-brown, 1.5-2 u thick, moderately ie ia
fie pores 3 or 4, approximately equato
On Sisyrinchium Bermudianum L. ein St. Davids,
Bermuda, February 26—March 9, 1908, Stewardson Brown 611.
A number of rusts have been described on Sisyrinchium, both
from North and South America, but representatives of them are
rare and scanty in herbaria. Judging from descriptions and
the small amount of material available this form should be con-
sidered distinct.
Uredo cumula sp. nov.
II. Uredinia amphigenous, somewhat aggregate or scattered,
round, applanate, small, 0.1-0.4 mm. across, soon naked, pulveru-
lent, cinnamon-brown, conspicuous, ruptured epidermis notice-
On Buchnera elongata Sw. (Sora phadoriieandy Herradura,
Cuba, March 24, 1921, John R. Johnston 2530. The species
is notable for its small delicate spores, although when massed on
the surface of the plant they are of ordinary conspicuousness.
The generic name of the host is sometimes written Buechnera.
Uredo curvata sp. nov.
Fi Nr grts ae ee meen prdliped Doe ara: ¢
discolored areas 3-5 mm. across, round, small -O
in diameter, soon naked, pubaerpleat, dark chestnut ieee
ruptured epidermis spermine paraphyses numerous, per-
i Bhaae cede d, hyphoid or somewhat See ge rete about
7 by 30 uw, the wall thin, less than I yp, slightly thicker above,
I-2 uw, colorless below, light chestnut-brown above, sometimes
Nir strongly curv wall chestnut-brown, uniformly thin,
I with one indistinct pore on the concave side below the
eqnabie evenly echinulate-verrucose
On Inga vera Willd. (Mimosaceae), San Diego de los Bajios,
Cuba, March 26, 1921, John R. Johnston 2540. The species
is remarkable for its very much curved spores with one pore on
the indented side, being in this respect similar to the uredinia of
Puccinia invaginata Arth., which occurs in Cuba on the rhamna-
ceous genus Gouania.
Aecidium Yuccae sp. nov.
O. ia amphigenous, in small and crowded groups,
inconspicuous, honey-yellow, subepidermal, small, globoid,
196 Artuur: New sPEcIes oF UrEDINAE—XIV
80-115 » in diameter; eee wean ped prominent, 45-65 wv
long, agglutinated into a colum
. Aecia atiphizenous; Seentarty a in loose groups,
cylindric, 0.3-0.5 mm. in diameter and somewhat higher; peridium
firm, erect, the margin finely erose; seridial cells angularly
globoid or ellipsoid in face view, rectangular or somewhat
rhomboidal in section, 16-18 by 23-35 yu, abutted or slightly over-
lapping, the outer wall 3-5 uv. thick, smooth, the inner wall thin-
ner, about 2 u, finely verrucose; aeciospores globoid or broadly
aap 16-20 by nein Bs wall colorless, thin, I uv. or less, very
closely and finely ve
be a glauca Ratt: (Dracaenaceae), Crawford, Nebraska,
June 28, 1921, A. O. Garrett. An interesting species, as it comes
from the semi-arid, sandhill region, where a number of grass rusts
of restricted range occur, to one of which it undoubtedly is
genetically connected. Only a few groups of aecia on the
upper part of a young leaf were secured by Mr. Garrett, who
kindly submitted all the material for study.
PURDUE UNIVERSITY,
LAFAYETTE, INDIANA
Studies in the genus Lupinus—VII. L. succulentus and
iveus.
CHARLES PIPER SMITH
(WITH FOUR TEXT FIGURES)
INTRODUCTION
In accounting for the North American annuals and biennials
of the subgenus Lupinus, I have found it advisable to recognize
six groups, as indicated and contrasted in the key below. These
groups, however, do not provide for the Mexican annuals
L. bilineatus Benth., L. Hartwegi Lindl., and L. Barkeri Lindl.,
nor for the Costa Rican L. Clarket Oersted, as satisfactory
material of these species has not been available for my study.
Keel petals ciliate on their lower edges near the claw, com-
Racemes shorter than their peduncles. STIVERSIANI.
Racemes longer than their peduncles
Flowers verticillate; leaflets eubrous above. UCCULENTI.
Flowers scattered; leaflets more or less hairy above. pcan
Keel petals non-ciliate on their lower edges
Keel petals ciliate above near the point. MICRANTHI.
Keel petals non-ciliate on both upper and lower edges.
Flowers verticillate; leaflets hairy above. MICcRANTAHI.
Flowers with scattere
Leaflets _ above, 2-5 mm. wide; pods 10-15
seeds 1.5-3 mm. long. CONCINNI.
Leaflets atin above, 6-12 mm. wide; pods 30-50
m. long; seeds 4-6 mm. long. SUBCARNOSI. |
Fic. 75. 1. STIVERSIANI; 2. SUCCULENTI; a ere 4. MICRANTHI;
CONCINNI; 6. SUBCARNOSI.
Fic. 75 is here inserted to ee and contrast the above
197
198 Smit: STUDIES IN THE GENUS LuPpInus—VII
indicated variations in the keel. Four of these groups, namely,
the Sparstflort, Stiversiani, Concinni, and Subcarnost, have been
treated in the last two papers of this series (Bull. Torrey Club
47: 487-509. 1920; 48:219-234. 1921). This paper will con-
sider the Succulenti and one species of the Micrantht.
SUCCULENTI
The one species included here is so distinct from all the other
annuals of our region that I do not feel justified in placing it in
any of the other groups recognized. The ciliation of the keel
is too constant to be ignored, though nowhere properly described.
This is the group some years ago indicated by me under the
name Affines (Muhlenbergia 6: 134. 1911).
a. Lupinus sucCULENTUS Dougl.; C. Koch, Wochenschrift
Gaertn. Pflanzenkunde 4: 277. 1861. ec 76.|
tout, succulent or fistulous, 2-6 dm. tall, branched, near]
glabrous or sparsely appressed-pubescent rarely anaes pte
several; petioles slender, 6-12 cm. long, o o three times as
long as their leaflets, ecla cgromaesion = mm.
long, the free part widely divergent; leaflets seven to nine,
at apex, usually grag glabrous above, sparsely appressed-
0-70 m i
2-8 cm. long, racemes 6-30 cm. long, flowers subverticillate in
subappressed- ore the upper lip deeply two-toothed or
bifid, about 5 mm. long, the lower lip lanceolate, entire and acute
or three- toothed, 7-8 mm. long; banner suborbicular, about
3 mm. including the claw, glabrous, blue with yellow
pose turning violet, or rarely bluish white, wings 12-14 mm.
long, about 8 mm. wide, blue or rarely nearly white, more or
€
Siherniss whitish; pods ab 50 mm lon g, 9-10 mm. wide,
loosely pubescent. or villods sith hairs o. 5 to 1.5 mm. long,
ovules eight to ten; seeds oblong, 3.5-5 mm. long, much marbled
-with dark brown, with a pair of contiguous whitish spots
embracing the raphe, the hilum deeply sunken in a protruding
ring: axial root normally vertical.
This is the plant that has been known as L. affinis Agardh,
since Watson reviewed the genus in 1873 (Proc. Am. Acad.
8: 517), but the specimens so marked in the Lindley Herbarium
SMITH: STUDIES IN THE GENUS LUPINUS—VII 199
(now at Cambridge University, England) are not this species.
Indeed, Dr. Greene, in 1891 (Flora Franciscana 1: 40), expressed
doubt that our robust, succulent annual is the true L. affinis,
and I have long felt that Agardh’s description does not satis-
factorily apply to same. It remained, however, for Miss Alice
Eastwood to unearth the needed evidence, and her photograph
of Lindley’s specimen labelled L. succulentus shows said specimen
to be of this species. Likewise, her photographs of the specimens
labelled L. affints Agardh show equally well that same should
not be referred to this robust species, but rather to L. nanus,
a species which will be treated in a later paper. These are the
conclusions arrived at by Miss Eastwood with the specimens
before her.
Koch knew the species as a garden plant under the name
L. succulentus and quotes Biedenfeld’s Garten Jahrbuch as, his
2 Fy:
Fic. 76. LUuPINUS SUCCULENTUS ayes . C. P. Smith 1406 a 2
K. Brandegee (UC 81964); 3. E 5 Hees (DS 87569); 4. R.
Abrams 4224 (DS); 5. K. pa “(UC 187796).
only authority for attributing the name to Douglas. He also
states that the same species was distributed from Darmstadt
as L. Liebmanni and from Geneva as L. purpureus Del. Con-
sidering the species to be undescribed, he drew up in 1861 a
description in German, accréditing the species to Douglas and
suggesting that its native home must be assumed to be California
or Oregon. As his description is not readily available in this
ee it is reproduced here:
ze, mehr neidrig bleibende Pflanze ist mit Ausnahme der
Obuedsebs der Blatter sehr fein behaart; 9 langlichkeilférmige Blattchen
mit zuriickgebogener Spitze; Nebenblattchen wenig angewachsen, borsten-
200 SMITH: STUDIES IN THE GENUS LuUPINUS—VII
férmig, aufrecht; Aehre gestielt, kurz, aus wenigen (meist nur 3) Quirlen
bestehend; Deckblatter linger als die Knospen; Deckblattchen klein und
oft undeutlich; Oberlippe 2 theilig, kiirzer als die ganze Unterlippe;.
i wimpert
Diese durch ihre dunkelen, gear een Bliithen ausgezeichnete
und sehr zu empfehlende Art scheint noch gar nicht beschrieben zu sein.
Obwohl Douglas als Autor eecene ist un ad man vermuthen muss, dass
Kalifornien oder das Oregon-Gebiet Vaterland sind, so haben wir den Namen
doch nur in Biedenfeld’s Garten Jahrbuche (im 1. Hefte S. 292) gefunden.
Aus welcher Zeitschrift der Verfasser die Pflanze entlehnte, wissen wir
nicht. Neuerdings ist sie weider aus dem Darmstadter botanischen Garten
als L. Liebmanni, aus dem Genfer als L. purpureus Del. ver-
brietet worden. Wir vermuthen, dass die Namen in dem Verzeichnisse irgend
eines botanischen Gartens aufgestellt wurden.
Koch also here recorded his opinion that the species seems
to be related to L. arvensis Benth. and to L. ramosissimus Benth.,
a Colombian species collected by Hartweg, and especially to
ee ‘plant illustrated in plate eleven of the Botanical Register
for 1847 and there named L. Ehrenbergii. Evidently some one
claimed this species to be the same as L. densiflorus Benth.;
for Koch, in 1867 (Ind. Sem. Hort. Berol., App. 1: 11), published
a Latin diagnosis and stated that it differed from L. densiflorus
in its fragile stems and branches and. violet flowers. Since this
description is likewise difficult of access, I reproduce most of
it here:
“Annu us, humilis, ramosus, autiti puberulus; stipulae setiformes,
erectae, paululum adnatae; folia 9- igcueny 3-pollicaris; petiolus long-
itudine folia aequans; : oliolis supra glaberrimus subtus puberulis
oblongis, sed ad basin magis stlenintls? s apice breviter cuspidato, recurvo;
spica verticillata, verticillis distantibus; . . - labium superius
Bind: sit lanceolatum, integrum; : vedi et alae violaceae,
a albida, ad partem anlage violacea, apice flavescente;
legumen tills, inter semina constric
s per longum tam tempu: < hoe culta e California nec non
terris mexicanis sine dubio allata, anitadit ad L. densiflorus Benth., differt
tamen fragilitate caulis ramosumque et floribus violaceis. Nusquam
descripta esse videtur, sed nupperime a in hortis botanicis nominibus:
Lupini Liebmannii et purpurei occu
There is a sheet of this species, aks “Lupinus succulentus
Douglas, 1843”, in a large old collection of European garden
plants recently secured by Stanford University. Thus, I consider
it advisable to accept this name for this species, which otherwise
is without a proper botanical name. Douglas could hardly have
overlooked the species during his sojourn in California, and
certainly a more appropriate name could not be found. I cannot
SMITH: STUDIES IN THE GENUS LUPINUS—VII 201
follow Watson (Proc. Am. Acad. 8:538. 1873; and Bib. Index
236. 1878) in assuming that this “is probably but a garden
form”’ of L. densiflorus, nor accept his application of Agardh’s
name to this species. Koch might have said more about the
pods and seeds, but his “legumen . . . inter semina
constrictum” and his comparison of the species with L. arvensis.
L. ramosissimus, and L. Ehrenbergit do not point to a confusion
with L. denszflorus.
The species is so common in the San Francisco Bay region
that I will omit citation of many specimens examined. An albino
form (petals pale bluish white) was common in 1921, in and near
the mouth of Alum Rock Canyon, Santa Clara County. This
was mostly in pure patches, but sometimes associated with
pale blue, or the normal, dark blue, form. Seeds were secured
from marked colonies, but those from the palest-flowered plants
were heavily pigmented, perhaps averaging even darker, rather
than paler, than is normal for the species.
CALIFORNIA. Alameda County: Berkeley, Feb., 1899,
H. P. Chandler 239 (UC); Livermore, April, 1904, A. A. Heller
7320 (B, UC); Niles, June, 1918, H. A. Walker 5006 (UC);
Sunol Valley, June, 1916, L. R. Abrams 5699 (DS). Butte
County: Chico, A. A. Heller 12983 (UCX). Colusa County:
College City, 1905, Alice King (UC); Sycamore Slough, Sacra-
mento River, April, 1917, R. S. Ferris 621 (DS). Contra Costa
County: Brentwood, May, 1893, A. Eastwood (CA); Bryon
Springs, March, 1914, A. Eastwood (CA); Concord, March, 1914,
A. Eastwood (CA); Martinez, April, 1862, W. H. Brewer 906
(UC). Fresno County: Alcalde, March, 1892, 7. S. Brandegee
(CA); Alcalde, March, 1893, A. Eastwood (CA); Huron, March,
1893, A. Eastwood (CA). Kern County: Sunset Oil Wells,
March, 1893, A. Eastwood (CA). Los Angeles County: Avalon,
Santa Catalina Island, May, 1920, G. L. Moxley 732 (CPS);
Claremont, April, 1904, C. F. Baker (CA); Elysian Hills, Feb., -
1903, E. Braunton (UC); Los Angeles, April, 1901, E. D. Palmer
(UC); Redondo, San Pedro Hills, March, 1903, L. R. Abrams
3141 (DS); Santa Catalina Island, May, 1916, K. Brandegee
(UC). Monterey County: Carmel-by-the-Sea, March, 1910,
Alice D. Randall (DS); Castroville, T. S. Brandegee (CA);
Jolon Grade from King City, April, 1920, C. D. Duncan 78 (DS);
Pacific Grove, Pebble Beach, April, 1909, L. R. Abrams 4224
(DS); Posts, June, 1893, A. Eastwood (CA). Riverside County:
202 SMITH: STUDIES IN THE GENUS LUPINUS—VII
Hemet, May, 1904, C. F. Baker 4189 (UC, DS). Sacramento
County: Sacramento, April and May, Edna Hannibal (DS).
San Benito County: Idria, May, 1893, A. Eastwood (CA);
San Benito, May, 1918, A. Eastwood (CA). San Bernardino
County: San Bernardino, April, 1888, S. B. & W. F. Parish
(UC), and March, 1901, S. B. Parish (DS). San Diego County:
Campo, April, 1920, A. Eastwood (CA); La Jolla, April, 1903, .
T. S. Brandegee (UC), and March, 1914, F. E. & E. S. Clements
(UC); Linda Vista, May, 1894, 7. S. Brandegee (UC); Point
Loma, April, 1913, A. Eastwood (CA); San Diego, June, 1885,
Fanny E. Fish (UC), and March, 1889, C. R. Orcutt (CA), also
April, 1905, K. Brandegee (DS, UC). San Francisco County:
Twin Peaks trail, April, 1921, Bertha Dold 105 (CPS). San
Joaquin County: Tracy, April, 1903, C. F. Baker 2908 (CA).
San Luis Obispo County: San Luis Obispo, May, 1882, M. E.
Jones (CA), April, 1886, M. Miles (CA), July, 1911, K. Brandegee
(UC), and June, 1914, C. P. Smith 2851 (CPS); San Luis Valley,
March, 1882, Mrs. R. W. Summers (UC). San Mateo County:
Crystal Springs Lake, April, 1896, A. Eastwood (UC); Portola,
May, 1903, A. D. E. Elmer 4827 (CA, DS); La Honda summit,
May, 1900, W. R. Dudley (DS); San Mateo ravine, April, 1894,
W. R. Dudley (DS); San Pedro, June, 1903, A. D. E. Elmer 4681
(CA, DS). Santa Barbara County: Ellwood, May, 1908, A.
Eastwood 2 (CA); Santa Cruz Island, April, 1888, 7. S. Brandegee
(UC), July-Aug., 1886, E. L. Greene (CA), and June, 1918,
A. Easiwood (CA); Santa Maria River, June-July, 1906, A. East-
wood 329 (CA). Santa Clara County: Alum Rock Park, April,
1907, A. A. Heller 8471 (CPS, DS), Feb., 1921, C. P. Smith 3212,
albino, and 3273, normal colored (CPS); Loma Prieta, April,
1894, J. B. Davy 631 (UC); Los Altos, April, 1894, W. R
Dudley (DS); Saratoga, June, 1915, L. R. Abrams 5260 (DS);
Stanford University, April, 1903, A. D. E. Elmer gozo (CA, DS),
and May, 1902, C. F. Baker 858 (CA, UC); Campbell, March,
1921, Claribel Boesch ror (CPS); Evergreen, March, 1921, Lotta
Bland tor (CPS); Madrone Packwood School, April, 1921, Mrs.
A. F. Cochrane to2 (CPS). Santa Cruz County: Chittenden,
April, 1921, C. P. Smith 3259 (CPS); Ellicott, June, 1908, C. P.
Smith 1455 (CPS). Shasta County: Anderson, April, 1914, L.
E. Smith 139 (CA). Solano County: Cordelia, April, 1902, Heller
& Brown 5369 (B, DS); Vacaville, May, 1891, W. L. Jepson
(DS), and May, 1903, C. F. Baker 5071 (DS, UC). Sonoma
SMITH: STUDIES IN THE GENUS LUPINUS—VII 203
County: Petaluma Valley, April, 1908, C. P. Smith 1382 (CPS).
Sutter County: Marysville Buttes, April, 1915, A. A. Heller
11,792 (CA, DS, UCX); West Butte, April, 1917, R. S. Ferris
690 (DS); Yuba City, April, 1891, W. L. Jepson (UC). Te-
hama County: Red Bluff, May, 1914, L. E. Smith 670 (CA).
Ventura County: Ojai Valley, April, 1896, F. W. Hubby 41
(UC). Yolo County: Davis, April, 1915, P. B. Kennedy 21
(UCX); Yolo, June, 1914, C. P. Smith, seed only (CPS).
LOWER CALIFORNIA. Carysito, April, 1885, C. R. Orcutt (CA).
ArIzonA. Gila County: Roosevelt Dam, April 1917, A. East-
wood 6286 (CA), and May, 1919, A. Eastwood (CA)
The distribution of this plant is thus seen to be from Shasta
County to northern Lower California, the occurrence at Roosevelt
Dam, Arizona, being certainly due to a casual introduction
of seed from California. Variations are not particularly well-
marked, except in two cases, as indicated below:
1b. “Lupinus succulentus Layneae var. nov.
Humilis ramis decumbentibus, vitlosus pilis pandentibus
2 mm. longis, leguminibus aeque villosis
Depressed, 1 dm. tall, with decumbent branches and short
racemes, densely villous with hairs 2 mm. long even on the pods.
CALIFORNIA. San Mateo County: Farallon City, June,
1918, K. Brandegee (TypE, UC 187796).
Mrs. Brandegee’s maiden name was Mary Katharine Layne.
Ic. Lupinus succulentus Brandegeei var. nov. [FIG. 77.]
Humilis, erectus, foliolis maximis prope 20 mm. longis,
floribus 9-10 mm. longis, vexillo 10 mm. longo, 7.5 mm. lato,
alis apice non truncatis, carina ad basin super paulum ciliata.
uch reduced in stature and size of flowers; about 1 dm. tall,
largest leaflets about 20 mm. long, flowers 9-10 mm. long,
banner about 10 X 7.5 mm., wings not truncate at the apex,
keel less ciliate above.
LOWER CALIFORNIA. Comondu, Feb., 1889, 7. S. Brandegee
Set
Fic. 77. Lupinus SUCCULENTUS BRANDEGEEI C. P. Smith. 7. S. Bran-
degee (UC 83408).
204 Smitn: STUDIES IN THE GENUS LupInus—VII
(Type, UC 83498);San Julio, April, 1889, 7. S. Brandegee (UC),
and April, 1919, T. S. Brandegee (CA).
MICRANTHI
L. micranthus and its nearest relatives comprise the largest
-and most difficult group of the annual species of the subgenus
Lupinus Watson. Variations in shape and size of flowers,
leaflets, pods, and seeds produce conspicuous extremes which
have suggested ‘‘new species’? not a few in number. That
botanists have appreciated these variations is evidenced by the
following list of published names:
bicolor Lindl. Bot. Reg. 13: pl. r109. 1827.
micranthus Dougl.; Lindl. Bot. Reg. 15: pl. 1251. 1829.
nanus Dougl.; Benth. Tran. Hort. Soc. II. 1: 409.. 1835.
affinis Aga rdh, Syn. Gen. Lup. 20. 1835
Aduebornii S. Schauer, Linnaea 20: 739. 1 1847.
micranthus microphyllus Wats. oe Am. Acad. 8: 535. 1873.
niveus W thid. 11: I 876.
irifidus Torr.; Wats. ibid. 12: 0, 1877.
chihuahuensis Wats. ibid. 21: 1886.
carnosulus Greene, Bull. Cal. Read 3 Rae 1886.
umbellatus Greene, ibid. 2: 145.
packs Greene, Pittonia I: 65. “187
polycarpus Greene, thid, 2: 171.
bee so Eastwood, Proc. Cal. Acad. IL ve Pa pl. 56. 1896.
persistens Heller, Muhlenbergia 2:62. 1905
vallicola Heller, ibid. 4: 40. 1908.
apricus Greene, Leaflets 2: 67. 1910.
vallicola apricus C. P. Smith, Muhlenbergia 6: 135. I9QII.
hirsutulus Greene, Leaflets 2: 152. IgII.
Ee oe eee
strigulosus Gandoger. Bull. Soc. i ree 461. 1913.
In addition to accounting for these published names, the
scope of these papers will necessitate disposing of some seven
herbarium names. Of the above, only one species will be treated
at this time, the remainder being scheduled for attention in
subsequent papers.
The name Bicolores would have been a more appropriate
appellation for this group; but the name here employed has been
in use for some time. The flowers measure 6-16 mm. in length
and are always bicolored but are certainly not always small.
They are usually verticillate; but the racemes are sometimes
reduced to one umbel-like whorl, and in one form the flowers
are strictly scattered. The keel is normally ciliate on the upper
SMITH: STUDIES IN THE GENUS LuPINUS—VII 205
margin near the point, a character not shared by any other group
of North American annual lupines; however, two of the forms-
lack this ciliation. The upper surface of the leaves is almost
always with at least a few short hairs.
2. LUPINUS NIVEUs Wats. Proc. Am. Acad. 11: 126. 1876. [F1G. 78.]
pparently annual or biennial with pace petioled
cotyledons, erect, or dm. tall, each well above the base,
densely velvety tomentose above : e cotyledons, “which are
fleshy, glabrous, the blades about 20 mm. long, 12 mm. wide:
leaves velvety on both sides, netioles 6-8 cm. long, leaflets
eight or nine, rounded or angle so at the apex, 30-40 mm. long,
e: peduncles 5-8 cm. long, racemes 8-12 cm. long,
loosely euedleeee: bracts lecture linear, 5-8 mm. long;
flowers 10 mm. long, spreading or eae pedicels 4-6 mm
long; calyx gree ebracteolate, the lips ahs in Bose
about 5 mm. long, the upper cleft or bifid, over 4 mm. wide,
the lower bluntly nese barely 3 mm. wide: Setaly broad,
deep blue, banner suborbicular, about 10 X IO mm., greenish
Fic. 78. Lupinus NivEus Wats. E. Palmer 861 (G, ime
fem:
This is a very distinct, isolated, insular species not closely
related to any other known species; but as I see it, better lined
up with L. nanus, at least for the present. For a long time
I was misled by Watson’s clause, “‘allied to L. leucophyllus”’,
ome nk had difficulty in classifying this plant.
WER CALIFORNIA: Guadelupe Island, 1875, E. Palmer 25.
oe tes E. Palmer 861 (G, US); 1893, F. Franceschi (US).
My thanks are especially due to Miss Alice Eastwood for
the use of unpublished notes and to Dr. J. H. packard for
important bibliographical help.
The abbreviations used herein in the citation of specimens
may be identified by reference to the following list:
206 SMITH: STUDIES IN THE GENUS LUPINUS—VII
B, Brooklyn Botanic Garden
CA, California Academy of Senne:
PS, Sgt herbarium of the writer;
DS, Dudley Herbarium, Stanford University;
G, Gra y Herbarium, Harvard Universi sity
UC, Departient of Botany, University of Californi ia;
ao + Hien of Agronomy, University of California Exper-
iment Stat
US, United States National Herbarium.
San Jose, CALIFORNIA
A new American fossil hepatic
MARSHALL A. HOWE AND ARTHUR HOLLICK
(WITH ONE TEXT FIGURE)
Fossil Hepaticae are among the rarest of known paleobot-
anical remains. The total number of described species is less
than twenty. Of these more than half are identifiable as living
species of Jungermanniaceae, all but one of which are represented
merely by fragments enclosed in amber of recent geologic age,
found in the Old World. The remainder probably all belong in
the Marchantiaceae.
Three species only have been heretofore recorded from
America,* all of which are referable, either definitely or provis-
ionally, to the latter family. It should be remarked, however,
that the identity, as well as the botanical relationship of the
species last cited, in our footnote, is questioned by Fontaine, who
says (1. c., p. 54): “It is similar to that of a Brachyphyllum and
the plant may be really a twig of that conifer.’’ Incidently it may
also be remarked that this is the only species recorded for any
geological horizon older than Tertiary, and that the type of the
species, from the Jurassic of England, was originally described
as an alga under the name Fucoides erectus Bean.}
Several years ago a collection of fossil plant remains from
Florissant, Colorado, was sent by Mr. T. D. A. Cockerell to the
was what appeared to be a fragment of a bryophyte; but it was
so small and imperfectly defined that it was not identified,
even as to its probable class relationship, until an enlarged pho-
tograph (Fic .1) was made. The specimen evidently represents a
*(a) Preissites Wardii Knowlton, Bull. Torrey Club 21: 458. pl. 279. 1894.
Eocene (Fort Union formation), Burn’s Ranch, thirty: miles from Glendive,
Montana.
(b) Marchantia Pealet Knowlton, Proc. U. S. Nat. ar 35: 157. pl. 25,
1908. Eocene (Lance formation), Custer County, Monta
(c) ‘‘Marchantites erectus (Bean) Seward?’’, fide Sicteive in Ward,
Monog. U. S. Geol. Survey 48:53. pl. 6, f. 1, 2. 1905. Jurassic, Douglas
County, oe
enue Quart. Jour. Geol. Soc. pri 90: 8r. pl. 71, f. 3a, 3b
ea ien numbered 2a, 2b on the plate). 18
207
208 Howe & Ho.uick: A NEW FOSSIL HEPATIC
member of the Bryophyta and probably belongs to the family
Jungermanniaceae of the class Hepaticae.
Jungermanniopsis gen. nov.
Plants slender and minute, consisting of stem and leaves and
showing a pronounced dorsive ntrality, with ventral (postical)
b
roothairs and dorsal (antical) apparently one-ranked but pro-
ably two-ranked ovate-acuminate, often subfalcate, assurgent
or subvertical euteauy secund) leaves ee ith occasional
n
ligulate leaves
Jungermanniopsis Cockerellii sp. nov.
Stem ascending at apex; leaves I-1.5 mm. long,
i 0.4
in maximum width, contiguous, Garonls ster | or slightly
overlapping at extreme base, the intervals between the leaves
s
sometimes accentuated by reflexions of the subapical margins,
the margins entire or with indistinct indications of small teeth.
COLLECTOR: T. D. A. Cockerell.
FORMATION AND LOCALITY: Miocene shale, Florissant,
Colorado (Station 14).
ype specimen in the Museum ot the New York Botanical
Garden.
The one fragment, upon
which the genus and species are
based, is about 12 or 13 mm. long
and evidently lies on its side.
The specimen shows a possible
forkingof the stem near the apex,
with one branch ascending and
Fic. I. JUNGERMANNIOPSIS COCK- the other descending, but the
EueLLn Howe & Hollick descending part is probably
only a collection of roothairs.
Among the living Hepaticae, the ne ae antically secund leaves
of Jungermanniopsis suggest the widely distributed Herberta
adunca (Dicks.) S. F. Gray; but there is no indication that the
leaves are bilobed as in the genus: Herberta. In its antically
secund leaves, it resembles also such leafy Hepaticae as James-
oniella autumnalis (DC.) Steph. and certain species of Nardia,
but in the form and looser disposition of the leaves it is very
different from any of these. The possible presence of smaller
Howe & Ho iick: A NEW FOSSIL HEPATIC 209
ligulate or lanceolate lateral leaves might, if fully substantiated,
result in placing the plant with the Musci rather than with the
Hepaticae. But the small size and general habit of the organism
suggest affinities with the Jungermanniaceae rather than with the
Musci. The specific name is in honor of Mr. T. D. A. Cockerell,
the collector.
A new genus of fossil Liliaceae
T. D. A. CocKERELL
(WITH ONE TEXT FIGURE)
In 1873* Lesquereux reported a singular fossil from the
Miocene shales of Florissant, Colorado, as Ophioglossum Allent.
In 1878 he figured itt and assigned it to the genus Salvinia,
on account of its-close resemblance to Heer’s European fossil
S. reticulata. Hollick{ gave a new figure, and placed the plant
in the genus Ymesipteris, where it remains in Knowlton’s
Catalogue.
This so-called Tmesipteris Alleni is common in the Florissant
shales but has remained an unsolved problem. A number of
years ago, at the British Museum, I showed specimens to Dr.
A..C. Seward and Dr. A. G. Nathorst. We could all agree that
there was no relationship with Tmesipteris, but it was im-
possible to even guess at the true position of the species. Seward,§
when publishing on the subject later, reject-
ed the reference to Tmesipteris, without
being able to suggest another alternative.
In 1913,{] to avoid perpetuating an obvious
error, I referred the plant to Carpolithes,
supposing it to represent some sort o
fruiting body. No seeds could ever be
seen, although the appearance suggested
a flattened pod; but the published figures
failed to show a thickening on the midrib, :
; i ALLEN! Cockerell
a little beyond the middle. In a very good Ais: Chuledesthbien
specimen found at Florissant in November, pistillate flower.
1921, there is a definite body at this point, flower, enlarged. C.
with a diameter of nearly3mm. It looks Fruit on a mature clad-
like a small berry, with the contents ex- 4 arginate
truded (Fic. r, C). a a cladode.
Probably the riddle would never have been solved but for
the discovery of a younger specimen in the Florissant shales,
Fic. 1. BRACHYRUSCUS
* For bibliographical references see Knowlton, Mesozoic and ee
plants of North America. U. = eo Survey Bull. 696: 6 1909
+ Tertiary Flora, pi. a 7 Be
t Bull. Torrey Club. sara ide oe 12. 1894.
§ Fossil ee a: 25: ge
Ser Vi: 86: 500.
212 COCKERELL: A NEW GENUS OF FOSSIL LILIACEAE
at Station 14. This (Fic. 1, A, B) shows that we have to
do neither with a leaf, frond nor pod, but with a cladode. At-
tached to the midrib is anindistinct mass, presumably a thin bract,
upon which can be seen a dark object (Fic. 1, B), which seems to
agree very closely with the flower of Ruscus. The pistil is
erect and distinct. The plant is dioecious, and the staminate
flower is as yet unknown. At first I thought there might be
two flowers, but the supposed second one is, I believe, part of
the midrib.
As I could not identify the plant with Rzscus, I consulted
Dr. Britton, who very kindly sent me material of Phyllonoma
ruscifolia Willd. and Ruscus hypoglossum L. It was at once
apparent that the fossil had nothing to do with Phyllonoma,
but returning to Ruscus as the only alternative, I was able
to recognize close morphological similarity. The genus Ruscus
is today limited to a few Palaearctic species, with lanceolate
to ovate sharply pointed cladodes. The apical points are stiff
and sharp, as any one who has handled butcher’s broom will
testify. At first sight the venation seems quite different from
that of the fossil, but if we imagine the Ruscus cladode broadened
and abbreviated until the principal veins are nearly or quite
transverse instead of longitudinal, the correspondence is exact.
The published figures of the fossil do not show that there are
principal veins, just as in Ruscus.
The morphological correspondence is so exact t that it becomes
a question whether the fossil can be referred to Ruscus. However,
considering the broad form of the cladodes, with truncate or
emarginate, mucronate apex, the essentially transverse veins,
and the remoteness in time and space, it appears justifiable to
establish a distinct genus, Brachyruscus, the species becoming
Brachyruscus Alleni sp. nov.* Whether Heer’s Salvinia ret-
* Dr. Hollick writes that he is not convinced that the flowering cladode,
which I have figured and described, belongs to the same plant as the ‘‘ Tmes-
tpteris” Alleni. 1 had considered the possibility of there being two different
things, and am personally entirely convinced that all the specimens belong
to a single species. However, I will designate the flowering specimen as the
type of the genus and of a species which I will name B. Alleni. This name
will remain valid if my view is correct, only B. Alleni Cockerell will be a
synonym of B. Alleni (Lesq.) Cockerell. If it should be decided that there
_ are two quite different plants, B. Alleni Cockerell will still stand, only to
fall in case some author considers that two distinct species of Brachyruscus
are present
COCKERELL: A NEW GENUS OF FOSSIL LILIACEAE 213
iculata is congeneric, it is impossible to say; the reticulation,
as figured, looks different
Ruscus today has its western limit in Madeira. The insular
form, which I saw in cultivation in Funchal, is regarded as a
race of R. hypophyllum L., but is perhaps specifically distinct.
The name R. latifolius ince is preoccupied by Rafinesque.
BRACHYRUSCUS gen. nov.
Cladodes pyriform to subeval in outline, with a strong midrib;
apex obtuse, emarginate, with a stiff mucro representing the
ite-shap ea m
midrib as in Ruscus, wie apparently a single flower, which (as
imperfectly preserved) presents no characters to differentiate
it from Ruscus. Type the following
Brachyruscus Alleni sp. nov.
Cladode in es ides aber broad-pyriform, with tapering
nd
stems unknown but doubtless ae
INDEX TO AMERICAN BOTANICAL LITERATURE
1921
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
America being used in the broadest sense.
Reviews, and papers that relate exclusively to foresty, agriculture, horti-
ae, reesei gane: pratanty of vegetaiee origins. or a oratory methods
nd f bacteriology.
An occasional spades is uate ¢ in favor of some paper appearing in an Ameri-
of the Index will call o attention of the editor to errors or omissions, their
og will be appreciated
s Index is ctatalk pene on cards, and furnished in this form to
Sondices at the rate of three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Kern, F. D. Distribution of Berberis vulgaris in Pennsylvania.
Bull. Torrey Club 48: 263-269. 31 D 1921.
Kern, F. D. The J. Roberts Lowrie Herbarium. Torreya 20:
79-81. O 1921.
Kern, F. D. Observations of the dissemination of the barberry.
Ecology 2: 211-214. 3 D 1921.
Keuchenius, P. E. Die Rindenbraune der Hevea brasiliensis.
Centralbl. Bakt. Parasitenk. u. Infektionskr. 55: 14-74.
f.- 1-33. 18 N 1921
Keys, A. The Agricultural Department, Dominica. Kew Bull.
Misc. Inform. 1921: 67-85. 1921.
Joss T. R. A sectorial chimera in maize. Jour. Hered.
a: 2B4c)> 32-0 25 S-1G2t.
A. B. The marine algae of the Passamaquoddy region,
New Brunswick. Contrib. Canad. Biol. 1917: 79-84. pl.
&; sory
Klugh, A. B. Springtime in the woods. Am. For. 27: 317-
319, 324. My 1921. [Illust.]
215
216 INDEX TO AMERICAN BOTANICAL LITERATURE
Knowlton, C. H. Herbarium of Rev. W. P. Alcott. Rhodora
23: 47« 5 Ap 1921.
Knowlton, F.H. Criteria for determination of climate by means
of fossil plants. Bull. Geol. Soc. Amer. 32: 353-358. 1
S 1921.
Knox, A. A. The induction, development, and_ heritability
of fasciations. Carnegie Instit. Washington Publ. 98: 1-18.
pl. 1-5 + f. 1. 1908.
[This is the second ‘paper of this publcatiion.].
Knudson, L., & Ginsburg, S. Suggestions with respect to the
measurement of osmotic pressure. Am. Jour. Bot. 8:
164-170. f. r. 3 Ap r92t.
Knunker, A. Hans Freiherr von Tiirckheim. Gartenflora
70: 19-22. 15 Ja 1921.
Von Tiirckheim collected chiefly in Guatemala
Koch, E., & Rumbold, C. Phoma on sweet sorghum. Phyto-
pathology 11: 253-268. pl. 9-11 + f. 1-3. 15 N 1921.
Kradusel, R. Ist Taxodium distichum oder Sequoia semper-
virens Charakterbaum der deutschen Braunkohle? © Ber.
Deutsch. Bot. Ges. 39; 258-263. f. 1-3. 10S 1921.
Includes a discussion and illustrations of ]the parenchyma of Taxodium
mexicanum.
Krieger, L. C. C. Is Amanita pantherina edible or poisonous?
Mycologia 13: 270, 271. 8 O 1921.
Lachmund, H. G. Some phases in the formation of fire scars.
Jour. For. 19: 638-640. O 1921.
Leighty, C. E., & Boshnakian, S. Genetic behavior of the spelt
orm in crosses between Triticum spelta and Triticum
sativum. Jour. Agr. Research 22: 335-364. pl. 33 + f:
I-3. 12 N 1921. :
Leonian, iL. H. Studies on the Valsa apple canker in New
Mexico. Phytopathology 11: 236-243. 21 O 1921.
Lindau, G. Eine neue Aphelandra-Art. Notizbl. Bot. Gart.
u. Mus. Berlin-Dahlem 8: 50. 1 S 1921.
Aphelandra Dusenii from Brazil.
INDEX TO AMERICAN BOTANICAL LITERATURE 217
Lipman, C. B. A contribution to our knowledge of soil relation-
ships with Citrus chlorosis. Phytopatholhgy 11: 301-305.
9 D Ig21.
Loeb, J. The quantitative basis of the polar character of re.
generation in Bryophyllum. Science II. 54: 521, 522-
25 N 1921.
Long, C. A. E. Some rare plants from Knox County, Maine.
Rhodora 23: 198, 199. 15 D 1921.
Lyon, C. J. Comparative studies on respiration, XVIII.
Respiration and antagonism in Elodea. Am. Jour. Bot.
8: 458-403. f, I, 2" 10° 104k:
Macoun, J. M., & Holm, T. The vascular plants of the arctic
coast of America west of the 1ooth meridian, collected by
the Canadian Arctic Expedition, 1913-18. Rep. Canad.
Arctic Exped. 1913-18. 5A: 5-25. pl. 1-3. 14 O 1921.
Magrou, J. Symbiose et tuberisation. Ann. Sci. Nat. Bot. X.
3: 181-275. pl. 1-9 + f.-1-9. N 1921. '
Mains, E. B. Unusual rusts on Nyssa and Urticasirum. Am.
Jour. Bot. 8: 442-451. f. 1-6. . 19 D 1921.
Includes A plospora, gen. nov.
Marsh, C. D., & Clawson, A. B. The Mexican whorled milk-
weed (Asclepias mexicana) as a poisonous plant. U. S.
Dept. Agr. Bull. 969: 1-16. pl. 1-3 + f. 1. 6O 1921.
Marsh, C. D., Clawson, A. B., Couch, J. F., & Marsh, H. West-
ern sneezeweed (Helenium Hoopesii) as a poisonous plant.
U.S. Dept. Agr. Bull. 947: 1-46. pl. 1,2 +f. 1-5. 1101921.
McWhorter, F. P. Destruction of mosses by lichens. Bot.
Gaz. 72: 321-325. pl. 13. 15 N 1921.
Mead, C. W. Indian corn or maize. Nat. Hist. 21: 408-413.
O 1921. [Illust.] -
Menzies, D. Sir Archibald Menzies, R. N. Gard. Chron.
702 324. 24 D 1921.
Merrill, E. D. Review of the new species of plants proposed
by N. L. Burman in Flora Indica. Philipp. Jour. Sci. 19:
329-388. S 1921.
218 INDEX TO AMERICAN BOTANICAL LITERATURE
Metcalf, W. Notes on the Bishop pine (Pinus muricata). Jour.
For. 19: 886-902. D 1921. [Illust.]
Miller, H. G. Further studies on relation of sulphates to ee
growth and composition. Jour. Agr. Research 22: 101-110.
8 O 1921.
Moffatt, W. S. Why Illinois prairie flowers have disappeared.
Nat. Stud. Rev. 17: 307-309. N 1921.
Molfino, J. F. Contribucién a la flora de la regién de Bahia
Blanca (Argentina). Physis 5: 1-27. 31 O 1ga2t._ [Illust.]
Mulford, F. L. Trees with bright autumn foliage. Am. For.
27: 648-655. O 1921. [Illust.] -
Munns, E.N. Coulter pine. Jour. For. 19: 903-906. D 1921.
Munns, E. N. Effect of location of seed upon germination.
Bot. Gaz. 72: 256-260. 15 O 1921.
Orton, C. R., & Weiss, F. The reaction of first generation
hybrid potatoes to the wart disease. Phytopathology
LEy 200-310... g LF. tot.
Osborne, T. B., Wakeman, A. J., & Leavenworth, C. S. The
proteins of the alfalfa plant. Jour. Biol. Chem. 49: 63-91.
N 192!.
Pascher, A. Uber die Ubereinstimmungen zwischen den Diato-
meen, Heterokonten und Chrysomonaden. Ber. Deutsch.
Bot. Ges. 39: 236-248. f.1-6. 10S 1921.
Pemberton, C. C. Overgrowth of stumps of conifers. Canad.
Field Nat. 35: 81-87. f. 1-4. 22 N 1921.
Perry, W. J. Some observations on the relation of soil moisture
to height growth of pine saplings. Jour. For. 19: 752,
wea IN 394; ,
Perry, W. J. Yellow pine reproduction. Jour. For. 19: 622-
631.. O: 19021.
Plaisance, G. P., & Hammer, B. W. The mannitol-producing
organisms in silage. Jour. Bact. 6: 431-443. S 1921.
Pomeroy, C. S. Bud variation in Eleagnus. Jour. Heredity
12: 227-230. f. 10, 20.° 34 S 1621.
INDEX TO AMERICAN BOTANICAL LITERATURE 219
Puttick, G. F. The reaction of the F, generation of a cross
between a common and a durum wheat to two biologic
forms of Puccinia graminis. Phytopathology 11: 205-
24%, 04) toot.
Raber, O.L. The effect upon permeability of polyvalent cations
in combination with polyvalent anions. Am. Jour. Bot.
8: 382-385. f.. 1... 14 N -t921.
Studies of Laminaria.
Raber, O. L. The effect upon permeability of (I) the same
substance as cation and anion, and (II) changing the
valency of the same ion. Am. Jour. Bot. 8: 464-470. f. 7, 2.
19 D 1921.
Rangel, E. Nota sobre dois fungos. Arch. Escol. Super. Agr.
e Med. Veter. 5: 35-40. pl. 3-5. S 1921.
Oswaldia, gen. nov., and Peniotrichum saccardinum, sp. nov.
Record, S. J. Figure in wood. Am. For. 27: 611-617. O
1921. [Illust.
Rehder, A. Plant nomenclature: more suggestions. Jour. Bot.
289-294. O 1921. ;
Reinking, O. A. Fiji disease of sugar cane in the Phillippine
Islands. Phytopathology 11: 334-337. pl. 15,10. 9D 1921.
Reinking, O. A., & Groff, G. W. The Kao Pan seedless Siamese
pummelo and its culture. Philipp. Jour. Sci. 19: 389-437.
pl. 1-16 + f. 1. O 1921.
Rendle, A. B..- sta an overlooked genus-name. Jour. Bot.
Be 261-264. SI
ludes reprint of the ee publication of this genus and its 3 species
by aoe in New Orleans Med. & Surg. Jour. 1847: 614-616. 1847.
Renner, O. Das Rotnervenmerkmal der Onotheren. Ber.
Deutsch. Bot. Ges. 39: 264-270. 105 1921.
Richards, B. L. A dryrot canker of sugar beets. Jour. Agr.
Research 22: 47-52. pl. 4-9. 1 O 1921.
Roark, E. W. The Septoria leaf spot of Rubus. Phytopath-
ology 11: 328-333. 9 D 1921.
Due to Mycosphaerella Rubi, sp. nov.
220 INDEX TO AMERICAN BOTANICAL LITERATURE
Roberts, J. W. Plum blotch, a disease of the Japanese plum,
caused by Phyllosticta congesta Heald and Wolf. Jour.
Agr. Research 22: 365-370. pl. 34 + f. 1, 2. 12 N I921.-
Robinson, T. R. The bud-sport origin of a pink-fleshed grape-
fruit in Florida. Jour. Hered. 12: 195-198. 23 S 1921.
[Illust.]
Rogers, E. C. Influence of the period of transplanting western
white pine seedlings upon their behavior in nursery and
plantation. Jour. Agr. Research 22: 33-46. f. 1-7. 1 O 1921.
Round, E. M. Odontopteris genuina in Rhode Island. Bot.
Gaz. 32: 397-403. f. I-53. 15 D 1921.
Rumbold, C., & Tisdale, E. K. Notes on Phoma insidiosa Tass.
found on Sudan grass. Phytopathology 11: 345 9 D 1921.
Ruschmann, W. Die Cohune- oder Corozoniisse Mittelame-
rikas. Tropenpflanzer 24: 147, 148. O 1921.
Satterthwait, A. F. Notes on the food plants and distribution
of certain billbugs. Ecology 2: 198-210. f. 7. 3 D 1921.
Schlechter, R. Die Thismieae. Notizbl. Bot. Gart. u. Mus.
Berlin-Dahlem 8: 31-45. 1S 1921.
ncludes Triurocoden, gen. nov., from Brazil, and new American species
in Sines (1), Myostoma (1).
Schiiepp, O. Zur Theorie der Blattstellung. Ber. Deutsch.
Bot, Ges. 39: 249-257. f. FT, 2. .10 S 1921.
Schultz, E. S.A transmissable mosaic disease of Chinese cab-
bage, mustard, and turnip. Jour. Agr. Research 22: 173-
178. pl. B. 15 O 1921.
Seymour, E. K., & McFarland, F. T. Loss from rye ergot.
Phytopathology 11: 285-289. f. 1.2. 15 N 1921.
Shapovalov, M., & Edson, H. A. Blackleg potato tuber-rot under
irrigation. Jour. Agr. Research 22: 81-82. pl. 13-16 +
pl. A. 8 O 1921.
Showalter, A. M. Chromosomes of Conocephalum conicum.
Bot. Gaz. 72: 245-249. pl. 4, 5. 15 O 1921
Sievers, A. F., & McIntrye, J.D. Changes in the composition
of paprikas (Capsicum annuum) during the growing period.
Jour. Am. Chem. Soc. 43: 2101-2104. S 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 227
Sinnott, E. W. The relation between body size and organ
size in plants. Am. Nat. 55: 385-403. f. 1, 2. O 1921.
Skinner, J. J.. & Reid, F.R. Nutrient requirements of clover
and wheat in solution cultures. Soil. Sci. 12: 287-296. pl.
£.. 42 4981,
Small, J. K. Along the Juniata River ie Jour.
New York Bot. Gard. 22: 168-171.
Small, J. K. Another Sonchus for America. Torreya 21:
100, 101. D Igat.
Records the finding of the European Sonchus uliginosus in Pennsylvania.
Small, J. K. Cercis chinensis. Chinese redbud. Addisonia
6: 33, 34. pl. 209. 24 O 1921.
Small, J. K. Monarda didyma. Oswego-tea. Addisonia 6:
47, 48. pl. 216 24 O 1921.
Smiley, F. J. A report upon the boreal flora of the Sierra
Nevada of California. Univ. California Publ. Bot. 9: 1-
423. pl. 1-7. -S 1921.
Includes 21 new names or combinations.
Spencer, E. R. uae of Brazil nuts. Bot. Gaz. 72: 265-292.
pl. 8-12 + f. 1-3 1921.
Includes new species in Pellioniella (1), Cephalosporium (1), Phomopstis
7);
Spink, H. M. The geographical distribution of commercial
timber in the Pacific States of North America and its
significance to the lumber industry. Scottish Geog. Mag.
37: 257-266. 15 O 1921. [Illust.]
Sprague, T. A. The generic name Schizonotus. Jour. Bot. 59:
249-251. S 1921.
Sprague, T. A. Plant dermatitis. Jour. Bot. 59: 308-310.
Steil, W. N. Vegetative reproduction and aposporous growths
from the young sporophyte of Polypodium irioides. Bull.
Torrey Club 48: 203-205. 265 1921.
Steil, W. N. The development of prothallia and antheridia
from the sex organs of Polypodium irioides. Bull. Torrey
Club 48: 271-277. pl. 4+ f.1-4. 31D 1921.
222 INDEX TO AMERICAN BOTANICAL LITERATURE
Sure, B., & Read, J. W. Biological analysis of the seed of the
Georgia velvet bean, Stizolobium Deeringianum. Jour.
Agr. Research 22: 5-15. f. I-15. 1 O 1921.
Swanson, C. O. Hydrocyanic acid in Sudan grass. Jour.
Agr. Research 22: 125-138. 15 O 1921.
Tavares, J. S. Cecidologia brazileira. Broteria, Zool. 19:
97-112. 1 D Igali. :
Taylor, F. B. The literature of diatoms. Trans. Am. Micr.
Soc. 40: 187-194. O 1921.
Taylor, M. W. Internal aecia of Puccinia albiperidia Arthur.
Phytopathology 11: 343, 344. f. z. 9 D 1921.
Taylor, N. Endemism in the Bahama flora. Ann. Bot. 35:
h23-632.: -O.-T62t.
Taylor, N. Plant immigrants and natives: a comparison.
Brooklyn Bot. Gard. Leafl. IX. 9: 1-7. 26 O 1921.
Thaxter, R. Lincoln Ware Riddle. Rhodora 23: 181-184.
15 D tIg21.
Thom, C., & Lefevre, E. Flora of corn meal. Jour. Agr. -
Research 22: 179-188. 22 O 1921.
Thompson, W.P. A botanical trip through German South-west
Africa. Canad. Field Nat. 35: 74, 75. 25 O 1921.
Townsend, C.O. The beet-sugar industry in the United States.
U.S. Dept. Agr. Bull. 995: 1-58. pl. r-r0. 14 O 1921.
Urban, I. Plantae Caribaeae. Notizbl. Bot. Gart. u. Mus.
Berlin-Dahlem 8: 25-30. 1 S 1921.
Includes new species in Capparis (1), Sloanea (1), Pachira (1), Basana-
cantha (1). :
Vaupel, F. Echinopsis mamillosa Giirke. Monatschr. Kakteenk.
31:152-155. O1g2t. [Illust.]
Vinall, H. W., & Getty, R. E. Sudan grass and related plants.
U.S. Dept. Agr. Bull. 981: 1-68. f. -25. 21 D 1921.
Wagner, E.. Einiges iiber Malacocarpus. Monatschr. Kak-
teenk. 31: 143, 144. S 1921.
BULLETIN
OF THE
TORREY BOTANICAL CLUB
AUGUST, 1922
Philippine Basidiomycetes—V*
Pau W. Grarr-
POLY POREAE (Con.)
_PoriA Persoon
PorIA LEUCoPLACA (Berk.) Sacc. Syll. Fung. 6: 322. 1888.
Polyporus leucoplacus Berk.; Hooker, Flora Nov. Zeland. 2:
180. 18
Luzon: Proves of Benguet, Pauai, June, 1909, R. C.
McGregor, Bur. Sci. 8720, at an elevation of 2,100 m., on decaying
timber.
Collected previously in New Zealand and Java.
TRAMETES Fries
TRAMEIES ASPERA (Jungh.) Bres. Hedwigia 53: 69. 1912.
Polyporus asper Jungh. Flor. Crypt. Javae 1: 61. 1838.
Polystictus asper Sacc. Syll. Fung. 6: 224. 1888.
Luzon: vicinity of Manila, Aasaants 1912, Sanchez 26, 27,
growing on decayed lo
Originally described from material collected in Java.
TRAMETES INCANA Lévy. Ann. Sci. Nat. Bot. III. 2: 196. 1844.
*The papers of this series already published have appeared as follows.
Additions to the basidiomycetous flora of the Philippines. Philip. Jour. Sci:
8: (Bot.) 299-309. pl. 8-ro. 1913; Philippine Basidiomycetes, IJ. Philip
Jour. Sci. 9: (Bot.) 235-255. pl. 2. 1914; Philippine Basidiomycetes—III.
Bull. Torrey Club 45: 451-469. pl. 15. dt Philippine Basidiomycetes—
IV. Bull. Torrey Club. 48: 285-295. 192
[The BuLLertin for July (49: 189-222) was issued August 10, 1922.|
223
224 GRAFF: PHILIPPINE BASIDIOMYCETES—V
Polyporus incanus Lév.; Gaudichaud, Bot. Voy. Bonite 1: 183.
eb. 297, J. 2. 1846.
Luzon: Province of Bataan, Mount Mariveles, November,
1912, P. W. Graff, Bur. Set. S157, S158.
This fungus was first collected in the Philippines by Gaudi-
chaud-Beaupré, in 1836-37, on the visit of the corvette “La
Bonite” to the port of Manila. The fungus has not been reported
since, and it is with considerable pleasure that one of these
three-quarter century old species is again collected.
TRAMETES PALEACEA Fr. Nov. Symb. Myc. 97. 1851.
Polyporus paleaceus Fr. Epicr. Myc. 471. 1838.
Luzon: Province of Bataan, Mount Mariveles, November
3-19, 1912, P. W. Graff, Bur. Sci. 19098, on a decaying log at
an elevation of 400 m.
Previously reported from New Guinea.
TRAMEIES CORRUGATUS (Pers.) Bres. Hedwigia 51: 316. 1912.
Polyporus corrugatus Pers.; Gaudichaud, Bot. Voy. Uranie
172. 1826.
Polyporus fusco-badius Pers.; Gaudichaud, Bot. Voy. Uranie
172. 1826.
Polyporus scabrosus Pers.; Gaudichaud, Bot. Voy. Uranie
172:
Daedalea sanguinea Klotz. Linnaea 8: 481. 1833.
Polyporus indecorus Jungh. Flor. Crypt. Javae 1: 51. 1838.
Polyporus tegularis Lév. Ann. Sci. Nat. Bot. III.5:131. 1846.
exagonia cruenta Mont. Syll. Gen. Spec. Crypt. 169. 1856.
Polystictus Persoonii Cooke, Grevillea 14: 85. 1885.
Trametes nitida Pat. Jour. de Bot. 4: 17. 1890.
Earliella cubensis Murr. Bull. Torrey Club 32: 479. 1995-
Earliella corrugata Murr. Bull. Torrey Club 34: 468. 1907-
Luzon: Province of Laguna, Mount Maquiling, February,
1912, P. W. Graff, Bur. Sci. 16027..
_ This is a much described species of universal tropical distri-
bution. It was first described from material collected in the
island of Guam and has later been reported from West Africa,
India, Ceylon, Java, Borneo, Sumatra, the Philippines, Australia,
and the American tropics.
GRAFF: PHILIPPINE BASIDIOMYCETES—V 225
DAEDALEA Persoon
DAEDALEA LURIDA Said Ann. Sci. Nat. Bot. III. 2: 189. 1844.
Daedalea sub Murr. Bull Torrey Club 35: 4F5. 1908.
Luzon: Provihce of Rizal, January, 1906, F. W. Foxworthy,
Bur. Sct. 22 (type of Daedalea subconfragosa Murr.). MINDANAO:
‘District of Davao, Lake Laneo, Camp Keithley, July, 1907,
Mary S. Clemens ‘“V”"’.
Murrill’s species, described from Philippine material, seems
to be without doubt identical with that described by Léveillé
from Javan specimens.
DAEDALEA PALIsoti Fr. Syst. Myc. 2: 335. 1821.
Daedalea amanitoides Beauv. Fl. Owar. 1: 44. pl. 25. 1804.
_— repanda Pers.; Gaudichaud, Bot. Voy. Uranie
1826
Sere eppionda Klotz. Linnaea 8: 481. 1833.
Lenzites Palisoti Fr. Epicr. Myc. 404. 1838.
Lenzites repanda Fr. Epicr. Myc. 404. 1838.
Lenzites applanata Fr. Epicr. Myc. 404. 1838.
Lenzites pallida Berk. Jour. Bot. 1: 146. 1842.
Lenzites platypoda Lév. Ann. Sci. Nat. Bot. III. 2: 180. 1844.
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sci. 1191; Province of Laguna, Mount Maquiling, Feb-
ruary, 1912, P. W. Graff, Bur. Sci. 15951, on fallen and decayed
tree trunk. CuLion: October, 1903, Merrill 3531. NEGROS:
Gimagaan River, January, 1904, Copeland 21.
Daedalea indica Jungh., described from Javan cea
should in all probability be added to the list of synonyms of
this species. Daedalea, in the Polyporaceae, and Lenzites, in the
Agaricaceae, are so closely related, and the species of one so
often show a strong tendency to a variation toward the other,
that considerable confusion has arisen in their determination.
This is especially true in the case of this species and D. flavida Lév.
in the Philippine material. In the latter species the variation
even goes further, producing a Trametes form.
The types of both Léveillé’s and Berkeley's species are
from material collected in the Philippines. This species is of
very general tropical distribution.
DaEDALEA PRUINosA Lév. Ann. Sci. Nat. Bot. IIT. 2: 199. 1844.
226 GRAFF: PHILIPPINE BASIDIOMYCETES—V
Hexagonia glabra Lév. Ann. Sci. Nat. Bot. III. 5: 143. 1846.
Luzon: Province of Laguna, Mount Maquiling, Kebuers
1912, P. W. Graff, Bur. Sci. 16015.
Collected previously in India and the Hawaiian Islands.
Daedalea versatilis (Berk.) comb. nov.
rametes versatilis Berk. Jour. Bot. 1: 150. 1842.
Hexagonia ciliata Klotz. Nov. Act. Acad. Nat. Cur. 19: (Suppl.)
235. pl. 5, f. 1. 1843.
Polystitictus versatilis Sacc. Syll. Fung. 6: 244. 1888.
Funalia versatilis Murr. Bull. Torrey Club 34: 469. 1907.
Luzon: Province of Bataan, Lamao, February, 1904, Williams.
As represented in Philippine material this species is of a de-
cidedly daedaleoid type. Occasionally Trametes-like forms are
collected, showing that it may merge into the type of plant
described by Berkeley. These forms are rare, however, as also
are such as might tend toward the genus Polystictus.
Collected in the American tropics and the Philippines.
ELMERINA Bresadola
ELMERINA CLADOPHORA (Berk.) Bres. Hedwigia 51: 319. 1912.
Hexagonia cladophora Berk. Jour. Linn. Soc. Bot. 16: 47.
1878.
Luzon: Province of Nueva Ecija, Cabanatuan, October, 1908,
R. C. McGregor, Bur. Sci. 5272; Province of Rizal, Jalajala,
October, 1910, C. B. Robinson, Bur. Sci. 11925.
Reported only from the Philippines.
ELMERINA VESPACEA (Pers.) Bres. Hedwigia 51: 319. 1912.
Hexagonia vespacea Pers.; Gaudichaud, Bot. Voy. Uranie
170
Lenzites aebers Klotz. Linnaea 8: 480. 1833.
Polyporus lacerus Jungh. Flor. Crypt. Javae 1: 65. 1838.
Daedalea inconcinna Berk. Jour. Bot. 1: 151. 1842. Not
Deless.
Lenzites platyphylla Lév. Ann. Sci. Nat. Bot. ILL. 2: 179. 1844.
Daedalea aulaxina Lév. Ann. Sci. Nat. Bot. II1.2:197. 1844.
Hexagonia Molkenboeri Léy. Ann. Sci. Nat. Bot. III. 2: 200
1844.
Hexagonia macrotrema Jungh.; Fries, Symb. Myc. tot. 1851.
GRAFF: PHILIPPINE BASIDIOMYCETES—V 227
Hexagonia albida Berk. Jour. Linn. Soc. Bot. 16: 47. 1878.
Hexagonia Cesatt Berk.; Cesati, Myc. Born. 8. 1879
Daedalea pruinosa Ces. Myc. Born. 7. 1879. Not Léveillé.
Daedalea intermedia Berk. Jour. Linn. Soc. Bot. 18: 385.
1879.
Lenzites nivea Cooke, Grevillea 15: 94. 1886.
NeGROs: Province of Negros Oriental, Dumaguete, April,
1908, Elmer 9860.
Collected in Hawaii, Mauritius, Surinam, and Java.
HEXAGONIA Fries
HEXAGONIA APIARIA (Pers.) Fr. Epicr. Myc. 497. 1836.
Polyporus apiarius Pers.; Gaudichaud, Bot. Voy. Uranie
£72... 1826,
Hexagonia Koenigit Berk. Ann. Nat. Hist. II, 10: 379. 1853.
Luzon: Province of Bataan, November, 1909, H. M. Curran,
For. Bur. 19220.
Collected in Java, Ceylon, and the Rawak Islands.
Hexagonia Clemensiae (Murr.) comb. nov.
Inonotus Clemensiae Murr. Bull. Torrey Club 35: 401. 1908.
Polyporus Clemensiae Sacc. & Trott.; Saccardo, Syll. Fung.
#1: 272. 10912.
Minpanao: District of Davao, Lake Laneo, Camp Keithley,
September-October, 1907, Mary S. Clemens, on dead wood.
Not reported as yet outside the Philippine Islands.
HEXAGONIA CUCULLATA (Mont.) Murr. Bull. Torrey Club 31:
332. 1904.
Favolus cucullatus Mont.; Ramon de la Sagra, Hist. Phys.
Polit. Nat. Cuba 9: 378. 1841
Favolus curtipes Berk. & Curt. te Bot. Kew Gard. Miscel.
1: 234. 1849.
Hexagonia Taxodii Murr. Bull. Torrey Club 31: 332. 1904.
Luzon, Province of Isabella, Palanan Bay, June, 1913,
Escritor, Bur. Sci. 211
Collected in Ceylon, the West Indies, — America, and
eastern United States.
Hexagonia subrubida (Murr.) comb. nov.
Hapalopilus subrubidus Murr. Bull. Torrey Club 34: 470.
1907.
228 GRAFF: PHILIPPINE BASIDIOMYCETES—V
Polyporus subrubidus Sacc. & Trott.; Saccardo, Syll. Fung.
Zt: 276. 1912.
Luzon: Province of Bataan, Mount Mariveles, November,
1904, Elmer 691z, on dead tree branches.
This species should have been transferred to the genus
Hexagonia by Saccardo and Trotter instead of to the genus
Polyporus, to which it does not belong.
So far only found in the Philippine Islands.
FAVOLuUs Fries
FAVOLUS MEGALOPORUS (Mont.) Bres. Hedwigia 52: 74. 1912.
Polyporus megaloporus Mont. Ann. Sci. Nat. Bot. IV. 1: 124.
1854.
— princeps Berk. & Curt. Jour. Linn. Soc. Bot. 10:
321.
a cte Province of Nueva Vizcaya, Mount Umuguen,
May 19, 1909, M. Ramos, Bur. Sci. 8282.
Reported from French Guiana, Cuba, and Java.
FAVOLUS MULITPLEX Lévy. Ann. Sci. Nat. Bot. III. 2: 203. 1844.
Luzon: Province of Bataan, Balanja Mountains, January,
1910, H. M. Curran, For. Bur. 19252.
According to the collector’s note this fungus is edible and
used by the Negritos of Bataan Province as a food
This species has been previously reported from Brazil and
Java.
FAVOLUS SPATHULATUS (Jungh.) Lévy. Ann. Sci. Nat. Bot. Ill,
m.. 203: 18
Laschia <piibalot Jungh. Flor. Crypt. Javae 1:75. 1838.
Hymenogramme spathulata Sacc. & Cub.; Saccardo, Syll. Fung.
5: 653. 1887.
Minpanao: District of Davao, Mount Apo. September,
1909, Elmer 11572
Previously collected in Java.
GLoEoporus Montagne
Gloeoporus reticulatus (Fr.) comb. nov.
Auricularia reticulata Fr. Epicr. Myc. 555- 1836.
Gloeoporus conchoides Mont.; Ramon de la Sagra, Hist.
Phys. Polit. Nat. Cuba 9: 385. pl. 15, fd; 1841.
GRAFF: PHILIPPINE BASIDIOMYCETES—V 229
Thelephora dolosa Lév. Ann. Sci. Nat. Bot. III.2:209. 1844.
UZON: Province of Laguna, Mount Maquiling, February,
1912, P. W. Graff, Bur. Sci. 15952.
Collected previously in Brazil, Cuba, and Java.
GLOEOPORUS DICHROUS (Fr.) Bres. pas pat: 52: 74: “1612.
Polyporus dichrous Fr. Obs. Myc. 1: 125. 1815.
Boletus thelephoroides Hook.; Kunth, Syn. Plant. 10. 1822.
Boletus dichrous Spreng. Syst. Veget. 4: 475.
Polyporus nigro-purpurascens Schwein. Syn. Fung. Fikiner.
Bor. 360. 1831.
Polyporus pelleporus Secr. Mycogr. Suisse Champ. 73. 1833.
Polyporus thelephoroides Fr. Epicr. Myc. 473. 1836.
Gloeoporus candidus Speg. Fung. Guar. Pug. 1: 30. 1883.
Polystictus thelephoroides Sacc. Syll. Fung. 4: 284. 1886.
ZON: Province of Rizal, January, 1906, F. W. Foxworthy,
Bur. Sci..79. MINDANAO: District of Davao, Mount Apo,
May, 1909, Elmer 10747.
Collected in southern Europe, the American tropics, and
ava.
LascuiA Fries
LASCHIA MINIMA (Jungh.) Sacc. Syll. Fung. 6: 409. 1888.
Polyporus minimus Jungh. Flor. Crypt. Javae 1: 64. 1838.
MINDANAO: Subprovince of Butuan, March-July, 1911,
Weber 1229, on decaying woods.
Reported previously from Java.
AGARICINEAE
PLEUROTUS Fries
PLEUROTUS FLABELLATUS Berk. & Br. Jour. Linn. Soc. Bot.
11; 529. 1871
Luzon: Province of Benquet, Baguio, December 5, 1910,
E. Fénix, Bur. Sci. 12048, on dead wood.
Collected previously in tropical America, Ceylon, and South
Africa.
LENTINUS Fries
LENTINUS FUSCO-PURPUREUS Kalchbr. Grevillea 8: 153. 1880.
230 GRAFF: PHILIPPINE BASIDIOMYCETES—V
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sct. 1196.
This species, while allied to Lentinus Zeyheri Berk., seems
to be distinct.
Described from material collected on the Richmond River,
Australia, and not previously reported from elsewhere.
LENTINUS javANicus Lév. Ann. Sci. Nat. Bot. III. 5: 118.
1846.
Lentinus Decaisneanus Lév. Ann. Sci. Nat. Bot. III. 5: 120.
1846
40.
Lentinus infundibuliformis Berk. & Br. Jour. Linn. Soc.
Bot. 14: 42. 1875.
Luzon: Province of Bataan, Lamao, February 29, 1904, Cop-
5:
Previously reported from Java.
LentTINUS PoLycHRoUs Lév. Ann.Sci. Nat. Bot. III. 2:175. 1844.
Lentinus praerigidus Berk. Jour. Bot. Kew Gard. Miscel.
6: 132. 1854
Lentinus Kurzianus Berk. & Curr. Trans. Linn. Soc. II.
Ly 120. $1.20, 2.2. 1376.
Luzon: Province of Cagayan, pp March, 1909, H. M.
Curran, For. Bur. 16819.
Collected in Java, Sumatra, and eastern India.
LENTINUS SAJOR-CAJU (Rumph.) Fr. Epicr. Myc. 393. 1838.
Agaricus sajor-caju Rumph. Herb. Amb. 11: 125, pl. 56,
fig. 1. 1741-1755.
Luzon: Province of Bataan, Mount Mariveles, epee
1904, Merrill 3704.
Collected in Amboinia and Java.
LENTINUS TANGHINIAE Lévy. Ann.Sci. Nat. Bot. III.5:119. 1846.
MINDANAO: District of Davao, Todaya, March, 1904,
Copeland 5106.
Described from material collected in Madagascar.
LENTINUS VELUTINUsS Fr. Linnaea 5: 510. 1830.
Scleroma velutinum Fr. Epicr. Myc. 392. 1838.
Lentinus setiger Lév. Ann. Sci. Nat. Bot. III. 2: 176. 1844.
Lentinus brachatus Lév.; Zollinger, Syst. Verz. Ind. Arch.
17. 1854.
GRAFF: PHILIPPINE BASIDIOMYCETES—V 231
MinpDanao: District of Davao, Todaya, April, 1904, Copeland.
Collected previously in Brazil, Cuba, and Guiana.
LENZITES Fries
LENzITES AcuTA Berk. Jour. Bot. 1: 146. 1842.
Luzon: Province of Cagayan, March, 1909, Becani, For. Bur.
1671z.
The gills in this species have a somewhat wavy margin as
well as being notched and, while the majority of the longer
ones are forked, most of the shorter are distinct.
This is the only collection which has been made of this
fungus in the Philippines since Cuming’s collection of the type,
between 1836 and 1840. In the meanwhile the species has been
collected in northern India.
LENZITES PLATYPHYLLA Lévy. Ann. Sci. Nat. sting Til. 22470;
1844. Not Cooke.
Luzon: Province of Nueva Vizcaya, vicinity of Dupax,
January—April, 1912, R. C. McGregor, Bur. Sci. 14353. 20284;
Province of Bataan, Lamao, November, 1909, H. M: Curran,
For. Bur. 19188; Mount Mariveles, January, 1910, H. M.
Curran, For. Bur. 19244a, MINDANAO: District of Davao,
Lake Lanao, Camp Keithley, September, 1907, Mary S. Clemens.
Collected previously in Java and Guiana.
LENZITES STRIATA (Swartz) Fr. Epicr. Myc. 406. 1838.
Agaricus striatus Swartz, Prodr. Fl. Ind. Occ. 148. 1788.
Merulius striatus Lam. Encyc. Méth. Bot. 4: 127. 1797.
Daedalea striata Fr. Syst. Myc. 1: 334. 1821.
Luzon: Province of Bataan, Mount Mariveles, November
3-19, 1912, P. W. Graff, Bur. Sci. 19044, at an elevation of 350
m., on decaying and burned logs.
Previously collected in tropical America, (including the
West Indies,) Australia, and Borneo.
SCHIZOPHYLLUM Fries
SCHIZOPHYLLUM COMMUNE Fr. Syst. Myc. 1: 330. 1821.
Agaricus multifidus Batsch, Elench. Fung. 1: 124. f. 120.
1786.
Scaphophorus agaricoides Ehrh.; Nees, Hor. Phys. Berol. 94.
1820
232 GRAFF: PHILIPPINE BASIDIOMYCETES—V
Schizonia vulgaris Pers. Myc. Eur. 3: 14. 1828.
Schizophyllum alneum Schrot.; Cohn, Krypt. Flor. Schles.
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sci. 1195; Panay, Uoilo, January 2, 1904, Copeland 29.
A fungus of tatversal distribution living on woody hosts of
all sorts, both as a sap yte and asa facultative parasite. After
gaining entrance beneath the bark of living trees, through wounds,
this species is able to slowly advance through the tissues which
are apparently killed in advance of the progress of the fungus.
CREPIDOTUS Fries
CREPIDOTUS POGONATUS Kalchbr. Grevillea 9: 131. 1880.
Luzon: Province of Nueva Vizcaya, vicinity of Dupax,
January 14, 1913, R. C. McGregor, Bur. Sci. 20251, on decaying
twigs.
Previously collected in East Africa.
PANAEOLUS Fries
PANAEOLUS PAPILIONACEUS (Bull.) Fr. Epicr. Myc. 236. 1836.
Agaricus equinus Alb. & Schw. Consp. Fung. Lus. 3. 1805.
Agaricus campanulatus Linn. Sp. Plant. 1175. 1753.
Agaricus carbon Batsch, Elench. Fung. 6. 1783.
Agaricus papilionaceus Bull. Hist. Champ. France 561.
1791-1798.
Agaricus varius Pers. Icon. Des. Fung. 40. 1800.
Panaeolus campanulatus Berk. Outl. Brit. Fung. 175. 1860.
Copelandia papilionacea Bres. Hedwigia 53: 51. 1912.
Luzon: Province of Cagayan, Aparri, March, 1909, H. M.
Curran, For. Bur. 17143; vicinity of Manila, September, 1905,
Merrill 5008. Mutnpanao: District of Davao, March 5, 1904,
Copeland 344.
The formation of the new genus Copelandia, in which the
sole distinction from Panaeolus is the presence of cystidia,
is inviting for the future a confusion similar to that existing
in the genera Panus and Lentinus, where the only difference
exists in the presence or absence of notched gills. This character-
istic, even though at present confined to this one species, is
not a sufficient distinction for a generic character, or to insure
GRAFF: PHILIPPINE BASIDIOMYCETES—V 233
the possible acceptance of this as the type of a new genus.
It seems the wisest course, therefore, to continue as before with
the use of the Friesian name, at least till some character more
positive may be found to warrant the change.
Previously reported from the United States, Europe, South
Africa, southern Siberia, Ceylon, Borneo, and Australia as well
as the Phillippines. :
UNIVERSITY OF MontTAna,
MIssouLa, MONTANA
Occurrence of the pycnial stage of Puccinia Taraxaci*
Louise DospALL
On April 29, 1920, dandelions were found on University Farm,
St. Paul, heavily infected with rust. In addition to the fact that
this was the earliest date on record at the University of the
occurrence of Puccinia Taraxaci (Rabent.) Plow., this collection
was of particular interest because pycnia were found accompany-
ing the uredinia. As far as could be discovered from the lit-
erature, this stage had never before been collected in North
America, although both Plowright (1889) and Grove (1913)
had reported its occurrence in England, and Jacky (1909) in
Germany.
In the spring of 1921 a careful watch was kept for the first
appearance of dandelion rust to determine whether or not the
pycnial stage was common in this region. The first pycnia
were found on University Farm, April 17, 1921. Subsequently
they were found in various localities around St. Paul and Minn-
eapolis. If a careful search was made, the pycnial stage could
be found in almost any cluster of plants. A difference was
noted in the abundance of the rust during the two seasons in
which observations were made. In 1920 the pycnia were found
accidentally in two localities. In both cases the infection on
the plants was so conspicuous that the rust was observed while
one was walking hurriedly through the field. In 1921 no pycnia
could be found without the most diligent search.
Through correspondence with Dr. H. S. Jackson, it was
learned that the pycnial stage of Puccinia Taraxaci had been
collected at Brookville, Indiana, in April, 1918, by C. A. Ludwig,
and the specimens deposited in the Arthur Herbarium. This
specimen was compared with the Minnesota collection and the
rusts found to be identical. In spite of the fact that collections
are rare, it would seem that the occurrence of the pycnial stage
must be rather common.
The pycnia of Puccinia Taraxaci are amphigenous on yellow,
slightly swollen spots, which occur frequently along the midrib
*Published, with the approval of the Director, as Paper No. 303 of the
Journal Series of the Minnesota Agricultural Experiment Station.
235
236 DosDALL: PYCNIAL STAGE OF PUCCINIA TARAXACI
of the leaf but are found sometimes on the blade. They are
reddish brown in color, subglobose in shape, about 9oyv. in diameter
and 76. in height. Ostiolar filaments are present. Uredinia
appear on the same spots soon after the pycnia.
Midwinter botanizing in southern Arizona
Epwin B. BARTRAM
As a large part of the literature relating to the desert flora
of Southern Arizona is not readily accessible to any but pro-
fessional botanists the following brief survey of the recognizable
winter plants in the region tributary to Tucson may prove
interesting and helpful to those fortunate enough to make the
ce mage a of this fascinating country in the early months
of the y
From a ee point of view the area is obviously more
closely related to the Mexican state of Sonora than it is to any
of our possessions so that many of its most characteristic plants
are not represented at all in Wooton and Standley’s invaluable
Flora of New Mexico.
The preparation of specimens in this dry sunny climate is a
simple operation; the surrounding mountains, or at least their
foothills, are easy of access; and, as the representation of Arizona
plants in most of our larger eastern herbaria is quite meagre,
the ambitious collector finds an almost unlimited field in which
to work. A surprising variety of flowers is evident even at this
season and many plants whose period of bloom is past are re-
cognizable by some peristent vestige of fruit or flowers.
Apart from such unique and unfamiliar plants as the sahuaro,
chollas, candle bush, prickly pears and palo-verde, which claim
the undivided attention of the newcomer until the novelty and
charm of the first impressions are to a certain extent dulled by
familiarity, the most striking feature of the desert vegetation is
the isolation of various clearly defined plant communities. It is
all desert, to be sure, but the wide gravel plains over which Covtllea
glutinosa is dominant and the barren spurs and talus slopes,
sun baked, arid and almost devoid of plant life, excepting the
hardiest drought-resisting species that have become inured to
this environment, seem to resolve themselves readily into a
series of barriers that from necessity restrict the distribution of
many species to quite definitely circumscribed limits.
Distances in Arizona are proverbially deceptive, but we were
hardly prepared to learn that the lower slopes of the Santa
Catalina Mountains, which looked so enticingly near, were at
237
238 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
least fifteen miles away instead of under five miles as we had
fondly imagined. The horses are willing and capable of covering
thirty miles or more in a day without discomfort, but the first
days experience proved conclusively that the one indispensable
article of equipment was a good canteen; for, excepting a few
canyons in the Catalina Range reaching well back into the forested
and snow-covered slopes, water is never available. The higher
peaks are heavily snow-capped during most of the winter months,
although snow rarely descends to the level of the plains. After
a rainy night in January it is not unusual to see the snow line
across the slopes as straight as though it had been drawn with
a ruler. In the sheltered canyons where snow rarely falls and
never persists we found many flowering plants in early January
which were materialy increased in number and variety with
each succeeding week. Many plants are distributed generally
over entire ranges but in each canyon, walled in by the steep
arid intervening ridges, we found some species that were wanting
in every other locality we visited.
Sabino Canyon in the Santa Catalina Mountains, distant
some sixteen miles from Tucson, is one that we happened to
reach quite frequently and will serve nicely as an example of
these interesting localities. Leaving early we drive across the
plains through miles of pungent greasewood with its curiously
folded leaves and ‘bright yellow flowers, which even now are
giving way to the little felted spheres of fruit. Thickets of
cholla, six or eight feet high, varying in color from pale green to
ruddy brown and brightened with pendant clusters of orange
colored fruit, glow in the early sunlight. Cirsium neomexicanum,
Lesquerella Fendleri, and Baileya multiradiata are in bloom along
the roadsides, the stiff wands of old fruit bend over colonies of
gray-green Atriplex canescens, and here and there an occasional
Yucca elata lifts its brown panicle of old fruit above the shrub
level. In a broad belt along the base of the mountains the
sahuaro gives a grass green tinge to the foothills ten miles away
where the arroyos are marked by wavering lines of mesquite and
desert willow. The hot rocky slopes flanking the mouth of
the canyon are spotted with Opuntia Bigelovii, glistening like
spicules of glass in the brilliant sunshine and undoubtedly the
most vicious of all the chollas. A carpet of detached joints
covers the ground about every plant, and woe to the unwary
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 239
walker who steps unguardedly into these precincts. The spines
penetrate shoe leather like a steel needle and refuse to be with-
drawn. Nature guards her wards well in these desert places and
the animals, profiting by the example, guard their dens with a
pile of these bristling joints, carried one by one at a cost easy
to be reckoned by anyone unfortunate enough to have brushed
against a stem in an unguarded moment.
Along the creek issuing from the canyon are many waifs
from the higher zones, such as Juniperus scopulorum, Yucca
baccata, Quercus Emoryi, Q. oblongifolia and Q.hypoleuca, but it is
among the rocks on sheltered slopes in the canyon that we find
the earliest flowers. Herethe fairy dusters, Calliandra eriophylla,
are just spreading their delicate balls of pink bloom, and clumps
of Lesquerella purpurea are established in many congenial crev-
ices. Hermannia paucifiora with the flowers well concealed
among the matted woody stems is easily overlooked, and a form of
Anislotus puberulus with showy golden yellow flowers and nearly
leafless stems helps to brighten the otherwise barren ledges.
Notholaena Lemmoni grows abundantly among the rocks of a
dry sunny slope, while the shaded crevices and fissures give
harbour to such species as Pellaea mucronata, Cheilanthes Lind-
heimert and various mosses, among which Tortula inermis, T.
ruralis, Barbula Manniae and B. chloronotus are the most in
evidence. Where almost any other living thing would wither and
perish Agave Parryi and Dasylirion Wheeleri spread over the
ledges and boulders in flourishing clumps that lend a charac-
teristic touch to this unique landscape.
The region about Picture Rocks was another productive
locality that proved interesting enough to warrant trips at
intervals of about ten days. Some twelve miles west of Tucson,
near the far end of the Tucson Mountains in a broad gully
fronting the old Indian pictographs, from which the place takes
its name, Anemone sphenophylla was just coming into bloom on
January twelfth, white clusters of Dryopetalon runcinatum
brightened many of the damp rock fissures, the trim little
Thysanocarpus amplectens grew sparingly in nearly every
depression where sufficient soil had accumulated to support a
short span of life during the season of winter rains, while various
winter annuals, such as Lepidium lasiocarpum, Draba cuneifolia
and Amsinckia Menziesii, were scattered sparingly in the shelter
240 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
of shrubs or rocks wherever a bit of moisture and shade was
obtainable. Along the dry arroyos leading off through the
mountains Anislotus brachycarpus and A. trispermus were just
opening their first flowers, and on the dry flanking talus slopes
the omnipresent Encelia farinosa in full bloom was temporarily
forgotten in the pleasure incident to finding a patch of Polygala
macradenia with inconspicuous little purple flowers barely
visible even at close range.
similar association of species, including Dryopetalon,
Anemone and Thysanocarpus, was observed on the shaded side
of a steep ridge at Robles Pass, where scattered pockets of soil
had been formed by the disintegrating mosses and lichens, and
no doubt a more thorough survey of the region would disclose
many similar localities; but the complete isolation of these
stations, one from another, is the one factor that seems to be
repeatedly emphasized.
Theorizing is always interesting, often productive, but some-
times misleading, yet in the light of these facts one can hardly
reject the inference that at some comparatively recent time the
forms of plant life, not fitted to thrive in the arid gravel and
rock formations of the desert as we find it today, were more
widely distributed than they are now, and by progressive changes
have been isolated by the rising tide of desert vegetation.
Such a condition is plain in outline but difficult to translate
into terms that are applicable when the available data are so
meagre. Inequality in the distribution of rainfall, the processes
of erosion and disintegration and variations in climatic conditions
have been some of the contributing causes to the spread and
contraction of the various plant communities as we now know
them, but to what extent these influences have operated and in
what manner the distribution has been modified by complicated
cross currents of action and reaction are queries that may be
more satisfactorily answered at some future time, when our
knowledge of the subject will be more intimate and compre-
hensive than it is now.
The following list of species, collected between December
23d, 1919, and January 26th, 1920, may be considered as
fairly representative of the region within a radus of fifteen
miles from Tucson but is necessarily incomplete. Many of the
plants listed were in flower and nearly all of them provided with
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 241
fruit in a sufficiently good state of preservation to make them
easily recognizable.
For assistance in the determination of many of the species
I am much indebted to the kindness of Dr. Paul C. Standley.
CHEILANTHES LINDHEIMERI Hook. Frequent on arid ledges and rocky
slopes.
CHEILANTHES MYRIOPHYLLA Desv. Less common than the preceding,
EILANTHES PRINGLEI Davenp. The finely dissected delicate fronds
of this species distinguished it at once from its associates in this region. In-
frequent in shaded rock clefts on the lower slopes.
NTHES WricHTI Hook. Infrequent but re recognized by
the smooth green pinnules and lustrous pee sti
AENA HooKerr D. C. Eaton. Rather common among rocks
on arid ‘hillsides The dahil outline of the fronds and the bright
yellow color of the lower surface are unmistakable marks.:
NOTHOLEANA SINUATA (Sw.) Kaulf. Frequent among dry rocks on the
lower mountain slopes.
OLAENA SINUATA INTEGERRIMA Hook. More widely distributed
than the cuca form of the sp
NOTHOLAENA BONARIENSIS Ot C. Chr. Rare among rocks near
the mouth of Sabino Canyon.
NoTHOLAENA LEMMoNniI D. C. Eaton. Abundant on a dry rocky slope
in the mouth of Sabino Canyon but not found elsewhere.
PELLAEA MUCRONATA D. C. Eaton. Ledges a crevices in the re
This seems to be the commonest fern of the regio
ELAGINELLA ARIZONICA Maxon. Arecently described species forming
flat green mats among rocks in the arroyos
SELAGINELLA RUPICOLA Underw. The grayish color and erect habit
are good field marks for this species. Near the mouth of Sabino Canyon
it forms dense turfy mats over the dry ledges
ERUS SCOPULORUM Sarg. Bank of stream near ov mouth of
Sabino iano pudeane a stray from the higher ridge
EPHEDRA TRIFURCATA Torr. Abundant along washes in the gravel
plains
eeceak ANTISYPHILITICA Meyer. Rare in an arroyo near Pictu
Rocks. The slender flagellate stems give this plant a very different as
from the more common, stiffly erect, E. trifurcata.
Epuepra viripis Coville. Collected but once in the dry foothills at
Robles Pass
ANDROPOGON SACCHAROIDES Sw. Frequent on the plain and foothill.
HETEROPOGON conTOoRTUS (L.) Roem. & Schult. More confined to the
foothill slopes.
242 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
CHAETOCBHLOA MACHROSTACHYA (H. B. K.) Scribn. & Merr. Rare on
talus aN at Robles Pas
CHAETOCHLOA virivIs (L.) Scribn. Introduced in fields and waste places.
ARISTIDA ADSCENSIONIS L. Frequent on hillsides in the Tucson Mount-
ains.
sansa Wricuti Nash. Uncommon on dry slopes in the Tucson
Mountain
ArIsTIDA BROoMoIDES H. B. K. Common in the foothills. Spikelets
developing through the winter.
ARISTIDA DIVERGENS Vasey. Dry slopes in the mouth of Sabino Canyon.
Muu RGIA DUMOSA Scribn. A unique species ee to ledges
along ie frail in Sabino Canyon at about 3000 feet eleva
MUHLENBERGIA MICROSPERMA (D C.) Kunth. Rare in oe oS foothills.
MUHLENBERGIA PoRTERI Scribn. Common in the shelter of shrubs
on the gravel plains.
EPICAMPES RINGENS Benth. Slopes in the mouth of Sabino Canyon.
OUTELOUA FILIFORMIS (Fourn.) Griffiths. Flowering plants collected
in Sabino Canyon on Janua d
BovuTELovuA RoTHROCKII Vasey. The dark brown spikelets persist
well mee the winter.
LOUA Parryi (Fourn.) Griffiths. This and the preceding species
are siisndant near Robles Pass.
TRICHLORIS FASCICULATA Fourn. Roadsides south of Tucson.
DASYOCHLOA PULCHELLA (H. B. K.) Willd. Generally distributed over
the plains.
Ko LERIA CRISTATA so Pers. Along streams in the foothills of the
Santa Catalina Mountai
ERAGROSTIS MEGASTACHYA (Koel.) Link. A weed of waste places.
Poa BIGELOVII Vasey & Scribn.
Rocks.
CYPERUS CYRTOLEPIS Torr. & Hook. Edge of stream east of Pima
Canyon.
Juncus NODOSUS L. Edge of stream in Pima Canyon.
Yucca ELATA te Single plants scattered over the plains but
nowhere abun
Confined to damp ledges near Picture
Yucca paccata Torr. Thickets in the mouth of Sabino Canyon. Ap-
parently washed down from the higher levels.
NOLINA MICROCARPA S. Wats. With the preceding.
DASYLIRION WHEELERI S. Wats. On dry ledges in the foothills.
DIPTEROSTEMON PAUCIFLORUS (Torr.) Rydb. Sparingly in flower along
an arroyo near Picture Rocks.
AGAVE Parryi Engelm. On dry ledges in the mouth of Sabino canyon.
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 243
PoPuLus —— (S. Wats.) Sarg. Catkins fully developed and
falling a late January.
RCUS ARIZONICA Sarg. Scattered along the creek near the mouth
of Sabino Can
QUERCUS eae Torr. With the preceding but more abundant and
larger
QUERCUS HYPOLEUCA Engelm. Sterile but easily recognized by the
leathery leaves densely white woolly beneath.
QUERCUS OBLONGIFOLIA Torr. This and the preceding species of oaks
are all confined to the lower end of Sabino Canyon, where they seem to exist
as waifs carried down by the stream from higher levels in the region about
Mt. Lemmon
CELTIS PALLIDA Torr. Frequent along arroyos in the Tucson Mountains.
PARIETARIA OBTUSA Rydb. Localized on a shaded ledge near Picture
Rocks.
PHORADENDRON MACROPHYLLUM (Engelm.) Cockerell. Abundant on
Populus in the plains and foothill
PHORADENDRON CALIFORNICUM ee The abundant small red berries
and bare stems make a very attractive combination. Chiefly on Mesquite in
the plains
ERIOGONUM POLYCLADON Benth. Recognizable but long past flowering.
ERIOGONUM PINETORUM Greene. Long past flowering but leaves and
fruit ere enough for identification
Erioconum Wricuti Torr. Widely distributed over arid rocky
slopes a “he foothills.
RUMEX HYMENOSEPALUS Torr. Common in sandy areas throughout
the —
PoL UM PUNCTATUM Ell. Edge of stream in the foothills of the
Santa Caidting ntain
ATRIPLEX CANESCENS ee. Nutt. Several forms of this species
are abundantly over the plains
ATRIPLEX ROSEA L. Young plants in the sandy bed of an arroyo near
Picture Rocks.
CHENOPODIUM PAGANUM Reichenb. Sparingly introduced in waste places.
AMARANTHUS PANICULATUS L. A common weed of fields and waste
places.
WEDELIELLA INCARNATA (L.) Cockerell. Flowering sparingly in sunny
talus slopes in the foothills.
HESPERONIA RETRORSA (Heller) Standl. Dry slopes near Picture Rocks.
COMMICARPUS SCANDENS (L.) Standl. Localized in rock clefts along
an arroyo west of Tucson.
ANEMONE SPHENOPHYLLA Poepp. Frequent among rocks along arroyos
in the foothills In full flower by mid-January.
244 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
Ciematis DrumMonpi Torr. & Gray. Frequent on the plains west of
Tucson. :
ESCHSCHOLTZIA MEXICANA Greene. A few flowers on sheltered banks
in January
YSANOCARPUS AMPLECTENS Greene. Locally abundant in damp
pockets near Picture Rocks and at Robles Pass. Sparingly in flower on
January 12th but plentiful and in mature fruit ten days later. The curious
disk-shaped pods with perforated wing margins mark this off at once from
all other crucifers of the region.
STREPANTHUS ARIZONICUS S. Wats. Frequent in the shade of bushes
near Picture Rocks.
LEPIDIUM HIRSUTUM Rydb. Infrequent on lower slopes of Santa Cat-
alina Mountains.
LEPIDIUM LASIOCARPUM Nutt. Common in sandy plains and dry washes.
LESQUERELLA FENDLERI (Gray) S. Wats. Flowering early along dry
roadsides and on open slopes in the foothills.
LESQUERELLA PURPUREA (Gray) S. Wats. An attractive early flower
of rock crevices in the foothills and lower slopes of the canyons. In vigor-
ous tufts the stems sometimes attain a length of nearly two feet.
DRABA CUNEIFOLIA Nutt. Widely distributed on shaded knolls and
banks along the a
ARABIS PERENNANS S. Wats. The purple medics Bike quite a touch of
coior to the ledges along the arroyos west of Tuc
BRASSICA NIGRA (L.) Koch. Occasional in cultivated fields.
SOPHIA OCHROLEUCA Wooton. Frequent in sheltered shady spots.
DRYOPETALON RUNCINATUM Gray. Rather local in damp shaded
tock pockets. The delicate white flowers form a pleasing contrast with their
rough arid setting.
SEDUM GRIFFITHSII Rose. Ledges in Sabino Canyon.
FENDLERA RUPICOLA Gray. Ledges in Sabino Canyon. Leaves and
old a persiste
MICRPOHYLLUS Gray. Frequent on open rocky slopes
HILAD:
in * foothills. “ sheltered places the flowers were opened in profusion by
uary.
mid Janu
PLATANUs WricutTi1 S. Wats. In the mouth of Sabino Canyon.
VAUQUELINA kee dela (Torr.) Sarg. Ledges in Sabino Canyon at
about sei feet elev
ANDRA ERIOPHYLLA Benth. One of the most attractive of the
early suihes flowers. Abundant on open rocky slopes in the foothills.
ACIA CONSTRICTA Benth. Frequent on the plains. Pods more per-
sistent than A. Greggii.
Acacia GreGccit Gray. This and A. constricta are character istic
thorny shrubs of the gravel plains.
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 245
PROSOPSIs GLANDULOSA Torr. This is the main source of firewood for
the region.
AcuaN JAMEsiI (Torr. & Gray) Kuntze. Pods persistent through the
winter
sta CovesiI Gray. Scattered sparingly over the open gravel plains.
Pods and leaves persistent.
HOFFMANSEGGIA DENSIFLORA (Benth.) Gray. On sandy plains. Pods
persistent.
PARKENSONIA MICROPHYLLA Torr. A thorny shrub or small tree of
the foothills and adjacent plains. On sheltered sunny slopes a few flowers
were in evidence by late January
RCIDIUM TORREYANUM Sarg. A low nearly leafless shrub occuring
sparingly along washes in the plains.
KRAMERIA GLANDULOSA Rose & Painter. A oe flowerless shrub
of the plains with soft silky oe young g
NISLOTUS BRACHYCARPUS (Benth.) Rydb. The broad leaflets and
villous sagminceds ‘Sidebee this Fes sharply from the following where
they grow toget
ANISLOTUS TRISPERMUS (Greene) Woot. & Standl. With the preceding
in a sandy bed of an arroyo near Picture Rocks. Just coming into flower
ANISLOTUS PUBERULUS (Benth.) Woot. & Standl. What seems to be
a form of this species was in full flower on the ledges of Sabino Canyon at
about 3000 feet elevation.
OSELLA Parry! (Torr. & Gray.) Heller. This inconspicuous little
ae is peer typical of arid rocky hillsides in the foothills.
ROSELLA GREGG (Gray) Heller. Abundant on rocky spurs and slop2s
near Agua Calgiente but not observed elsewhere
Nissoiia Scuotti (Torr.) Gray. Pods and leaves persistent. Frequent. —
AsTRAGALUS NuTTALLiANus DC. Local on the plains. In flower
in January.
CRACCA TENELLA Gray. Localized on dry sunny ledges near Robles Pass.
GaALacTIA WricHtu Gray. Twining over low bushes in the foothills.
Leaves and pods persistent.
OLNEYA TESOTA Gray. Foothill slopes and nearby plains.
ERODIUM CICUTARIUM (L.) L’Her. Abundant in waste places and
on open sandy
ERopiIuM TEXANUM Gray. Gravelly Knolls near Picture Rocks.
LEA GLUTINOSA (Engelm.) Rydb. The most abundant and char-
os eal of the plains.
KOEBERLINIA SPINOSA Zucc. Frequent on knolls and in thickets in
‘the plains.
JANuSIA GRACILIS Gray. Twining about bushes on open rocky hillsides.
246 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
RUTOSMA PURPUREA Woot. & Standl. Frequent on banks of the
arroyos west of Tuc
POLYGALA MACRADENIA Gray. Confined to a rocky hillside near Picture
Rocks. .
UPHORBIA CAPITELLATA Engelm. Forming —— showy mats on ledges
and in crevices along the arroyos west of Tucson
EUPHORBIA PEDICULIFERA Engelm. A ae with densely glandular
pubescent stems and leaves is not uncommon
EUPHORBIA POLYCARPA Benth. This and the two preceding species
of the genus are freely distributed over the arid ridges and talus slopes. They
all belong to the group of perennials with entire pubescent leaves and vary
considerably in habit and appearance.
DITAXIS NEOMEXICANA (Muell. Arg.) Heller. The lax straggling habit
of this species is very different from the stiff erect form of D. sericophylla.
DITAXIS SERICOPHYLLA (Gray) Heller. With the preceding on rocky
slopes at Robles Pass.
CALLITRICHE PALUSTRIS L. In shallow water at the mouth of Sabino
Canyon.
SIMMONDSIA CALIFORNICA Vutt. A striking cea: bush of the dry
plains and goers with leathery leaves and yellow flowers in drooping
axillary clusters.
Doponaka viscosa L. Abundant along the stream bank in the mouth
of Sabino Canyo
CONDALIA SPATHULATA oe: Banks of arroyos near the foothills.
Leaves and fruits persisten
RHAMNUS TOMENTELLA one Ledges in Sabino Canyon at about
3000 feet elevat
HIBIscus else Harv. Flowering sparingly through the winter.
: HIBIscus DENUDATUS Benth. The pale almost leafless stems and
cae tinted pink flowers give this plant an attractive personality. Rare
n slopes at Robles Pass
GAYOIDES CRISPUM (L.) Small. Frequent on rocky slopes.
ABUTILON INcANUM (Link) Sweet. Common on rocky hillsides. Leaves
and carpels persistent.
SPHAERALCEA AMBIGUA Gray. A polymorphous species widely scattered
over the region. Flowezs white or pink and leaves varying greatly in outline.
' SPHAERALCEA GROSSULARIAEFOLIA York. On rocky slopes at Robles Pass.
ERMANNIA PAUCIFLORA S. Wats. Confined to rocky slopes near the
mouth of Sabino Canyon.
,_ AYENIA PUSILLA L. Frequent on the dry gravel plains.
FOUQUIERIA SPLENDENS Engelm. Although the | ll developed
in sunny sheltered places no flowers were in evidence by the end of Jan uary.
OpunTIA FULGIDA Engelm. Common on the plains.
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 247
OPUNTIA ARBORESCENS Engelm. Forming impenetrable thickets on
the plains.
OPUNTIA LEPTOCAULIS DC. The common _ slender stemmed cholla
of the regio
parent: BicELovil Engelm. Frequent on arid rocky slopes in the
foothills.
OPUNTIA CHOLOROTICA Engelm. & Bigel. On ledges near the mouth of
Sabino Canyon
OpuntTiA TouMEYI Rose. Frequent on the gravel plains.
MAMILLARIA GRAHAMI Engelm. Frequent in small colonies among
rocks in ee foothills. The diminutive scarlet pipe-shaped fruits are very
attractive
Ecu ACTUS WISLIZENI Engelm. Eola sparingly over the foot-
hill sie een plants are well over a meter eight.
ECHINOCEREUS FENDLERI (Engelm.) Riimpl. Forming clumps in
gravel plains near the foothills.
CEREUS GIGANTEUS Engelm. The sahuaro well merits the reputation of
being one of the most unique and striking elements of the desert vegetation.
Daucus PpusmLLus Mx. Scarce on shaded knolls in the washes.
PLUMBAGO SCANDENS L. In the foothills of the Santa Catalina Moun-
tains. Leaves and fruit persistent.
FORESTERIA PHILLYREOIDES Torr. This is a Mexican type and Dr.
Standley informs me they have but one other specimen from the United
States, collected at Tucson by Pringle. The staminate flowers were fully
developed by December 3oth and a few leafy shoots supplied the foliage
characters. On rocky slopes along the Ajo road west of Tucson.
FRAXINUS ATTENUATA Jones. Inthe mouth of Sabino Canyon.
MENODORA SCABRA Gray. On arid slopes of the Tucson Mountains.
HAPLOPHYTON CImMicIpDUM A. DC. Rare on rocky talus slopes at
Robles Pas
ASCLEPIAS LINARIA Cav. Frequent on ledges in the lower slopes of
the Santa Catalina Mountains
PHILIBERTELLA LINEARIS Geass Twining over bushes on dry rocky hill-
sides
EVOLVULUS LINIFOLIUS L. An inconspicuous little perenial with fugitive
flowers. Frequent among rocks and dry ledges in the foothills.
JACQUEMONTIA PRINGLEI Gray. Not uncommon on rocky hillsides.
Leaves and capsules persistent.
ECTOCARYA LINEARIS (R. & P.) DC. Locally abundant on a gravel
shelf near Picture Rocks.
AMSINCKIA Menzigesu (Lehm.) Nels. & Macb. Flowering with Thys-
anocarpus and nue on shaded ledges at Picture Rocks.
248 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
VERBENA WricHTU Gray. This is one of the showiest of the early
flowers in the foothills, where it blooms profusely in January.
VERBENA XUTHA Lehm. Rare and local on dry slopes of the Tucson Mts.
Lippra Wricutu Gray. Rocky slopes in the foothills. Leaves and
ae persis
eae VULGARE L. A very common weed of roadsides and waste
places
ea Ay Emorvyi (Torr.) Kuntze. Flowering oo the winter
and very characterisric of arid rocky slopes in the foothilis
HEDEOMA OBLONGIFOLIA (Gray) Heller. Ledges in the mouth of Sabino
Canyon
Scsuoua NANA (Torr.) Greene. In rock crevices along arroyos west
of Tucson. The leaves and fruit of this and the preceding are persistent
through the winter.
SALVIA PINGUIFOLIA (Fernald) Woot. & ae eye rocky hillside at
Robles Pass. Buds well developed but not in flow
NICOTIANA TRIGONOPHYLLA Dunal. A common plant of dry ledges in
the foothills. In flower throughout the winter.
NICOTIANA GLAUCA Graham. An introduced shrub of waste places.
Lycrum Cooperi Gray. Frequent on rocky slopes.
Lycium Fremontit Gray. Ledges at the mouth of Sabino Canyon.
In full aiber on January 6th
LycIUM PARVIFLORUM Gray. A thorny straggling shrub of the gravel
plains.
Lycium Torrey Gray. Confined to the slopes near Picture Rocks.
PHYSALIS IXOCARPA Brot. Infrequent in the Tucson Mountains.
RHINUM ANTIRRHINIFLORUM (Willd.) Hitchc. On slopes of the
Tucson Nacsa. Leaves and capsules persistent
SCROPHULARIA PARVIFLORA Woot. & Standl. istics sparingly on
January soit in the mouth of Pima Canyon, Santa Catalina Mountains.
PENTSTEMON SUPERBUS A. Nels. Sandy washes in the foothills. Flow-
ering Feeehy in January.
MIMULUS —— Don. Shallow water in a brook east of the
mouth of Pima Cany
MECARDONIA PEDUNCULARIS 2 gnienrseiil — Abundant in a brook
east of the mouth of fo —— to flower and mature
fruit throughout the win
STEMODIA DURANTIFOLIA (L.) Sw. In shallow water with the two
gia
Dia um TorreEyi (Gray) Woot. & Standl. Arid slopes near Agua
Caliente. "The large cordate bracts, fading white and papery. give this plant
a unique appearance in the field.
BARTRAM: BOTANIZING IN SOUTHERN ARIZONA 249
CARLOWRIGHTIA ARIZONICA Gray. Leaves and capsules persistent
enough for identificatio
SIPHONOGLOSSA LONGIFLORA (Torr.) Gray. Frequent on arid slopes
west of Tucson. Leaves and capsules persistent.
RUELLIA TUBEROSA OCCIDENTALIS Gray. Infrequent near Picture Rocks-
Old capsules persistent.
CHILOPSIS LINEARIS (Cav.) Sweet. Along arroyos in the foothills-
TENOLOBIUM INCISUM Rose & Standl, Rocky ledges and crevices
among the foothills. The persistent leaves and showy yellow capsules make
this a clean and attractive plant throughout the winter.
M STELLATUM Kellogg. This plant makes considerable growth
from a diffuse woody base during the winter, but there was no evidence of
bloom by January 30th. Frequent on ledges and rocky hillsides.
SAMBUCUS MEXICANA Presl. oe along water courses and in
cultivation. In bloom by mid-Janua
PTILORIA PAUCIFLORA (Torr.) Raf. A common plant of the arid plains
and foothill blooming through ea winter
PEREZIA WricutTi Gray. In full eect on January 21st. Infrequent
on arid rocky slopes near Picture Rocks.
TRIXIS CALIFORNICA Kellogg. In full flower during January. Abundant
and quite showy among rocks in the foothills.
HYMENOCLEA MONOGYRA Torr. & Gray. A tall slender weed-like
shrub with persistent filiform leaves and winged fruit. In the mouth of Pima
Canyon.
FRANSERIA DELTOIDEA Torr. One of the characteristic plants of dry
gravel . near the foothills. Buds well developed by January 30th
but no flow
pasipie SOLIDAGINIFOLIUM Gray. Ledges at the mouth of Sabino
Canyon
LEOSANTHUS BACCHARIDEUS (Gray) Kuntze. Infrequent on ledges
near the mouth of Pima Canyon. Leathery leaves and flower heads very
persistent.
COLEOSANTHUS COULTERI (Gray) Kuntze. Common on rocky hillsides.
CoLEOSANTHUS WRIGHTII (Gray) Britton. Ledges in the mouth of
Sabino Can
Se ccs coryMBosuM DC. Confined to the mouth of Pima
Canyon
GUTIERREZIA GLOMERELLA Greene. Leaves and old heads persistent.
Frequent on the plains.
SIDERANTHUS AUSTRALIS (Greene) Rydb. Common on the gravel plains.
coma Hartweci (Gray) Greene. Forming dense clumps on ridges
and knolls in the gravel plains
CHRYSOMA LARICIFOLIA (Gray) Greene. Rock clefts and ledges in the
arid foothills. Flowering casually through the winter months.
250 BARTRAM: BOTANIZING IN SOUTHERN ARIZONA
BACCHARIS GLUTINOSA Pers. Common along the banks of Rillito Creek.
BACCHARIS SAROTHROIDES Gray. Abundant on the sand bars and
banks = Rillito Creek.
ERIGERON DIVERGENS Torr. & Gray. This is apparently an abnormal
winter form of the species but very constant throughout the region. It is
scattered freely over the rocky foothill slopes and flowers during January.
GNAPHALIUM WriGHTII Gray. Frequent on arid rocky slopes.
PARTHENIUM INCANUM H. B. K. A common low shrub of the foothills.
In good conditions through the winter,
CRASSINA PUMILA (Gray) Kuntze. Occurring sparingly on dry gravelly
ENCELIA FARINOSA Gray. The distinctive gray color of many of the
desert slopes is derived from the lustrous silvery leaves of this species. The
showy yellow flowers are locally abundant, even in January, and it ma
well be numbered among the most satisfying and attractive plants of the
region.
LEPTOSYNE ARIZONICA Gray. Bank of a stream in the mouth of Pima
Canyon
Poi: ROPHYLLUM JUNCIFORME Greene. A rush-like plant of ledges and
rocky slopes in the foothills.
DYSODIA POROPHYLLOIDES Gray. Rare on ledges near Picture Rocks.
PsILOSTROPHE CoopErI (Gray) Greene. Fresh flowers among the
)
persistent papery old blooms are found sparingly in January. Frequent on
the gravel plains.
BAILEYA MULTIRADIATA Harv. & Gray. Frequent and showy
sandy plains, especially along wiathaldes: Flowering freely by a January.
PERITYLE DissEcTA Gray. An attractive little plant with curiously
cleft leaves and persistent involucres. In rock clefts near the mouth of
Pima Canyon.
BAHIA DEALBATA Gray. Arroyos in the Tucson Mountains.
ELENIUM MONTANUM Nutt. Localized along the bank of a stream
east of the mouth of Pima Canyon
ARTEMISIA ALBULA Wooton. Arid foothill slopes Abundant near
Agua Caliente.
ARTEMISIA Britton Rydb. Ledges in Sabino Canyon.
BEBBIA JUNCEA (Benth.) Greene. An attractive slender stemmed
plant of the sandy plains.
ENECIO LEMMONI Gray. Flowering rather sparingly on rocky slopes
in the foothills.
CIRSIUM NEOMEXICANUM Gray. Infrequent along roadsides in the
gravel plains.
BUSHKILL, PENNSYLVANIA
Two new genera of Cactaceae
N. L. Britton AND J. N. Rose
In our recent study of the genus Echinocactus, we have
segregated a number of new genera, and restored several previ-
iously published, among which are Malacocarpus, Gymnocalycium,
and Discocacius, all composed of species native to South America.
Among the North American segregates are the two described
below, which do not belong even to the same subtribe as Echi-
nocactus.
, THELOCACTUS (Schumann)
Echinocactus subgenus Thelocactus Schumann, Gesamtb.
Kakteen 429. 1808.
Cacti of medium size, one-jointed, solitary or caespitose,
globose or somewhat depressed, spiny, often densely so; ribs
few, low or even indistinct, often spiraled, divided into large
tubercles, the flowering ones more or less grooved above; flowers
from near the center of the plant, borne on very young tubercles,
rather large for the subtribe, campanulate, diurnal; scales on
the ovary usually few, their axils naked; fruit, so far as known,
dry, dehiscing by a basal pore; seeds numerous, black, finely
tuberculate, with a large basal hilum.
Type species: Echinocactus hexaedrophorus Lemaire.
The generic name refers to the tubercled ribs.
1“ Thelocactus hexaedrophorus (Lemaire) comb. nov.
Echinocactus hexaedrophorus Lemaire, Cact. Gen. Nov. Sp.
27. 1839.
2.‘ Thelocactus bicolor (Galeotti) comb. nov.
Echinocactus bicolor Galeotti in Pfeiffer, Abbild. Beschr. Cact.
a: ht. 25. 4648
3 ‘Thelocactus lophothele (Salm-Dyck) comb. nov.
Echinocactus lophothele Salm-Dyck, Allg. Gartenz. 18: 395.
1850.
’ NEOLLOYDIA gen. nov. |
all, more or less caespitose plants, fibrous-rooted, cylindric,
one-jointed, densely spiny, tubercled; tubercles borne on spiraled
ribs, grooved above; radial spines numerous, widely spreading ;
251
252 +.Britron & Rose: NEW GENERA OF CACTACEAE
central spines one to several, much stouter and longer than
he radials; flowers large, pink or purple, subcentral from the
axils of nascent tubercles, their segments widely spreading;
fruit compressed-globose, dull colored, thin-walled, becoming
papery, , with few scales or none; seeds globose, black, dull,
tuberculate-roughened, with a large light basal scar
Type species: Mammillaria conoidea De Candolle.
The genus is dedicated to Professor Francis E. Lloyd.
1.’Neolloydia conoidea (De Candolle) comb. nov.
Mammillaria conoidea De Candolle, Mém. Mus. Hist. Nat.
Paris: 17: 112. 1828.
Echinocactus conoideus Poselger, Allg. Gartenz. 21: 107. 1853.
2.‘ Neolloydia Beguinii (Weber) comb. nov.
Echinocactus Beguinit Weber in Schumann, Gesamtb,
Kakteen 442. 18098.
INDEX TO AMERICAN BOTANICAL LITERATURE
1908-1921
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
America being used in the broadest sense.
Reviews, and papers that relate exclusively to foresty, agriculture, horti-
culture, manufactured products of vegetable origin, or laboratory methods
t included, and tempt i de to index the literat f bacteriology.
An occasional exception is made in favor of some paper appearing in an Ameri-
can 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.
s Index is reprinted monthly on cards, and furnished in this form to
subscribers at the rate of three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Andrews, F. M. The effect of aeration on plants. Proc. Indiana
Acad. Sci, 1920:.147, 148. 1921.
Andrews, F. M. The effect of centrifugal force on plants.
Proc. Indiana Acad. Sci. 1920: 143-145. 1921.
Andrews, F. M. Phyllotaxis of mee iovevat Proc.
Indiana Acad. Sci. 1920: 149, 1 1921
Andrews, F. M. Studies on es III. Proc. Indiana Acad.
Sci. 1920: 155-156. 1921.
Baerthlein, K. Uber bakterielle Variabilitat, insbesondere
sogenannte Bakterienmutationen. Centralbl. Bakt. Par-
asitenk. u. Infekt. 81'!: 369-435. 31 JI 1918.
Baker, F. C. The life of the Pleistocene or Glacial Period.
Univ. Illinois Bull. 17: 1-476. pl. 1-56. 7 Je 1920.
Bardell, E. M. Production of root hairs in water. Univ. Wash-
ington Publ. Bot. 1: 1-9. 1 F 1915.
Barrus, M. F. Bean anthracnose. Cornell Univ. Agr. Exp.
Sta. Mem. 42: 101-214. pl. 1-8+f. 10-19. Jl 1921.
253
254 INDEX TO AMERICAN BOTANICAL LITERATURE
Barton, J. E. The amount of standing timber in Kentucky.
Min. and For. Resources Kentucky V. 1: 251-284.
Beccari, O., & Rock, J. F. A monographic study of the genus
Pritchardia. Mem. B. P. Bishop Mus. 8: 1-77. pl. I-24.
1921.
Includes 13 new species from the Hawaiian Islands.
Belling, J. The behavior of homologous chromosomes in a
tpl canna. Proc. Nat. Acad. Sci. 7: 197-201. f. 1. 2.
Jl.
ae H. M. Further consideration of the sizes of vein
islets of leaves as an age determinant. Science II. 55: 399,
400. 14 Ap 1922.
Benedict, R. C. Is Botrychium dissectum a sterile mutant of
B. obliquum? Am. Fern. Jour. 11: 53-55. 1921.
Benedict, R. C. Nephrolepis nutrition. Am. Fern Jour. 11:
41-43. 1921. [Illust.]
Berry, E. W. Tertiary fossil plants from Venezuela. Proc.
U. S. Nat. Mus. 59: 553-579.. pl. 107-109. 1921.
Bowman, H. H. M. Histological variations in Rhizophora
Mangle. Michigan Acad. Sci. Rep. 22: 129-134. pl. 9-12.
Au 192I.
Britton, N. L. George Valentine Nash. aa New York Bot.
Gard. 22: 145-147. pl. 261. Au 1921
Brown, F.-B. H. Origin of the Hawaiian flora. Proc. Pan-
Pacific Sci. Confer. 1: 131-142. 1921
Brown, L. L. Canadian Sitka spruce, its mechanical and phy-
sical properties. Canada Dept. Inter. For. Bull. 71: 1-39.
f. I-20. ‘1921.
Brunswik, H. Uber Hesperidinspharite im lebenden Hautgewebe
von Antherium Binotii Lenden. Ber. Deutsch. Bot. Ges.
39: 208-212. Je 1921.
Buscalioni, L., & Muscatello, G. Studio monografico sulle
specie americane del gen. ‘“‘Saurauia’’ Willd. Malpighia
: I-32; 97-112. I92I.
Includes Saurauia intermedia sp. nov. from Guatemala.
INDEX TO AMERICAN BOTANICAL LITERATURE 255
Buscalioni, L. II legno crittogamico del fascio vascolare seminale
di talune angiosperme considerato nei suoi rapporti colle
moderne teorie filogenetiche. Malpighia 29: 46-80; 113-207.
f. 1-30: 3
Butters, F. K. A new western species of Pellaea. Am. Fern
Jour. 11: 39, 40. 1921.
Pellaea Suksdorfiana from Washington.
Butters, F. K. Salvinia in Minnesota. Am. Fern Jour. 11:
45-50. i921.
Chapman, G. H. Tobacco investigations. Massachusetts Agr.
Coll. Bull. 195: 1-38. pl.z-zB. [1920]
Chiovenda, E. La culla del cocco. Webbia 5: 199-294. 1921.
Concerning the origin of the coco-palm.
Christensen, C. An overlooked species of Dryopteris Am.
Fern Jour. 11: 44-46.
Dryopteris crypia (Underw. & Maxon) from Cuba.
Conzatti, C. Monografia del Arbol de Santa Maria del Tule.
Pp. 1-65. Oaxaca, Mexico. Mr 1921. [Illust.]
Account of a giant tree of Taxodium mucronatum Ten,
Coons, G. H., & Levin, E. The relation of light to pycnidium
formation in the Sphaeropsidales. Michigan Acad. Sci.
Rep. 22: 209-213. Au 1921.
Corrington, L. J. The Peltigeraceae. [The Ascomycetes of
Ohio V]. Ohio Biol. Surv. 2: 354-362. pl. 14, 15. 28 My
1921,
Coville, F. V. The influence of cold in stimulating the growth
of plants. Ann. Rep. Smithsonian Inst. 1919: 281-291.
pl. I-27. 1921.
Dahlgreen, U. Phosphorescent animals and plants. Nat. Hist.
22: 1-26. Ap 1922.
Dallimore, W. The yellow pines of North America. Kew Bull.
Misc. Inform. 1921: 330-335. 1921.
Dalton, L. V. The plants and animals of Venezuela. In Dalton
L. V., Venezuela, 47-60. London. Ig912. [Illust.]
256 INDEX TO AMERICAN BOTANICAL LITERATURE
Darlington, H. T. Contributions to the flora of Gogebic County,
Michigan. Part I. Michigan Acad. Sci. Rep. 22: 147-176.
Au 1921.
Deam, C. C. Plants new to Indiana—IX. Proc. Indiana
Acad. Sci. 1920: 225-228. 1921.
Dow, C. M. Anthology and bibliography of Niagara Falls.
Pp. 1-1423. Albany. 1921. [Illust.]
peracki! VI (pages 439-491) gives a chronologic discussion of the “flora
and fauna
lacy: J. Anaerobic bacteria in plant tissues. Phyto-
pathology 11: 344. Au 1921.
Dupret, H. Notes about the Drepanocladi of the vicinity of
Montreal, Quebec, Canada. Bryologist 24: 36-39. 21 5
1921.
Earle, F. S. S. M. Tracy as a botanist. Torreya 21: 64, 65.
Au 1921.
East, E. M. A study of partial sterility in certain hybrids.
Genetics 6: 311-365. f. 1-17. Jl 1921.
Studied crosses of Nicotiana.
Emerson, R. A. Heritable characters of maize—IX. Crinkly
leaf. Jour. Hered. 12: 267-270. f. 24-26. 23 S 1921.
Enock, C. R. Natural History [of Peru]. In Enock, C. R.,
Peru 192-205. London. 1908
Espinosa Bustos, M.R. Una enfermedad grave del trigo Chileno.
Revista Chilena Hist. Nat 24: 79-86. pl. 34+/.8: 31 Au 1920.
Essary, S. H. Lespedeza err clover). Univ. Tennessee
Agr. Exp. Sta. Bull. 123: 1-28. Ap 1921. [lIllust.]
Evermann, B. W., & Clark, H.W. The flora of Lake Maxinkuckee
and vicinity. In Evermann & Clark’s Lake Maxinkuckee
[Indiana], a physicial and biological survey 2: 117-447. 1920.
Farwell, O. A. Notes on the Michigan flora—III. Michigan
Acad, Sci. Rep. 22: 177-185. Au 1921.
Includes new varieties and forms in various genera.
Fink, B. The Lecideaceae. [The Ascomycetes of Ohio IV.]
Ohio Biol. Surv. 2: 334-353. pl. 13. 28 My 1921.
Includes new species in Lecidea (1), Bacidia (1), Buellia (1), Rhizocarpon (1).
INDEX TO AMERICAN BOTANICAL LITERATURE 257
Fink, B. - Lincoln Ware Riddle, lichenist. Bryologist 24:
33-36. 21'S 1921.
Fisher, G.C. A station for the ramshead lady’s-slipper. Torreya
21: 63, 64. Au 1921.
Forbes, C. N. New Hawaiian plants—VII. Occ. Pap. B. P.
Bishop Mus. 7: 33-39. pl. 3-1r. 1920.
Includes 7 new species.
Forbes, C. N. Notes on Marsilea villosa Kaulf. Occ. Pap. B. P.
Bishop Mus. 7: 47-49. pl. 13, 14. 1920.
Forbes, C. N. Salient features of Hawaiian botany. Proc.
Pan—Pacific Sci. Confer. 1: 125-130. I921.
Forbes, C. N., & Munro, G. C. A new Cyanea from Lanai,
Hawaii. Occ. Pap. B. P. Bishop Mus. 7: 43. pl. 12. 1920.
Cyanea Baldwinit, sp. nov.
Gardner, M. W. Indiana plant diseases, 1920. Proc. Indiana
Acad. Sci. 1920: 187-208. f. 1-12
Gassner, G. Ueber einen eigenartigen Uromyces auf Passiflora
foetida L. Ber. Deutsch. Bot. Ges. 40: 64-68. f. I-3. 30
Mr 1922.
Includes Uromyces Appelianus sp. nov. from Uruguay.
Girola, C. D. Cultivo de la yerba mate (Ilex paraguariensis St.
Hil.) Cap. IV. Bol. Minist. Agr. [Argentina] 26: 128-142.
Je 1921.
Grabou, A. W. A textbook of geology, Part II. Historical
Geology. Pp. 1-976. f. 735-1980. New York. 1921.
Contains much false botanical matter.
Grinnell, J., & Storer, T. I. Life-zones of the Yosemite Region.
In Hall, A. F., Handbook of Yosemite National Park [Cal-
ifornia], 123-132. New York. 1921.
Hall, A. F. Trees of Yosemite National Park [California].
In Hall, A. F., Handbook of Yosemite National Park
[California], 219-234. pl. 18-21. New York. 1921.
Harshberger, J. W. Geographic names and terms of significance
in plant geography and rtd Bull. Geogr. Soc. Phila-
dephia 19: 13-18. O 19
258 INDEX TO AMERIC AN: BOTANICAL LITERATURE
Harshberger, J. W. The old gardens of Pennsylvania. Gard.
920): 78-80; 137-139; 257-258; 326-328; 44-46;
120-123; 195-196; 255-256; 326-329; 374-377. 1921.
[Illust.]
Heimlich, L. F. Native plants of White County [Indiana]—III.
Proc. Indiana Acad. Sci. 1920: 219-224. 1921.
Hitchcock, A. S. Foral aspects of British Guiana. Ann. Rep.
Smithsonian Inst. 1919: 293-305. pl. 1-1z2+/f. I-3. 1921.
Hoffmann, R. Flora of Berkshire County, Massachusetts.
Proc. Boston Soc. Nat. Hist. 36: 171-382. Mr 1922.
Hollister, B. A. The relation between the common weeds of
Michigan and those found in commercial seed. Michigan
Acad. Sci. Rep 22: 187, 188. Au 1921.
Holm, T. Studies in the Cyperaceae, XXXII. Carices aeroast-
achyae: Phacotae nob. and Ternariae nob. Am. Jour. Sci.
202? 322-329. ff. F-#. DD 1621.
Hopkins, E F. The Botrytis blight of tulips. Cornell Univ.
Agr. Exp Sta. Mem. 45: 315-361. pl. 33+f. 22-50. Au 1g2!.
Hotson, J.W:. Sphizgnum used asa surgical aressing in Germany
during the world war. Bryologist 24: 74-78, 89-96. f. 1-8.
Py. Fo. 1022.
House, H. D. Nomenclatorial notes on certain American
~ plants—I. Am, Midl. Nat. 7: 126-135. S 1921.
Includes Trianthella gen. nov. and new combinations in many genera.
Howell, A. H. A biological survey of Alabama. N. Am. Fauna
45: 1-88. pl. 1-11+ f. I-10. 1921. ~
Illick, U. S. Big trees in Pennsylvania. For. Leaves 18: 24,
25. Ap 1921. [Illust.]
Jackson, H. S. The Uredinales of Indiana III. Proc. Indiana
Acad. Sci. 1920: 164-182. f. 7. 1921.
jackson, H. S. The Ustilaginales of Indiana II. Proc. Indiana
Acad. Sci. 1920: 157-163. 1921.
Jepson, W. L. Flowers of Yosemite National Park. In Hall,
A. F., Handbook 2f Yosemite National Park [California],
247-267, pl 24-27 New York 1921.
Vol. 49 | No. 9
BULLETIN
OF THE
TORREY BOTANICAL CLUB
SEPTEMBER, 1922
New species of trees of medical interest from Bolivia
H. H. Russy
The botaincal work of the Mulford Biological Exploration of
1921-22 was directed specially toward the investigation of
medicinal plants. The distinctly pharmacological features of
these plants will be discussed in the medical and pharmaceutical
literature, but such unknown species as are encountered should
be cited in botanical publications. Two problems of exceptional
interest were connected with the botanical origin of the two
drugs, “‘coto” and ‘‘cocillana” or “‘guapi,” and their spurious
substitutes.
It has never been known what trees yield the coto and paracoto
barks, although evident that they pertain to the Lauraceae.
For many years, no genuine bark of either has reached our
markets, and the use of their worthless substitutes has resulted
in destroying the medical reputation of these valuable drugs.
Through the careful work of my associate, Dr. O. E. White, of
the Brooklyn Botanic Garden, it is now possible to describe the
genuine coto tree, as well as its principal substitute, and to provide
for the supply of the genuine drug in future. The bark of a
second substitute, with leaf-bearing branches, was also collected,
but the species cannot be determine d from this material, though
undoubtedly pertaining to the genus Nectandra.
Of cocillana or guapi bark, I collected not only the genuine’
species, but also the only substitute that has as yet been seen in
commerce, besides two other species which, for special reasons,
are liable to so appear in future.
The BuLLetin for August (49: 223-258) was published August 31, 1922.
259
260 NEW SPECIES OF TREES FROM BOLIVIA
Those interested further in these subjects should consult
future numbers of the Journal of the American Pharmaceutical
Association.
Nectandra coto Rusby, sp. nov.
Young parts, inflorescence and lower leaf-surfaces densely
and finely yellowish-tomentellate, the twigs stout. Mature
leaves, including the petiole, 10-20 cm. long, 3~7 cm. wide, the
petiole 2-3 cm. long, stout, sharply channelled above when dry,
reddish brown; blade thi ck and coriaceous, 23 Sess er
prominent, the secondaries twelve to fourteen on each side,
the margin, the 'interspaces faintly reticulate. Inflorescence
. : : ze
it when young. Fruiting pedicel of variable length,
sicketed upward and becoming continuous with the calyx.
roughish, single but showing two distinct layers, the light brown
fruit slightly projecting when mature, its summit lightly convex,
finely granular or lightly mottled, bearing the very short, stout,
acute style.
Mulford Biological Sapa Nos. 1048 and 1050. Collect-
ed by Dr. O. E. White as ‘‘Coto No. 1” (No. 1048) and “Coto
No. 1a” (No. 1050), and a the native names “coto piquiante”’
and ‘‘coto fino.”” Mountains south of Huachi, near the Cocha-
bamba River, Bolivia, at an altitude of 2500 to 3000 feet,
_September 9, 1921
Dr. White’s field notes are as follows:—
No. 1048.—A tree 50 to 60 feet high, with few branches except near the
sandal-wood yellow and a satiny luster, the trunk turning brown after being
peeled. Leaves whitish beneath, dark- to light-green and
Young growth cinnamon-brown. Flower buds whitish. Cup of fruit brown
NEW SPECIES OF TREES FROM BOLIVIA 261
green with brown spots, the fruit green, evidently of two generations on the
same tree
It is not possible to say with absolute certainty, from speci-
mens collected in this stage of growth, whether this is a species
of Nectandra or of Ocotea. The slender midribs and veins of the
leaves are more those of Ocotea than of Nectandra, but the tomen-
tose flower-buds are characteristic of Nectandra, the flowers of
Ocotea being usually glabrous. There is no species of either
Nectandra or Ocotea with which this can be identified.
Ocotea pseudo-coto Rusby, sp. nov.
oung growth and inflorescence whitish-puberulent, the
lower surfaces of the mature leaves minutely and _ sparsely
ag lal the branchlets elongated and stout, little spreading,
deep
red, more or less angled. Leaves, including the —
at cm. roe 1.5-3 cm. broad, the petioles I-1.5 cm. long,
deep red, the upper surface moi plane. Blades diel oblong,
acuminate at base and wit a very abrupt short and stout
obtuse point at the summit, the margin entire, not series the
upper surface brown when dry, glabrous, slightly lustrous, the
midrib slightly prominent, sharply so underneath, the six to
near the margin. Panicles axillary, less than half the length o
their leaves, sessile or etoaccile. broad, rather dense as seen 3
the young state. Bracts thickish, red, broadly ovate. Perigo
thin, glabrous, ean oped in the specimen. The other deel
parts are too young for characterization
Collected by s O. E. White in the mountains south of
Huachi, near the Cochabamba River, Bolivia, at an altitude of
2500 to 300 feet, September 9, 1921. Mulford Biological
Exploration No. 1051, collected as ‘‘Coto No. 2” and under the
name “‘coto ordinario.”’ It is to be noted that the term “ordin-
ario,’’ as applied by these natives to any substance or article,
has the significance that the article is of inferior quality.
Dr. White contributes the following field notes:—
large forest tree, growing with Nos. 1048 and 1050 and of the same
general habit. Young buds only, and no fruit encountered. Bark thick and
easily peeled, not specially aromatic and not pungent. The peeled wood
remains white. Wood reddish-white internally.
The same difficulty of identification presents itself with this
species as with No. 1050. The buds are a little more developed
and the perigone characters are found to correspond with those
of Ocotea.
262 NEW SPECIES OF TREES FROM BOLIVIA
The bark of this species is one of those that has very commonly
occurred in commerce in this country under both names, coto
and paracoto, but sold under either name it is absolutely
spurious and medicinally worthless, so far as known.
Aerodiclidium benense Rusby, sp. nov.
Specimen in mature fruiting stage. Glabrous, ae twigs
numerous, rather slender, terete, densely leafy at the summit.
Leaves, including the petiole, 10-15 cm. long, 3-5 cm ts oad.
Petioles 5-10 mm. long, slender, sharply channelled above.
Blades thick and coriaceous, lustrous above, lanceolate, abruptly
th
the margin, the reticulation fine and sharp. Margin entire, very
slightly revolute. Panicles ian. less fae half the length of
their leaves, stoutly peduncled, having one to jour shortly and
stoutly pedicelled fruits, the pedicels strongly thickened upward.
Calyx-cup subhemispherical, about 1.5 cm. broad, hard a nd thick,
projecting paid pevond the strongly recurved limb so as to
mouth, the limb thin, irregularly crenate-dentate.
Fruit about Z cm. long cad two-thirds as broad, oval to ovoid,
light brown, glabrous, strongly wrinkled at the base and very
shortly stipitate, the rounded summit topped by the very short
stout style
Collected by Dr. O. E. White at the second portage of the
Bopi River, Bolivia, at an altitude of about 2000 feet, August
8, 1921. Mulford Biological Exploration No. 649.
Dr. White says: ‘‘A small evergreen tree 20 to 30 feet high,
trunk 6 inches in diameter, growing in damp, partly shady
places near the river bank. The fruit is green with white specks,
the cup brilliant scarlet-red. Not common.”
Guarea Bangii Rusby, sp. nov.
Only fruiting eS seen. Glabrous, with the exception
NEW SPECIES OF TREES FROM BOLIVIA 263
abruptly contracted into a short and ein mostly obtuse
lu Midrib
sessile, rather densely fruited, the rachis and pedicels very stout
the latter little longer than broad, annular-roughened, articulated
with the very short stipe of the fruit, which is not so thick as its
pedicel. Mature fruit usually a little more than 2 cm. long and
nearly or quite as broad, pear-shaped when young, becoming
nearly ee —— for the narrow base, which remains
unchanged to a short thecaphore. Body of fruit obscurely
costate or ela riberealae teeta and soitly and finely
tomentose.
First collected by Mr. Miguel Bang, on the hills near Tipuani.
Collected also by H. H. Rusby in October, 1921, on the Mulford
Biological Exploration, at Rurrenabaque, Bolivia, Nos. 779 and
1590; also No. 2178 (‘‘Guapi B’’). All the spurious cocillana
bark that has reached our market up to the present has been
derived from this species.
A good-sized forest tree of the Mapiri and Beni River region,
growing mostly on the hillsides in relatively dry soil, the bark
roughly striate and shreddy, and peeling in long, fibrous, thin
strips.
~
Guarea alborosea Rusby, sp. nov.
Glabr ge te ees and lower surface of midrib finely
verrucose pillose in the dried state. ok and rachis
rather stout: doas ur ele: together 2-4 dm. long, the petiole
about one-fifth of the total length, flat on the bn surface, the
leaflets five to ten pairs, nearly opposite, the lower somewhat
smaller. Petiolules 4-8 mm. long, stout, lightly channeled above,
like the midrib. Blades io-38 cm. long, by 4-8 cm. wide, thick-
ish, pale underneath, oval or slightly narrower either below or
side, not opposite, straight and slender, moderately spreading,
falcate wens the margin, connected by the coarsely reticulate
ertiari Inflorescence axillary, racemose, simple, or occasion-
ally slightly branched = the base, the racemes recurved-pendu-
lous, often 2-3 dm. long, 1.5-2 cm. ee densely flowered, the
rachis slender, loaaty 1 nodose by th e flower-scar s, the bractlets
minute, the flowers divaricate. Pedicels lender articulated
264 NEW SPECIES OF TREES FROM BOLIviA
into the broad concave base, bse pedicel proper scarcely one-
fifth - = of the flower, obscurely angled. Unopened
calyx n Mature dawers about 6 mm. long, the erect
cai rg coils 2.5-3 mm. broad. Calyx crateriform, 3
mm. broad, closely investing the base of the corolla, deeply
ong ‘the ae oe purple, broadly ovate and obtuse.
Petals 6 mm. long m. broad, oblong, obtuse, slightly thick-
ened at Go pucdco or rose colored summit, the basal portion white.
Stamen-tube three-fourths the smi ‘of the petals, broadly
b h h
Oo
uite so broad as the lower half, the margin shallowly eight-
appressed-pilose, the ovoid ovary about equalling the stout
Immature fruit fig-shaped, hee contracted at the base, then
tapering into the stout short pedicel. Mature fruit not seen.
A small tree in the forest a Rurrenabaque, Bolivia, at an
altitude of about 1000 feet. Collected by H. H. Rusby in
flower and fruit, on the Mulford Biological Exploration, October
6, 1921 (No. 797).
e species is peculiar in its very long slender raceme and in
the handsome rose and white coloration of its flowers.
Notes on trees and shrubs of the southeastern United States
W. W. ASHE
1. THE CASTANEA PUMILA GROUP
Castanea pumila Margaretta var. nov.
A tree, sometimes attaining (in Arkansas) a height of 15 m.
and a diameter of 1 m., but usually smaller and often a large
single-stemmed shrub. Leaves sinuate-serrate with ten to
on the midrib and veins; shade leaves 5-12 cm , o-6-8
oad, obovate or sometimes oblong-obovate, obtuse, or rounded
and abruptly pointed at the apex, cuneate at t , thin,
aaa
with branched rigid canescent spines, 3-6 mm. long, which as a
rule are not sufficiently dense to conceal completely the involucre.
—-I15 mm. long, 7-12 mm. thick, pointed
at the tip, shining, blackish brown.
Dry soils or, at the upper edge of the longleaf pine lands of
Texas and Louisiana, along the edges of sandy hummocks.
The following collections have been made by the writer: ALA-
BAMA, Winston County; Mussissrppi, Jackson; LoursIANA,
Natchitoches and Sabine (Tyre) Parishes; TExAs, Sabine County;
OKLAHOMA, McCurtain County; and ARKANSAS, common on the
mountains. Named for Margaret Henry Newland.
This variety largely replaces the typical C. pumila-(L.) Mill.
west and southwest of Tennessee and is separated from it by hav-
ing the much shorter and smaller shade leaves of an obovate
outline, thinner and often green beneath ; by the closer white pube-
scence on the narrow and acute sun leaves (never velvety and
rarely dull grayish); by the spines of the involucre being
canescent well toward their tips (those of the typical C. pumila
being nearly glabrous when mature) and seldom so dense as to
conceal the involucre. Nut apparently not quite so plump as
that of the type.
265
266 NOTES ON TREES AND SHRUBS
CASTANEA ALNIFOLIA Nutt.
The western recorded limit of this dwarf shrub with its sub-
terraneous stems is Louisiana.* It was collected by the writer
in Newton County, Texas, during October, 1921, under the
same conditions as it grows further east and not exceeding 18
inches in height. C. alnifolia pubescens Nutt. has all of the leaves
pubescent beneath.
v Castanea floridana (Sarg.) comb nov.
Castanea alnifolia floridana Sarg. Bot. Gaz. 67: 242. 1919.
I have not seen authentic specimens of the tree which Dr.
Sargent has recently described and which he separates from the
typical C. alnifolia by its arborescent form, but it is believed
that material collected near Charleston, South Carolina, in 1915
represents it; and, if so, the differences of smaller fruit (which
is more densely hispid), somewhat larger leaves with about two
more pairs of lateral veins, and entirely different habit of growth
justify specific rank.
x CASTANEA NEGLECTA Dode, Bull. Soc. Dendr. France 8: 155.
May, 1908. C. pumila x C. dentata
According to the description this hybrid is in general like C.
pumila. It differs, however, in having larger and less densely
pubescent leaves, a larger involucre with irregular spines, longer
aments, and a single larger nut. The distribution is given as
extending from Maryland to North Carolina. As a rule hybrids
between C. pumila and C. dentata have the nuts in pairs, although
this is not necessarily the case.
M. Dode has been kind enough to furnish the writer with a
list of the specimens at the Museum of Paris, which he refers to C.
neglecta. Of the seven specimens listed, the locations of five are
well within the ranges of both C. pumila and C. dentata. These
are: Small & Heller 152, and their collection of June 22, 1891,
both from Blowing Rock, North Carolina; Pearson, Fairfax
County, Virginia, 1844; Green, Maryland, 1839; Bonpland,
Washington, D. C. Since the states only are given for the other
two (Gibbs, South Carolina, and Vasey, Alabama, 1878), it is
not possible to determine this.
* Small, Flora S. E. U. S. Ed. 2, 348.
NOTES ON TREES AND SHRUBS 267
Material collected by the writer in the mountains of Rabun
County, Georgia, and Yancey County, North Carolina, largely
agrees with Dode’s description of C. neglecta. In its gross
characters it shows, as Dode states, a superficial resemblance
to C. dentata, having large pointed leaves, not very densely
pubescent, and a large involucre with long, irregular, nearly
glabrous and closely crowded spines; but there are two nuts to
the involucre. Other material collected in flower in Sullivan
County, Tennessee, shows similar leaf character.
’ Castanea Ashei Sudworth, comb. nov.
Castanea pumila Ashei Sudworth, Am. For. 28: 300. 1922.
This is the common chinquepin of the coastal plain of the
Southern States. Its distribution begins in northeastern Nort
Carolina and extends to Louisiana (and probably to south-
eastern Texas). It is separated from C. pumila by its smaller
foliage, shorter and more slender aments, canescent spines on
the involucre and prevailingly thicker nuts. Walter (Fl. Car.
233. 1788) notes two forms (without description), praecox and
serotina. C. Ashet is possibly his forma praecox.
The type of C. pumila (L.) Mill. (Fagus pumila L.) came from
eastern Virginia (Gronovius, Flora Virginica 118. 1739), within
which region the ene known form is that which is here regarded
as typical C. pumi
Key to Castanea pumila group (one nut to the involucre)
Al=Alleghanian; C=Carolinian; A=Austral.
Spines of mature fruit densely set, glabrous or
glabrate.
Leaves soft-pubescent beneath............. C. pumila (Al & C).
Leaves 10-20 cm. long, more or less glabrate.. x C. neglecta (Al).
Spines of mature fruit not densely set, canescent or
Leaves 10-15 cm. long; 12-20 pairs of veins.
All leaves closely white-pubescent be-
RABE a ee ee C. Ashet (A).
Interior leaves green beneath or glab-
MitG ee ee C. pumila Margaretta (C).
Leaves 6-10 cm. long; 8-16 pairs of veins.
Only uppermost leaves on twigs white-
pubescent beneath.
10-16 pairs of veins, non-stoloniferous. C. i ilecosd (A).
8-12 pairs of veins, a dwarf stoloniferou
OMIM re ae C. alnifolia (A).
All leaves white pubescent beneath...... C. alnifolia pubescens (A),
268 NOTES ON TREES AND SHRUBS
2. MISCELLANEOUS SPECIES
v
x Quercus coloradensis sig nov. Q. virginiana Mill. x Q.
macrocarpa Mx.
Leaves aero eagteae fe and oblong or undulate, sinuate
or lobed, especially near the apex, with shit abruptly acute
or obtuse lobes, 4. es cm. long, 2-4 cm. thic
coriaceous, dark gr een above, strongly pedeulated aid deheely
close white- canines beneath, revolute on the margins; twigs
3-4 mm. thick, permanently close pale gray pubescent. Fruit
mm. long, excluding pe eduncle (which i is often of half the
length of the fruit); cup covering one-half or more of the oblong
nut, obconic or slightly rounded, covered with slender acute,
closely imbricate scales, either thin at the top or much thickened
by several overlapping rows of scales and incurved at the edge.
Along the Colorado River above Marble Falls, Texas. While
the overcup oak is rare in this region, the cup-character points
to that species rather than to Q. acuminata Houba or Q. stellata
Wang, as a possible parent.
Quercus nigra elongata comb. nov.
Quercus aquatica elongata Ait. Hort. Kew. ed. 1, 3: 357. 1789.
In the third edition of Aiton’s work there is a reference to a
plate (Abbott, Insects Ga. 2: pl. 29.), which figures the leaf-
form described bis Sargent under his Q. nigra tridentifera (Bot.
Gaz. 65: 429. 1918).
v Malus elongata pubens var. nov.
In general similar to the type but having somewhat broader
leaves, pernianently soft-pubescent beneat
ead of Chattoga River, Rabun County, Georgia, October,
ors; W. WwW. A.
“Malus platycarpa parvula var. nov.
Foliage in general similar to that of type, but the leaves more
ovate in outline and smaller; fruit of the same shape but much
smaller, no larger than that of M. elongata.
Habersham County, Georgia, October, 1913, W. W. A.
Commelinantia, a new genus of the Commelinaceae
B. C. THARP
(WITH PLATES IO AND IT)
Few if any families of plants have members in more urgent
need of field study than are those of the family Commelinaceae.
Their evanescent corollas, which it is difficult even to bring
from field to laboratory without injury; the delicate shades of
color which are lost in pressing; and their succulent stems,
which shrink tremendously in drying, make field study imperative
if one is to get any adequate idea of these delicate details as they
appear in the growing plants. Since 1919 I have had abundant
opportunity for field study of a very interesting member of the
family, a species first described by Torrey* as Tradescantia
anomala and latter transferred by C. B. Clarket to the genus
Tinantia, where it has remained to the present time.
Torrey’s description, based upon specimens from Texas, (one
of Wright’s, and one of Thurber’s), preserved in the herbarium
of the New York Botanical Garden, is surprisingly good, con-
sidering that it was drawn wholly from dried material. Suc
details of general habit, sepals, petals (except color), filaments
(except that of the posterior stamen), anthers, style, stigma,
and ovules, as are pointed out, are correctly described. The
greatest error into which his material led him seems to have
been in considering the ovary and capsule two-celled. He also
described the seeds as being ‘‘as in the rest of the genus,’’ which
is incorrect. The wonder is that, having examined his material
so carefully as to be able to write such an accurate description,
he could feel justified in assigning this species to the genus
Tradescantia. That he was none too confident of its having been
correctly assigned is abundantly testified by the specific name,
by legendst on the sheets containing his specimens, and by his
*U. S. and Mex. Bound. i 225. geo
{t DeCandolle, Monog. Phaner. 3: 28
t Wright ’s specimen is labelled * Wright, pee Commelyna anomala, Torr.
Tradescantia anomala Torr.,” the word “Commelyna’’ being stricken out.
Thurber’s specimen is labelled “Thurber, San Antonio Texas, May 1853.
Commelyna rhodantha, Torr
269
270 A NEW GENUS OF THE COMMELINACEAE
observation that ‘“‘the species is intermediate between Trades-
cantia and Commelyna.”’
Clarke’s description of Tinantia anomala appears to be based
neither upon the excellent original by Torrey, nor upon any
critical examination of material of the species. He characterizes
the plant as glabrous, the leaves as ovate-cordate, the three
longer stamens simply as ‘‘naked above,” the three shorter as
having ‘‘golden bearded anthers,’’ the stems as “‘ dichotomously
branched,” the inflorescence as being ‘‘almost included in the
subcomplicate uppermost leaf,’”’ the petals as being ‘‘obovate-
elliptic, bluish”’ (by inference all alike in size and appearance).
ith the exception of the capsule, the measurements of all
structures are under-stated. His specimens may not have shown
the basal linear-lanceolate, ciliate, leaves; his scant measure-
ments may have been due to his materials having shrunken in
drying, and his representation of the “uppermost leaf’’ as
“‘subcomplicate”” may have been due to his specimens having
been somewhat wilted before being pressed; but there seems no
valid excuse for so inaccurate a description of stamens, or for
characterizing the stems as dichotomously branched, or for
totally ignoring the description (excellent so far as it goes) of
Torrey. His observation, concerning the relationship existing
between the type species (7. -fugax) and the other species, that
“caeterae species . . cum J. jupace ... . ... militant;
et ad Tinantiam ex hetomitate referendae sunt, nisi genus Jinantia
ad Tradescantiam reducatur,’’ seems to be in the nature of an
apologetic attempt at justification, though it is difficult to see
the logic in his statement.
My own experience with an abundance of fresh material of
the species in question constrains me to call attention to the
following characteristics, which keep it from being properly
placed in the genus Tinantia as characterized by Clarke: branches
which break through the leaf sheaths; flowers in a simple ter-
minal scorpioid raceme subtended by a single, broad, erect, flat,
cordate, clasping spathe; colored lateral petals much larger than
the very small anterior white petal; stamens of four distinct
kinds as regards size and pubescence.
The character of the stamens and of the spathe and the habit
of the branches breaking through the leaf sheaths are sufficient
to exclude it from either Commelina or Tradescantia. The
former has difformed stamens, all of whose filaments are entirely
A NEW GENUS OF THE COMMELINACEAE ° 271
glabrous and some of whose anthers are sterile; a spathe which is
folded so as to protect alike the flowers bud and the ripening
capsule, the flower being projected above the edges of the spathe
only during the very brief period of bloom: the latter has stamens
all alike in size and pubescence and all fertile; the inflorescence
subtended by one or more (usually two) unlike bracts, never by
a spathe. Neither Commelina nor Tradescantia has branches
which break through the leaf sheaths. This habit, however, is
held in common with certain tropical genera to which the species
has never been heretofore compared. Dr. Pennell has kindly
gone over the whole family as represented by specimens in the
New York Botanical Garden Herbarium and reports the following
genera as having this habit:
1. Callisia (tropical American) lacks spathe, has uniform petals and
stamens,
2. Campelia (tropical American) has spathe, but uniform petals and
stamens, white.
3. Cyanotis (tropical Asia) has series of overlapping spathes.
4. Dichorisandra (tropical American) lacks spathe, inflorescence a panicle.
5. Floscopa (tropical American and Asia) lacks spathe, inflorescence a
panicle.
6. Forrestia (tropical Asia) lacks spathe, inflorescence congested erupts
spathe.
7. Pollia (tropical Asia) lacks spathe, inflorescence a panicle.
8. Sauvellea (Cuba) has spathe, but petals and stamens uniform; repent.
9. Streptolirion “age has spathe below cymes or panicle, but petals
ei stamens uniform, the filaments bearded. Stem twinin
Pradesconies (Florida) lacks spathe, but petals and cannens uniform.
Racor
tT: Zebrinn (tropical American) has two bracts below the congested in-
florescence; petals uniform, at base approximating to form a corolla-tube.
Repent.
The species differs from any of these generic groups in its
combination of erect habit, single, flat, erect, spathe; unequal
petals; and stamens of four forms.
In addition to the several genera listed above, Dr. Pennell
finds that Tradescantia Pringlei S. Wats.* also has leaves
which break through the sheathes. Moreover, it has the following
additional characters which ally it very closely to T. anomala:
general appearance very similar; inflorescence a simple scorpoid
raceme (or frequently reduced to only one flower); stamens
probably like those of 7. anomala in that there are four kinds,
*Trans. Am. Acad. 26: 157. 1891.
272 . A NEW GENUS OF THE COMMELINACEAE
implied by the following statement: “stamens 6, unequal, the
longer with filiform filaments bearded in the middle (or one
naked) and broadly oblong anthers, the shorter filaments densely
bearded in the middle with green gland-tipped hairs, the anthers
or bicular; anther cells contiguous.’’ The stamen corresponding
to the posterior stamen in 7. anomala might, by casual examina-
tion, have been counted as being like the postero-lateral ones.
But 7. Pringlei has the following characters in conflict with
T. anomala: plant green, not glaucous; cauline leaves uni-
formly ovate, abruptly acuminate, none modified to form a
definite “‘spathe’’ (though the slender peduncles of T. Pringlei
frequently have one or more much reduced sheathes below the
inflorescence), inflorescence axillary, petals uniform, all blue
(“‘purple” ?). Though 7. Pringlei thus shows such strong
resemblance to 7. anomala in certain characteristics, yet
because of these important points of dissimilarity and because
of insufficient material for critical study I feel constrained to be
content with pointing to it as perhaps the closest existing relative
of 7. anomala, leaving the question of its assignment to the
proper genus to be settled by a more critical study of more
abundant material. Certainly it can not properly belong in
the genus Tradescantia!
Confining my attention, therefore, in the present paper to
the placing of T. anomala, it seems not only justifiable but
necessary to make it the type of a new genus.* Following the
suggestion of Torrey that the species has certain characteristics
which ally it to Tradescantia and Commelina and recognizing
the fact that it was assigned to Tinantia more than forty years
ago, I propose a sort of hybridization of these names, giving the
new genus the name Commelinantia. The generic description
which follows is made broad enough to include some of the dis-
tinctive characters of T. Pringlei, acting on the probability that
this species will ultimately be assigned to the new genus.
COMMELINAN TIA gen nov.
Annual succulent plants, somewhat resembling species 24
Commelina: stems 2-8 dm. long, erect, ascending, or decumben
*Since I began the study of this species with the view of determining the
propriety of retaining it in Tinantia, I have learned that Dr. J. K. Small had
decided to segregate it as a distinct genus in the next edition of his Flora of
the Southeastern United States.
+
A NEW GENUS OF THE COMMELINACEAE 274
glabrous, much branched at maturity, the branches ascending,
breaking through the close-fitting leaf-sheaths: basal leaves 0.5—
3.5 dm. long, acute or acuminate, smooth or slightly pubescent
bract; calyx of three similar, green, ovate, imbricated, persistent
sepals; corolla showy, the petals equal or very unequal; stamens
six, all fertile but dissimilar in size and in degree of pubesc cence;
ovary oblong, three-celled, with two superimposed ovules in
the transverse embryo.
“ Commelinantia anomala (Torr.) comb. nov.
Tradescantia anomala Torr. U.S. & Mex. Bound. Bot.
225. 1859.
Tinantia anomala C, B. Clarke; De Candolle, Monog, Phaner.
3: 287. 1881.
Pea green, usually glaucescent SOs: stems clustered,
at first simple but later much branched, more than one branch
frequently arising at a ee nig Sanaa. in the upper part
of a stem late in the season) ; basal leaves 2—3.5 dm. long, linear-
spatulate, attenuate into long ciliate petioles below, rather aes
ches glaucescent, slightly os with a few long hair.
above; lower cauline leaves 1.5-2.5 dm. long; upper pees
pees sessile at “the top of the sheaths or eked. 0.5-2
ng, broadly or narrowly lanceolate or somewhat cordate
and clasping at the base, acute or acuminate; inflorescence
erect, shorter (4-8 cm. long), broader, and strongly cordate-
clasping at the base and in having no sheath; flowers three to
very scrote lavender blue one an avery small and in con-
aus anterior white petal, the showy petals almost rhomboid,
1.5-1.8 cm. long and somewhat broader than long, the small
petal ovate-rhomboid, 3-4 mm. long, and longer than broad ;
stamen densely bearded (except at the extreme apex, at the
274 A NEW GENUS OF THE COMMELINACEAE
extreme base and in the lower two-thirds of the inner face) with
showy lavender-purple hairs, (b) the two postero-lateral densely
bearded above the middle with a collar of short lavender-purple
yellow-tipped hairs, (c) the two antero-lateral bearded below
the middle on the posterior margin with a tuft of short lavender
hairs, and (d) the anterior stamen glabrous throughout—anthers
of the three anterior stamens oblong, these stamens being mani-
festly larger throughout than the posterior stamens, the latter
with orbicular anthers; ovary constricted in the middle, the
stigma terminating a slender bent white style; capsule oblong,
ong; seeds 3-3.5 X 2-2.25 mm., rounded at one end,
) ear
eccentric perforation containing the minute embryo; endosperm
not continuous but separated by an obscure commissure on one
side and bearing a group of low broad, tuberculate ridges radi-
ating from the sides of the embryo cavity, where it comes through
to the outer surface.
In rich, moist, shaded soil of ravines and rocky hillsides of
the Edwards Plateau, Texas; February to October, but princi-
pally March to May. Original localities, according to Torrey:
“shady woods on the Blanco, Comale, and other rivers,
Texas; Wright, (No. 699.) San Antonio, Texas; Thurber.”
A comparison of Torrey’s description with that submitted
herewith will reveal several places wherein certain differences
appear. These differences are: (1) in the height of the plant;
(2) length of basal leaves; (3) number of flowers per raceme;
(4) I can find no evidence of wings on the margin of the posterior
filament and am at a loss to account for Torrey ’s mention of such
structures unless he examined the filament in question only
from the flat naked inner face of pressed material; (5) the ovary
is uniformly three-celled with two superimposed ovules in each
cavity, though failure of fertilization will of course cause abortion
of any one or more of the six, with a consequent great variation
in the appearance of the capsule; (6) the seeds, though super-
ficially resembling those of Tradescantia, are different from any
Tradescantia seeds I have examined in that the cavity containing
the embryo extends entirely through the seed, the endosperm
instead of being continuous around the embryo is separated by
a commissure that extends to the embryo-cavity on one side,
and the seeds of C. anomala are of greater size. Other points
which are brought out in the description submitted herewith
are not mentioned in that of Torrey.
VOLUME 49, PLATE 10
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COMMELINANTIA ANOMALA (Torr.) THARP
A NEW GENUS OF THE COMMELINACEAE 275
J am indebted to Dr. F. W. Pennell, now of the Philadelphia
Academy of Sciences, for helpful criticism and advice, for
assistance in looking up the literature, and for access to a speci-
men of 7inantia Pringlet.
UNIVERSITY OF TEXAS.
Explanation of plates 10 and 11
PLATE 10
Plant of Commelinantia anomala (Torr.) Tharp, pitt typical appearance
of a flourishing specimen in bloom. The specimen is somewhat wilted causing
the spathe to fold slightly. The linear-spatulate basal es. characteristic
of plants during winter months, have disappeared by the time the stems
have reached this stage of maturity. Scorpoid racemes seen in profile. Scale
in inches.
PLATE II
eg 1. Inflorescence of C. anomala, its the position assumed by each
it comes into bloom. The spathe is somewhat wilted, causing it
= t0 voll “slightly at the edges. er ‘metric.
Fic. 2. Dissected flower to show f part
Fic. 3. Stamens from a bud several hours before anthesis; magnified hie
diameters. The filaments have not attained nearly the length of those in an
shape of mature anthers, and also the nature of the pubescence of filaments
well shown.
INDEX TO AMERICAN BOTANICAL LITERATURE
1912-1922
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or = upon American material; the word
America being used in the broadest se
Reviews, and papers that relate ae to foresty, agriculture, horti-
culture, poponmners products’ of vegotabie origin, or Poets ratory methods
nd literature of bacteriology.
An occasional Et Re is made in favor of some paper appearing in an Ameri-
can 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 three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Johnson, A. G. Wheat scab and its control. U.S. Dept. Agr.
Farmers’ Bull. 1224: 1-16. f. 1-12. O 1921.
Juday, C. The plankton. In Evermann, B. W., & Clark, H. W.
Lake re a physical and biological survey 2:
105-116. 1920.
Kaiser, G. B. Little journeys into mossland, IV. Bryologist
24: 41-43. 21 S 1921.
Kasai, M. On the morphology and some cultural results of
Fusarium Solani Mast. Appel et Wollenweber, an organism
which causes dry rot in the Irish potato tubers. Ber. Ohara
Instit. Landw. Forsch. 1: 510-542. pl. 8-11. 1920.
Kauffman, C. H. A black rot of squash. Michigan Acad.
Sci. Rep. 22: 201, 202. Au 1921
Kauffman, C.H. Collybia strictipes, developed in the Peseaseie
Michigan Acad. Sci. Rep. 22: 203, 204. Au 1921
Kauffman, C. H. Mortierella Baineri. Michigan Acad. Sci.
Rep. 22: 195-198. pl. 16. Au Ig2I.
Kurtz, F. Atlas de las plantes fosiles de la Republica Argentina.
Actas Acad. Nac. Ci. 7: 133-153. 1921.
277
278 INDEX TO AMERICAN BOTANICAL LITERATURE
Laibach, F. Untersuchungen iiber einige Septoria Arten und
ihre Fahigkeit zur Bildung héherer Fruchtformen III und LV.
Zeitschr. Plfanzenkr. 31: 161-194. 1921.
Lee, H. A. Citrus-canker control: a progress report of experi-
ments. Philippine Jour. Sci. 19: 129-175. pl. 1-2. Au 1921.
Lloyd, J. U. Plant pharmacy. Am. Jour. Pharm. 94: 238-244.
922.
Lowe, E. N. Plants of cea Mississippi Geol. Surv.
Bull. 17: 1-293. F 1921
Ludwig, C. A. A curious variation in the common milkweed.
Proc. Indiana Acad. Sci. 1920: 243-245. f. 1-6. 1921.
Lyon, T. L., & Wilson, J. K. Liberation of organic matter by
roots of growing plants. Cornell Univ. Agr. Exp. Sta. Mem.
40: 1-44. f. 1-9. Jl 1921.
McLean, F. T. The permeability of Citrus leaves to water.
Philipp. Jour. Sci. 19: 115-124. f. z. Jl 1921.
Marchionatto, J. B. Acacia visco Lorentz, Syn. A. platensis
Manganaro. Revisto Facult. Agron. Univ. Nac. La Plata
42: 90-95. 1921.
Marcou, P. Cacao, cacahult ou cacaouete. Jour. Soc. Ameri-
canistes Pasia II. 12: 65-67. 1920.
Maxon, W. R. Notes on American ferns—XVII. Am. Fern
Jour. 11: 33-39. I921.
Includes Selaginella scopulosum, sp. nov., from Montana.
Molisch, H. Der Naturmensch als Entdecker auf botanischem
Gebiete. Schr. Ver. Verbr. Naturw. Kenntn. Wien 54: REA:
je 1, 2:40
Muir, F. The origin of the Hawaiian flora and fauna. Proc.
Pan-Pacific Sci. Confer. 1: 143-146. 1921.
Murrill, W. A. John Macoun. Mycologia 13: 264. 80 1921.
Murrill, W. A. Edward T. Harper. Mycologia 13: 264, 265.
80 1921.
Nestler, A. Einige Beobachtungen an der Paprikafrucht. Ber.
Deutsch. Bot. Ges. 39: 230-234. Je 1921. .
INDEX TO AMERICAN BOTANICAL LITERATURE 279
Newcombe, F. C. Response of sensitive stigmas to unusual
stimuli. Michigan Acad. Sci. Rep. 22: 145, 146. Au 1921.
Patouillard, N. Une nouvelle lepiote du Bresil (Lepiota Putte-
maneit). Bull. Trimest. Soc. Mycol. 37: 81-83. 20 Jl 1921.
Pemberton, C.C. Some notes of the growth of Arbutus Menziesii
Pursh. Canad. Field-Nat. 36: 21-26. f. 1-4. F 1922.
Petry, E. J. Germination and growth of Ceanothus americanus
as affected by heated soils. Michigan Acad. Sci. Rep. 22:
135-143. pl. 13, 14.. Au 1921.
Pfeiffer, H. Conspectus Cyperacearum in America meridionali
nascentium. I. Genus Heleocharis R. Br. Herbarium 1921:
65-69. 1921.
Praeger, R. L. An account of the genus Sedum as found in
cultivation. Jour. Roy. Hort. Soc. 46: 1-314. f. 1-85. 6 Je
1921
Reinking, O. A. Higher Basidiomycetes from the Philippines
and their hosts, V. Philipp. Jour. Sci. 69: 91-114. ji 1921.
Renner, O. Heterogamie im weiblichen Geschlecht und Em-
bryosackentwicklung bei dem Onotheren. Zeitschr. Bot.
13° 006-021. - 7. 2-$. .. 1021:
Rhoads, A. S. Some new or little known hosts for wood-des-
troying fungi. III. Phytopathology 11: 319-326. Au 1921.
Ridlon, H. C. A new eee from Vermont. Am. Fern
Jour. 11: 46-48. pl. 7 a:
Polypodium vulgare f. anager
Rodway, J. Fauna and flora [of Guiana]. In Rodway, J.,
Guiana 237-248. London. 1912.
Rudolph, B. A., & Franklin, H. J. Studies of cranberries during
storage. Fungi studies. Massachusetts Agr. Coll. Bull.
198: 88-92. f. I, 2. 1920.
Safford, W. C. Cultivated plants of Polynesia and their vern-
acular names, an index to the origin and migration of the
Polynesiana. Proc. Pan-Pacific Sci. Confer. 1: 183-187. 1921.
280 INDEX TO AMERICAN BOTANICAL LITERATURE
Sampson, H. C. An ecological survey of the prairie vegetation
of Illinois. Bull. Illinois Nat. Hist. Surv. 13: 523-577. pl.
48-77 + f. 1-8 + maps I-3. Au 1921.
Saunders, E. R. Address to the Botanical Section [British
Association for the Advancement of Science, 1920]. British
Assoc. Adv. Sci. Rep. 88: 169-190. 1920.
A genetical address.
Sax, K. Sterility in wheat hybrids. I. Sterility relationships
and endosperm development. Genetics 6: 399-416. Jl 1921.
Scala, A. C. Contribucién al estudio histolégico de la flora
chilena. Revista Chilena Hist. Nat. 24: 73-78. f. 5-7. 31 Au
1920.
Schiiroff, P. N. Die Teilung des vegetativen Pollenkerns bei
Eichhornia crasstpes. Ber. Deutsch. Bot. Ges. 40: 60-63.
fe Pe 25 FF 1gz2.
Serre, P. La culture du cacaoyer au Costa-Rica. Bull. Mus.
Hist. Natur. 1921: 260-264.
Showalter, A.M. An orthotropous ovule in Hyacinthus orientalis
Ls Torreya 21+ 62,64. f. 7, 2. Aw’ 1921.
Silveira, A. A. Especies novae civitatis Minas Geraes |Brazil}.
Arch. Mus. Nac. [Brazil] 23: 159-171. pl. 1-5. 1921.
New species in Aspidosperma (1), Eriocaulon (4).
Small, J. K. Seminole bread—the conti. A history of the genus
Zamia in Florida. Jour. New York Bot. Gard. 22: 121-137.
pl. 258-259. 1921.
Includes Zamia umbrosa, sp. nov.
Smith, C. P. Studies in the genus Lupinus—VI. The Stiver-
stant, Concinni and Subcarnost. Bull. Torrey Club. 48:
219-234. 07-74. 210: 1921; :
Smith,A.L. Lichens. Pp. 1-464. f. 1-135. | Cambridge,
England. 1921. Inseries of Cambridge Botanical Handbooks.
Smith, L. Sugar cane in St. Croix. Virgin Is. Agr. Exp. Sta.
ce 2s es. Bd fo 1921.
Spegazzini, C. Mycetes chilenses. Bol. Acad. Nac. Ci. [Are-
entina] 25: 1-124. 1921. [Illust.]
Includes 5 genera and 113 new specie and forms of Fungi.
INDEX TO AMERICAN BOTANICAL LITERATURE 281
Spegazzini, C. Una nueva especie argentina del genero “ Pro-
sopanche.”’ Anal. Soc. Ci. Argentina 92: 251-256. f. 1-4.
D 1921
Prosopanche Mazzuchii sp. nov.
Spegazzini, C. Plantas nuevas o interesantes. Anal. Soc. Ci
Argentinisa 92: 77-123. S Igat.
Includes new species in Atriplex (1), Portulaca (2), Talinum (1), Echin-
ocactus (1).
Sprague, T. A. Alphabetical list of nomina conservanda [Phan-
erogamae]. Kew Bull. Misc. Inform. 1921: 321-326. 1921.
Sprague, T. A. A revision of the genus Belotia. Kew Bull. Misc.
Inform. 1921: 270-278. 1921.
Includes 5 new species from tropical America.
Sprague [T. A.| & Riley [ |. Petastoma Broadwayi, Sprague
et Riley [Bignoniaceae]. Kew Bull. Misc. Inform. 1921:
S14, ia 1971.
Sprague [T. A.]. Rinorea Hummelii, Sprague [Violaceae]. Kew
Bull. Misc. Inform. 1921: 307, 308. 1921.
Stevens, O. A. Pants of Fargo, North Dakota, with dates of
flowering Am. Midl. Nat. 7: 135-156. S 1921
Stewart, F. C. The velvet-stemmed Collybia—a wild winter
mushroom. Ann. Rep. (Geneva) New York Agr. Exp. Sta.
37: 208-227. 1919. [Illust.]
St. John, H. A critical revision of Hydrangea arborescens.
Rhodora 23: 203-208. 9 Ja 1922.
Stout, A. B. Sterility and fertility in a of Hemerocallis.
Torreya 21: 57-62. f. 1-8. Au 192
Stout, A. B. Studies of grapes in cooperation with the State
Experiment Station at Geneva, N. Y. Jour. New York
Bot. Gard. 22: 148-156. pl. 262. My 1921.
Tahara, M. Cytologische Studien an einigen Kompositen.
Jour. Coll. Sci. Tokyo Imper. Univ. 437: 1-53. pl. 1-4.
30 Mr 1921.
Tilden, J. E. The study of Pacific Ocean algae. Proc. Pan-Pacif.
Sci. Confer. 1: 207-209. 1921.
282 INDEX TO AMERICAN BOTANICAL LITERATURE
Torrey, G. S. Les conidies de Cunninghamella echinulata
Thaxter. Bull. Trimest. Soc. Mycol. 37: 93-99. pl. 10, f.
9-13. 20 Jl 1921.
Torrey, G. S. Coronella nivea Crouan. Bull. Trimest. Soc.
Mycol. 37: 88-93. pl. 10, f. 1-8. 20 Jl 1921. ,
Turrill, W. B. Plants introduced to horticulture from Chile and
Argentina (including Patagonia and Fuegia). Jour. Roy:
Hort. Soc. 46: 346-350. 6 Je 1921.
Urban, I. Flora domingensis. Symb. Antill. 8: 481-860.
1 Jt 1621.
Includes new species in Lyonia (3), Bumelia (1), Matastelma (1), Cordia
(1), Tournefortia (1), Ruellia (1).
Hook, J. M. Indiana fungi—V. Proc. Indiana Acad.
Sci. 1920: 209-214. 1921.
Van Hook, J. M. The pycnidium of Cvtcinnobolus. Proc.
Indiana Acad. Sci. 1920: 215, 216. f. 7. 192
Van Hook, J. M. A tricotyledonous bean. Proc. Indiana Acad.
ees, 1080s 207. fe tc. 1981
eo Melocactus depressus Hooker. Monatsh. Kakteenk.
Mr 1922. [Illust.]
— a
Walker, J. C., & Jones, L. R. Relation of soil temperature and
other factors to onion smut infection. Jour. Agr. Research
22: 235-262. pl. 25—27 + f. 1, 2. 29 O 1921.
Warnstorf,C. Die Unterfamilie der Scapaniodeen (Spruce 1885).
Hedwigia 63: 58-116. Ig921.
Includes 7 new American species.
Weatherby, C. A. Old time Connecticut botanists and their
herbaria Il. MRhodora 23: 122-125. Au 1921; 171-177.
O 1921.
Weatherby, C. A. Other records of Salivina natans in the United
States. Am. Fern Jour. 11: 50-53. 1921.
Additional tothose from Minnesota;see Butters, “‘Salvinia in Minnesota.”
Weatherwax, P. Anomalies in maize and its relatives—I. Bull.
Torrey Club 48: 253-255. f. 1-5. 1921.
Vol 49. No. 10
BULLETIN
OF THE
TORREY BOTANICAL CLUB
OCTOBER, 1922
The genus Costus in Central America
W. W. ROWLEE
(WITH PLATES 12-15)
As at present organized the genus Costus L. is composed of
about one hundred species, divided almost equally between the
Eastern and Western Hemispheres. Very few species occur in
both hemispheres. The American species are most abundant
in regions of heavy rainfall, maximum humidity and high
temperature of both air and soil. The genus ranges from one
Tropic to the other; it occurs wherever conditions are favorable
in hot, warm and temperate regions, but is not found in high
and cold districts.
In any given locality the species do not usually appear abun-
dant, but are scattered about in deep woodlands and undisturbed
jungle. The plants are herbaceous perennials with spirally
arranged leaves. The stems or culms are simple and represent
branches arising ‘from a system of underground rootstocks.
They thus form gregarious clusters, some of the culms 1emaining
sterile and others producing spikes of flowers. In some cases the
mats are more extensive than in others, this depending partly
upon age and partly upon the habit of the plant. The leaves are
ovate to lanceolate in outline and invariably entire.
In Spanish American countries members of the genus are
called ‘‘canagria’”’ or ‘‘cana agria,”” meaning “bitter cane,”
[The BULLETIN for September (49: 259-282. pi. 10, 11) was issued Sep-
tember 22, 1922.]
283
284. RowLee: GENUS Costus IN CENTRAL AMERICA
all jointed and unbranched stems being known as ‘“‘cana”’ and
the tissues in the present instance being very bitter. The plants
are fleshy throughout their whole body structure, and their
tissues are saturated with water, so that it is difficult to make
satisfactory herbarium specimens. Certain parts of the plant
and especially the spikes are so saturated as they grow in the
field that it is easy to wring water from them. In drying they
change their form markedly, and it is therefore most desirable
to make observations on specimens growing under natural
conditions. The flowers in many cases are delicately colored
and become fragile when pressed.
The genus Costus, together with two or three other genera,
belongs in the subfamily Costoideae of the family Zingiberaceae,
the group being very distinct in its general characteristics from
the other genera of the family. Two types have been recognized
in the genus: in the first the floral spike is borne upon what may
be called a normal leafy culm; in the second the floral spike
develops upon a modified culm. In the first type the culms are
designated as ‘‘not diverse;’’ in the second, as “diverse.”
The species heretofore recorded from the Western Continent
outside of South America have all belonged to the group in
which the culms are not diverse. In the present paper two
Central American species with diverse culms are reported and
proposed as new. They are not closely related to each other,
but each is represented by allied species in South America. The
paper reports in addition an undescribed species from Central
America in which the culms are not diverse and discusses the
previously described Central American species.
Key to the Central American species
A. Culms diverse, the spikes borne on short bracteate
pes.
B. Bracts of spikes with foliaceous appendages
CO AG 1. C. bracteatus
BB. Bracts of spikes not appendiculate (Guate-
Maley oo Sree) ee Basics 14 2. C. sepacuitensis.
AA. Culms not diverse, the spikes borne on ordinary
leafy ones.
B. Bracts of spikes ae! a appendages.
L. _ ovoid,
. Plants i 2-3 m. tall; leaves
about 30 cm. long, fuscous-hirsute,
especially on the sheaths Fae 3. C. villosissimus.
ROWLEE: GENUS CosTUS IN CENTRAL AMERICA — 285
DD. tee small; leaves about 16 cm.
g (Mexico, Costa Rica and Pana-
pice ae eke a ele eet ee. eee 4. C. hirsutus.
GG; Spikes teased red.
D. Appendages ie bracts lanceolate,
ae ee upper sterile erect;
leaves glabrous (Guatemala and Costa
Ri
1A enters iert oan amet eS es 5. C. Bakert.
DD. Appendages of bracts broadly ovate;
leaves scabrous (Costa Rica) ...... 6. C. lima.
BB. Bracts of spikes not appendiculat
&: a lacking on back of aie (Costa
PUM) Chee eee eee icicle ea 7. C. Malortieanus.
CL. Callose present on back of bra
D. Spike ovoid; leaves sessile yee 8. C. pulverulentus.
DD. Sih fusiform (Costa Rica an
(suatemiala) sce a ae he ties
DDD. sae eplindinical or spherical.
E. Plants small, about 2 m. high.
. C. sanguineus.
©
us.
Spike red, cylindrical
(Costa Ricaand Panama) Io. C. spicatus.
GG. Sp eo green, spherical
(Guatemala) i. 25 oboe: 11. C. congestus.
FF. Leaves oe beneath (Costa
Patiama ) 0s. 12. C. nutans.
Ric
EE. Plant large, much more than 2 m.
high.
F. Labellum orange-red
yellow markings (Costa Rica
and Guatemala). <. 4.4.2: 13. C. splendens.
. Costus bracteatus sp. nov.
Plant clothed a long black hairs; —— culms up to 2 m.
high, the lower leaves reduced to sheathing scales; flowering
culms 40 cm. tall, the leaves geetich eines except the four
or five uppermost, which bear small baie and closely surround
the spike. Uppermost teaver Di on the sterile culms large and thin,
32 cm. long, 7 cm. wide, oblanceolate, acuminate at the apex
and tapering below evenly to an acute base, ~~ hairy
on both sides, the hairs arising from a bulbous midrib
ue
"oO
=
Qo
om
je.
=
ie)
pe
fe)
8
©
ne
— ts
“én
Soe
—
i
Bae,
oO
c.
2
oO
W
—
QO
a
o
S
ge
Q
¢°)
> ©
Hi
- &.0
am
ae
=
f
ginous-tomentose; ligules 5-10 mm. long, tomentose; sheaths
* This fraction indicates that there are 3.5 primary and 23 secondary veins
to the centimeter.
286 RowLeEE: GENUS CosTUS IN CENTRAL AMERICA
overlapping, spreading-hirsute where exposed and with long
appressed needle-like hairs where included in the next lowe
sheath: lower leaves reduced to sheaths with oblique orifices:
|
most leaves forming a transition to the bracts of the spike.
Spike ovate-cylindrical, 13 cm. long, 6 cm. wide, conspicuously
bracteate, the bracts oblong, 2-5 cm. long, 1-8 cm. wide, puberu-
ent on the back and along the margins, strigillose within;
appendages of the bracts narrowed at the base, ovate, acuminate,
rusty puberulent: bracteole on the right side of the flower,
the back near the ee acuminate, rusty-puberulent. Flower
3-5 cm. long; ovary 5 cm. long, three-ribbed, the ribs densely
lobed, very thin and delicate, nearly equalling the labellum,
nearly as broad as long, reddish orange in the upper part: sta-
minodia equalling the labellum, also reddish orange at the tip;
stigma spoon-shaped with a notched appendage on the back.
[PLATE 12
Costa Rica: in woods on the hillside about a mile south of
Siquirres, H. E. Stork (Rowlee & Stork 675). Specimen pre-
served in the herbarium of Cornell University.
Only a single mat of the species, including one flowering culm
and several leafy culms, was found, although careful search
was made for additional material. The plant differs from any
other species known to us and is remarkable in its peculiar
vesture and delicate texture. It appears to be most closely
related to K. Schumann’s Costus Dinklagei from Cameroon,
Africa. The locality where this novelty was found is interesting.
The old Spanish trail from the mouth of the Matina River leads
westerly along the foot of the mountains to near Siquirres.
It then ascends the mountains along the south side of the gorge
of the Reventazon River to the central plateau, where the cities
of Cartago and San Jose are located. Our plant was found near
this trail. There was also found near the same place a spur
(for horse-back riding) quite unlike any known to the present
generation and probably dating back to the days of the buccaneer.
2. Costus sepacuitensis sp. nov.
Vegetative and fruiting culms diverse: leafy culms with large
elliptic leaves (rese esembling those of C. maximus), at least 30 a
lon 9 cm. wide, softly pubescent on both sides, margi
ciliate; petiole 5 cm, long; ligule 1.5 cm. long, ciliate on tie
RowLeEE: GENUS Costus IN CENTRAL AMERICA 287
margin, pubescent on upper part, glabrescent below; venation
/ long
Spike not much exceeding the culm in thickness and appearing
like it in texture et in color (dark chestnut in dried specimens) :
bracts broadly o , 4 cm. long, and 3 cm. wide, glabrous,
highly polished yoga within. Flowers not seen. Fruit
obovoid, pubescent, 1.5 cm. long and 0.8 cm. wide, crowned with
a deeply parted calyx; calyx lobes unequal, 1.3 cm. long, separated
to near the base, ovate, acute. [PLATE 13.]
GUATEMALA: near the Finca Sepacuite, Alta Verapaz, April
13, 1902, O. F. Cook & R. F. Griggs 596. Known only from the
type specimen, which is in the U. S. National Herbarium.
‘This remarkable plant is related to C. geothyrsus K. Schum.
of Ecuador and C. erythrocoryne K. Schum. of Peru but differs
from both in the shape and size of the spike, in the calyx, and
in the vesture of the plant. Other species with diverse culms
occur in South America and in the Eastern Hemisphere.
3. COSTUS VILLOSISSIMUS Jacq. Fragm. 51. pl. So. 1800-1810
Specimens referable to this species from Central America
and the West Indies exhibit considerable variation, although
agreeing in stature and in having appendiculate green bracts
and ovate spikes. The hairiness varies, for example, from tawny
shagginess to an almost smooth condition, while the color of
the flower varies from sulphur yellow to white. In all cases,
however, the margin of the labellum is tinted with pink. The
most typical specimen collected by H. E. Stork and the writer
is No. 401, found in June, 1918, on the west side of the Panama
Canal, opposite Balboa. The flowers are large and showy, the
labellum being tubular, about 7.5 cm. and ivory white
except for the rose red border. The whole plant and particularly
the upper leaves are clothed with long tawny hairs. This
specimen was 3 m. tall. Another specimen, also in bloom but
only half as tall, is No. 325a, collected in September, 1918, at
Quiriga, in Guatemala. It agrees with No. 401 in having tawny
hairs and in the color of its flowers.
The species, although apparently rare, is widely distributed in
tropical America. The following additional specimens from
Central America may be cited: near Balboa, Canal Zone,
Panama, H. Pittier fae 3779, 6690; Province of Alta Verapaz,
Guatemala, O. F. Cook & R. F. Griggs 308.
288 Row.EE: GENuS Costus IN CENTRAL AMERICA
4. Costus HIrsutus C. Presl, Reliq. Haenk. 1: 112. 1830
This is a much smaller plant than the foregoing, according
to the writer’s interpretation, and is evidently not more than
50 cm. high. The spike also is smaller. The following specimen
seems to be characteristic: on the cattle trail between the
Chiriqui Lagoon and the divide, western Panama, H. E. Stork
(Rowlee & Stork 1045). Another specimen to be associated with
the species is the following: upper stretch of the Saraquipi
River, H. Pittier 14166. Both represent a diminutive C. villo-
sissimus. The published descriptions of C. hirsutus are meager,
and no drawing or photograph of the species has been seen by
the writer.
5. Costus BAKERI K. Schum.; Engler, Pflanzenreich 4'*°:
387. 1904
The following specimens of this species, all from the uplands
of central and western Guatemala, have been studied: Barranca
del Samala, Dept. Retalhulen, 1700 ft. alt., J. Donnell Smith
2801; Barranca de Eminencia, Dept. Amatitlan, 1400 ft. alt.,
J. Donnell Smith 2802 (Type); Volcan Tecuamburro, Dept.
Santa Rosa, J. Donnell Smith 4650. The following specimen
from central Costa Rica has likewise been examined: A. Ton-
duz 282.
C. Bakeri is characterized by having a cylindrical and appendi-
culate spike, more or less red in color; the appendages of the
bracts being acute and longer than wide. In these respects it
agrees with C. comosus (Jacq.) Roscoe, of northern Colombia
and Venezuela, a species with which it was originally confused.
In C. comosus, however, the upper surface of the leaves is softly
hairy, while in C. Bakeri it is perfectly glabrous. It is probable
that intermediates between these two species will be found.
6. Costus LIMA K. Schum.; Engler, Pflanzenreich 44%: 388. 1904
Dr. Schumann apparently knew C. lima from the type speci-
men only, which was collected at the following station on the
Pacific side of Costa Rica: near Punta Arenas, January, 1854,
Scherzer. The species is frequent however, throughout the
lowlands in the northern part of Costa Rica, being found also
on the Atlantic side. The following additional specimens may
be cited: Buenos Aires, near the continental divide, A. Tonduz
6659; Livingston on the Reventazon River, Rowlee & Siork 648;
Siquirres, Rowlee & Stork 1318.
Row.LEE: GENus Costus IN CENTRAL AMERICA — 289
It agrees with C. Bakeri in having an appendiculate, cylindri-
cal spike, often much shortened, but the color is more brilliantly
red, making the spike very conspicuous. The appendages,
moreover, differ in being shorter and less striking. The species
is further characterized by the peculiar vesture of the very
opaque leaves, the upper surface being clothed with shorter
bulbous hairs, all pointing toward the apex, while the lower
surface is densely covered by a tawny tomentum. The plants
are large, often 5-7 m. high, and the spike is about the size of
an ear of corn.
According to the original account C. lima is related to C.
Friedrichsenit O. G. Peters., a species described from plants
cultivated at Berlin. Dr. Schumann states that it was probably
introduced from Central America, but the writer has seen no
specimen agreeing with the description.
7. Costus MALORTIEANUS Wendl. MHamb. Garten- u.
: Blumenzeit. 19: 30. 1863
So far as the writer’s observations go C. Malortieanus is
confined to the coastal plains of northeastern Costa Rica. The
type locality is near the Sarapiqui River at the head of navigation,
and the original description was drawn from specimens cultivated
in Germany. The species grows also along the Reventazon
River, where it emerges on to the lowlands. It is the gem of
the genus. The plants are small, being less than 1 nt. high, and
grow in wide spreading mats, flowering in July and August.
The flowers, which are not freely produced, are very beautiful,
suggesting in their tints some of the tropical orchids. The
leaves are broadly obovate, densely hairy, and obscurely variegat-
ed with darker and lighter threads of green. Specimens in the
greenhouses of the New York Botanical Garden, although under
cultivation for several years, have not as yet produced flowers.
Two other species of tropical North America with unappen-
daged bracts and no callose have beendescribed under the names
C. pictus D. Don, and C. mexicanus Liebm. Specimens of the
first two have not been found by the writer, but both are ap-
parently of Mexican origin, C. pictus being based on plants
grown in England.
290 RowLeE: GENUS CostUs IN CENTRAL AMERICA
8. CosTUS PULVERULENTUS C. Presl, Reliq. Haenk. 1:41. 1830
The original specimen of C. pulverulenius was collected in
Mexico by Haenke, no more definite locality being mentioned.
The writer has identified with it, somewhat doubtfully, the
following specimen from Guatemala, origina'ly referred to C.
Malortieanus: Rio Sis, Dept. Suchitepequez, 1300 ft. alt., J.
Donnell Smith 4800. This and the following six species agree
in having a callose on the back of the bracts.
g. CosTUS SANGUINEUS Donn. Sm. Bot. Gaz. 31: 122. 1901
This species is the most frequent representative of the genus
in northern Costa Rica, and a photograph of it has been repro-
duced in Calvert’s ‘A year of Costa Rican Natural History,”’
opposite page 257, under the incorrect name C. Malortieanus.
It is not found on the beach but inhabits higher ground, often
at an elevation of as much as 2000 feet. The type specimen
was collected in 1901, at Cubilquitz, Dept. Alta Verapaz,
Guatemala, about 1000 ft. alt., by H. von Tuerckheim (No.
7686), but the species has been found in other Central American
countries bounding the Carribbean Sea. The following specimens
may be cited: Peralta, Rowleé & Stork 45; Livingston on the
Reventazon River, Rowlee & Stork 653; and Siquirres, Rowlee &
Stork 1311, 1312. The plant is about 1.5 m. high, and the
leaves are softly pubescent. The red spikes are fusiform and
acute, when perfectly developed, as shown in Calvert’s figure,
but they are frequently deformed and much shortened.
10. Costus SPICATUS (Jacq.) Sw. Prodr. Fl. Ind. Occ. 11. 1788
The original material of C. spicatus came from the West
Indies, where the species is widely distributed. It is found,
also, in the close vicinity of the seashore in northern Costa
Rica, between Limon and the mouth of the Matina River. The
following specimens may be referred to it: Chargres, 1850.
A. Fendler 446; Rio Dulce, March, 1889, J. Donnell Smith;
vicinity of Frio, province of Colon, August, 1911, H. Pittier
4137; Limon, Rowlee & Stork 674. The field notes that follow
were made from No. 674.
Culms 1-2 m. high, many from a rather dense mat, leafy to near the base,
whole plant glabrous. .Leaves 18 cm. long, 7 cm. wide, elliptic, short-acuminate
ROWLEE: GENUS CosTUs IN CENTRAL AMERICA = 291
at the apex, subauriculate at the base, shining green above, giacuous be-
neath, petioles 6 mm. long, sheath shorter than the internodes, ligule 5 mm.
long, three or four leaves approximate to the base of the spike, venation 7735.
Spike oblong, 7 cm. long, 2.3 cm. thick, very compact. Bracts nearly orbi-
cular, 2.5-3 cm. in diameter. Bracteole 1.5 cm. long, relatively broad, boat-
siaped dextrad. Ovary 5 cm. long, glabrous, subtrigonous, seeds very numer-
ous, angular with elater-like aril. Calyx 5 cm. long, campanulate and with
very short broad lobes and red margins. Corolla orange-red, 2 cm. long,
dilated at the apex, lobed to the middle, lobes oblong, acute. Labellum 3 cm.
long, obovate, subtrilobed, yellow.- Stamen 2.5 cm. long, lanceolate, obtuse,
anther adnate above the middle.
. Costus congestus sp. nov.
Stem less sa 1 m. high. es akin near the summ mit;
m.
m. lo
base, the lobes unequal, oblong, pended at the ora petals
large, Ryaline, obtuse; labellum 4 cm. long. [PLATE 14.]
GUATEMALA: Escuintla, 1100 ft. alt., March, 1890, J. Donnell
Smith 2036 (Type); Santa Lucia, Dept. Escuintla, 1045 ft. alt.,
March 3, 1905, W. A. Kellerman 5284; Santa Barbara, Dept.
Solola, 1370 ft. alt., August, 1891, J. Donnell Smith 137. Type
specimen in the herbarium of Cornell University.
Mexico: vicinity of Choapam, pees, 3800-4500 ft. alt.,
July 28, 1894, E. W. Nelson git.
The species is glabrous in all its parts. It differs from C.
spicatus (Jacq.) Sw., to which Baker referred some of the speci-
mens, in the size of the flower and in the shape and size of the
spike.
12. Costus NuTANS K. Schum.; Engler, Pflanzenreich 4**: 407.
1904
The type material of C. nutans came from the following local-
ity: near Aguacate, Costa Rica, September, 1857 (in flower), C
Hoffman 727.
The only specimen seen by the writer was collected on Mount
Pirri, Panama, on June 12, 1912, by E. A. Goldman (No. 1963)
at an altitude of 1350 m. [PLaTE 15.
292 ROWLEE: GENUS CosTUS IN CENTRAL AMERICA
13. COSTUS SPLENDENS Donn. Sm. & Tuerckheim; Donnell
Smith, Bot. Gaz. 33: 260. 1902
Costus maximus K.Schum.; Engler, Pflanzenreich 44°: 405. 1904.
Costus giganieus Kuntze, Rev. Gen.2:687. 1891. Not Ridley,
1887.
The species was based on a specimen collected in Guatemala
at an altitude of 350 m., by H. von Tuerckheim (No. 8015), no
more definite locality being mentioned. The writer has not seen
the type specimen, but other material collected in Guatemala by
Von Tuerckheim and referred to C. splendens has been available
for study. Jn this material the bracts bear a callose below the
apex, although no structure of this sort is mentioned in the
original description. Since these specimens are indistinguishable
from C. maximus K.Schum., the name C. splendens will have to
supplant the later name C. maximus, and the known range of
the species will extend from Guatemala to Panama.
14. Costus LAxus O. G. Peters.; Martius, Fl. Brasil. 3%: 56.
1890
The type specimen was collected in Costa Rica by Aented
in 1847, the name of the locality being illegible. The species
is known also from Venezuela.
CORNELI. UNIVERSITY
Explanation of plates 12-15
PLATE 12. CosTUS BRACTEATUS Rowlee.
PLATE 13. COSTUS SEPACUITENSIS Rowlee.
PLATE 14. Costus CoNGEsTUS Rowlee.
PLATE 15. Costus nuTANS K. Schum.
BuLL. ToRREY CLUB VOLUME 49, PLATE I2
COSTUS BRACTEATUS ROWLEE
BuLL. Torrey CLuB VOLUME 49, PLATE 13
COSTUS SEPACUITENSIS ROWLEE
Butt. TorREY CLUB VOLUME 49, PLATE 14
COSTUS CONGESTUS ROWLEE
BuLL. TORREY CLUB VOLUME 49, PLATE I5
COSTUS NUTANS_K._Scuum.
Quercus lyrata in Iowa
B. SHIMEK
(WITH PLATES 16 AND 17)
The discovery of the overcup oak, Quercus lyrata Walter,
in lowa, by the writer, so far extends the range of this species
that it seems worthy of more detailed record. Sargent says*
that the species “is distributed from the valley of the Patunxent
River in southern Maryland southward near the coast to western
Florida, through the Gulf States to the valley of the Trinity
River in Texas, through Arkansas and southwestern Missouri,
where in a swamp near Allentown, there is a single specimen,
the most northern known representative of the species west of
the Mississippi River, to central Tennessee, southern Indiana,
and Jasper County, Illinois.”’
Other authors give the distribution less fully, but in all the
references consulted the range falls within that here given. The
herbarium of the Missouri Botanical Garden at St. Louis
contains several specimens from the southern counties of Illinois,
and from Butler County, Missouri, but these localities also
lie within the territory given by Sargent.
The Towa specimens are located in lowa County, near the
town of Amana. This not only extends the northern range
west of the Mississippi, but the locality is more than two degrees
of latitude north of the Maryland and Lllinois localities, making
this the northernmost point from which the species is known.
Thus far three trees have been found. They are located in
the low bottom land timber along the Iowa River, the locality
being subject to overflow and always quite moist. They are
associated with the large, bottomland form of the bur oak,
Quercus macrocarpa, a few trees of Q. bicolor, and the ordinary
bottomland species of Iowa valleys, such as Ulmus americana,
Betula nigra, Populus deltoides, Acer saccharinum, A. Negundo,
Plaianus occidentalis, and Juglans nigra. Other trees of the
species will probably be found among the “bur-oaks.”’
he finest of the three specimens (shown in PLATE 16, FIG. 1)
is about 70 feet high, and the trunk measures 22 inches in diam-
eter at a height of 3 feet. Its rate of growth is probably approxi-
*Silva of North America 8: 48. 1895.
293
294 SHIMEK: QUERCIS LYRATA IN IOWA
mately the same as that of the large bottomland form of Q.
macrocarpa, and a nearby specimen of the latter, recently cut,
measured 27 inches in diameter and exhibited about 120 rings,
those from about 55 to 85 showing the most rapid growth. Our
overcup oak trees are’ probably not less than a hundred years
old, thus antedating the coming of the white man to this section.
The trees have been observed both in flower and fruit. Sar-
gent gives the time of flowering as March or April, but in this
northern locality the flowers do not appear until about the
middle of May.
Two of the trees produce acorns which are entirely enclosed
in the cup; the third has a partly open cup, a form not infrequent
in the South. The peduncles of the cups are 6-16 mm. in length.
The wall of the cup is 3-6 mm. thick or even thicker at the
base, but thins upward. The scales of the cup are prominently
tubercular, the tubercles becoming smaller upward. The small
opening has a short fringe.
The nut is short-ovate, with a broad basal scar and is 16-19
mm. in length, with about the same diameter. Its tops and
sides are covered with a short, grayish pubescence. For cup
and acorns see PLATE 16, FIG. 2.
The general aspect of the tree is not unlike that of the large
form of Q. macrocarpa which grows in somewhat better drained
spots in the same timber. The latter is here not gnarled or
stunted, as is usually the case in more exposed localities. It
is probable that our species has been mistaken for the latter in
other localities.
The leaves are distinctly different from those of the nearby
bur-oaks. As compared with the latter they are thinner, with
more distinct veinlets; lighter green; more irregularly lobed,
with the terminal lobe rarely coarsely crenate; usually smaller
and narrower; and the upper surface is more likely to have scat-
tered short hairs at maturity, while the pubescence of the lower
surface is much less dense. PLATE 16, FIG. 3, shows young
leaves, and PLATE 17 shows mature leaves taken in different
years. The leaves and acorns shown on the plates are from the
same tree.
On the whole the Iowa specimens of Q. lyrata are quite
typical. ;
Nom»
SHIMEK: QUERCUS LYRATA IN Iowa
Explanation of plates 16 and 17
PLATE 16
Quercus lyrata ieee a leafless tree.
ups and acorns, x .55.
Young leaves aid ef a twig, < 42;
PLATE 37
Mature leaves of Quercus lyrata, x .24.
Old leaves, taken a year earlier, x .30.
295
Buti. Torrey CLuB VOLUME 49, PLATE 16
3
QUERCUS LYRATA WaALTER
BuLL. TORREY CLUB VOLUME 49, PLATE 17
2
QUERCUS LYRATA WaALrTER
References to the algae in the Chinese classics
W. M. PorTERFIELD, JR.
(WITH ONE TEXT FIGURE)
The following notes on the algae referred to in ancient
Chinese literature are at best sketchy, but are offered neverthe-
less for what they are worth. The references to the texts and the
translations thereof are quoted on the authority of Mr. Y. T.
Chu, Instructor of Biology at St. John’s University, Shanghai,
China, and Mr. C. F. Wu, formerly of St. John’s, now of Cornell.
The writer presents this paper with a view to throwing some new
light on the development of science in general, and to intro-
ducing to the West, besides, some evidences of Far Eastern
activity in this line, which parallels, if not predates, that of
Europe.
Agriculture in China dates back to Shen Nung,* an emperor
of the legendary period, 3000 B. C. He was said to be the first
farmer and taught the people to till their fields. Since this
mythical age, the people of China have been farmers primarily.
As the Old Testament was essentially the expression of an
agricultural and pastoral people and is, in consequence, re-
plete with similes and references to plants and animals, so in the
Chinese classics we find the farming life of the people, a life
of continuous touch with nature, coming to expression in the
frequently recurring allusions to animals and plants, and in the
use of terms connected therewith. Long before the Aristotelian
age of hearsay and philosophical conjecture, the ordinary facts
of farming and floriculture had entered the realms of Chinese
literature in the form of terms serving as figurative expressions
for desirable characters and virtues. In order to have given
time for everyone to become so familiar with these original
terms that they crept unconsciously into speech and literature
as specific classifiers, observation of the form and structure of
plants must already have proceeded far. This is offered as
one evidence of age-old familiarity with the facts of nature.
The second is coincident with the first and deals with the ideo-
graphs representing these terms and ideas. The discovery and
use of the facts of nature called into being special ideographs to
* Appendix, “Peking,” by Juliet Bredon.
297
298 PoRTERFIELD: ALGAE IN THE CHINESE CLASSICS
represent them, so that owing to the character of the language
and its slow evolution, we feel that the appearance of these
specific terms in the ancient literature places the knowledge
back of them at a very early date
We shall confine ourselves to the algae in this paper. The
Chinese character, as we are accustomed to term the ideograph,
is built up of many radicals or elementary symbols supposed
formerly to have been pictures which taken together represent
an idea. The character for algae is
vat Tsao (Fic. 1) and may be resolved
* oo--- a into four radicals, the ones for grass,
caiidan! = water, wood, and mouth, the latter
: ~eceaes, ©. being repeated three times. The
Fic. 1. Tsao mouth radical is a simple rectangle,
a, grass radical; 6, mouth which when repeated three times and
radical; c, wood radical; e, water placed in the form of a pyramid,
radical; f, small ideograph mean-
iii Gee mechiant. two at the bottom and one on top,
€ up an ideograph which means
rank or character. This ok smaller ideograph seems to
carry with its meaning the idea of segmented, or possibly cellular
(!) structure because of the shape of the thrice repeated box-
shaped radicals. Evidently the idea in their minds from the
character used was that of a grass-like, fibrous or stringy,
cellular plant that grows in the water. Such an analysis leads
us to believe that they had in general a good idea of what an
alga is.
The character for Tsao first appeared in one of the five classics,
the Canon of Boems (Fic. 1). In the chapter Chao Nan there
appears the following passage, the romanized version of which
is here given: ‘‘‘yii bih bien tsao”’ (with respect to the collecting
of algae). This term is still used today. The term for water
plants is a much simpler character and therefore is much more
general in its meaning. It has the grass and water radicals but
no specifically descriptive ones. There can be no confusion
then in the use of these terms.
In support of the cellular or segmented idea of algae brought
out in the analysis of the character, K’ung An-kuo, in the Canon
of History, says that ‘‘an alga is an aquatic plant that has
systematically arranged branching parts, and is used, therefore,
(figuratively) to denote literature.”” He may here be referring
PoRTERFIELD: ALGAE IN THE CHINESE CLASSICS 299
to the diagrammatic structure of Hydrodictyon. Nearly two
thousand years ago in a pond beside the temple of Han Voo Tee,
algae were said to have grown to a length of nine feet. Among
other characteristics they were reported as having the appear-
ance of a net from which certain water birds were said to have
had great difficulty in extricating themselves. The people called
this the water-net alga. The structure of such types was
plainly visible under careful scrutiny, hence a netted or reti-
culate structure was specifically attributed to algae. _
About 600 B. C., in a book entitled Sze tsen, occurs the follow-
ing reference to algae: ‘“‘Some algae are a delicacy fit for the
most honorable guest, even for the king himself.”” At the
present time country people gather Nostoc for food. This is
called ‘‘Heaven vegetable.’’ Red algae are dried and eaten
by farmers who live ear the sea. It is to some of these doubt-
less that the quotation refers.
The real knowledge in Chinese medicine as it exists today is
based on the Chinese ‘‘ Materia Medica,”’ the edition of which
was begun four thousand years ago. The present edition was
written two hundred years ago in the Ming dynasty. Among
other plants, Kw’un Boo, or Laminaria is mentioned as being
useful for medicinal purposes, for which it is calcined after being
washed and sun-dried. It is a common practice in China to
pack open cuts with ashes in order to stop the bleeding. Whether
there is any discrimination shown as to what kinds of ashes are
used the writer is not prepared to say. If there is, it is interesting
to note that in case preference was shown for ashes of kelps,
they must have realized that some medicinal virtue was con-
tained in them. Iodine as an element was probably not known
to the Chinese but they may have — its presence as a
virtuous remedy in other.things.
sin western countries the sea weeds of the China coast are
used by the farmers nearby for fertilizer and also, when dried
for fuel. Agar-agar is made by the Chinese out of certain species
of sea-weeds, a well known fact to most scientists and technicians.
e morphological characters of these marine forms seem to
have been as well known as the freshwater forms, if not better.
Many were said to be attached to stones. The large leafed
forms were known as ox or horse algae. Some float on the top
of the water, others live at the bottom. Of the smaller forms,
300 PoRTERFIELD: ALGAE IN THE CHINESE CLASSICS
many have long silk-like filaments, the longer ones more than
thirty “segments” each, and others have the appearance of
“uncombed hair.”
Metaphorically the term for algae is used in a very compli-
mentary sense. The elegance and beauty of essays was often
designated by or compared to that of the algae. The term was
used in praise of the thinking of a learned man, signifying that
his thoughts were as systematically ordered as the parts of an
alga. Judgment was in like manner complimented. The
algae contributed not only to the language and literature of
the Chinese but also to their art and superstitions. In an old
book called Zong Shu, we find reference to conventionalized
designs derived from algae being included in the embroidery of
their garments. When the figures of algae appeared on the ends
of the roof beams of their houses in brilliant colors, we find that
the underlying idea was protection from fire. Because algae
were known to be water plants, any evidence of them on houses
was a protection against fire in that the former invoked the aid
of their native element to drive away the fire spirit.
From this discussion we begin to realize that from direct
references in ancient Chinese literature and an analysis of the
ideograph, there is a possibility, if not a probability, that the
knowledge of the algae as a distinct morphologic unit in the
plant kingdom dates back to very early times, as compared
with the state of knowledge in western countries. Moreover
this knowledge seemed to be more wide spread amongst the
people from the use of Tsao in a metaphorical sense, in its
practical use as medicine, as food and fuel, and as fertilizer; as
the basis of commercial products; and in the realization of its
decorative value and its superstitious meaning.
A Lachnea with a botryose conidial stage
B. O. DopGEe
(WITH SEVEN TEXT FIGURES)
While engaged in culturing species of Discomycetes for the
purpose of studying the nature of the origin of the ascocarp, an
interesting Lachnea was collected several different times in 1912
on a variety of substrata in the vicinity of New York City. The
apothecium of this fungus, which was identified* at the time as
Lachnea abundans Karst., originates in a long winding multicel-
lular ascogonium, so similar to that described by Fraser} for
L. cretea that, when we consider also the characters of the fruit
bodies and mycelium, there appears to be little doubt that the
fungi with which both of us were working are at least very
closely related species. Believing that the characteristics of the
primordia are of fundamental importance in determining re-
lationships, further discussion, other than the mention made
in a paper published at that time,t seemed unnecessary.
Miss Fraser does not state that she grew L. cretea in pure
cultures. Her cultures were originally derived presumably by
transfers from a plate culture overrun with moulds and some
hyphomycetous fungus. It would be necessary to know whether
L. cretea has a conidial stage like that which we have connected
with L. abundans before we can be certain that the species are
identical. In view of the discussions that have arisen since the
publication by Seaver and Horne* of a paper on Sclerotinia
Geranti and another paper by Godfrey} on Tite Ricint,
both of these forms having Botrytis conidial stages, it has been
thought advisable to point out that there are other types of
Discomycetes which also have asexual fructifications closely
resembling Botrytis.
* The Pasaentay iss has been confirmed ns Dr. F. J. Seaver, who is con-
sidering L. crefea as a possible synonym
ft Ann. Bot. py 553-563. 1913.
poses 9: 565- 567. 1910.
301
302 Dopcre: LACHNEA WITH A BOTRYOSE CONIDIAL STAGE
The conidia of Lachnea abundans are smooth at maturity
while those of Sclerotinia Geranii are rough or warted, otherwise
their conidial stages are very similar. The connection between
the apothecial and the conidial stage of L. abundans has been
established repeatedly by cultures from single conidia and from
single ascospores, apothecia arising in each case within a week
or two after showing the spores.
Fic. 1. Part of a dichotomously branched conidiophore showing the en-
larged ends of the ultimate branchlets upon which spores will be borne.
Fic. 2. End branches of a similar stage more highly magnified, spore
buds arising from two of the globular heads.
Fic. ortion of an old: conidiophore showing botryose clusters of
spores, A, developed on about sixteen fertile ends; B, shows the collapsed
ends of the conidiophore after the spores have been dislodged.
Fics. 4 and 5. Germinated ascospore becomes a cell in the hypha; F1G. 5
also shows ungerminated ascospores.
Ascospores germinate very readily when the inoculated culture
DopGE: LACHNEA WITH A BOTRYOSE CONIDIAL STAGE 303
medium is heated to 70° C. for about fifteen minutes. This
method has been found effectual in inducing germination of
ascospores of certain other species of Discomycetes.{ The out-
lines of the spore are generally obscured as it germinates and
becomes a cell in the hypha (Fics. 4, 5,). The conidia as well
as the ascospores remain viable for a long time if kept dry in the
laboratory, the former, sometimes living three or four
years. Should several conidia be sowed together in a culture
there follows at once a great amount of anastomosing of the
germ tubes.
The fungus grows well on almost any of the ordinary culture
media. Conidiophores arise the second day in cultures on milk,
bouillon, potato agar, etc. An agar medium in which the nutrient
is a decoction from heated soil is especially favorable for the
development of ascocarps. A potato dextrose agar gives an
abnormal amount of conidial development. The conidia are
formed on the spherical or knob-shaped ends of regularly dichoto-
mously branched conidiophores, seven or eight such divisions
often occurring. One might consider an aerial hypha whose
main axis ends in a pair of sporophores as a part of the coni-
diophore system even though branches ultimately bearing conidia
arise at irregular intervals from it. Fic. 1 shows a branch of the
third order at the time spore formation is just beginning. A
portion of such a branch is further enlarged in Fic. 2, showing
conidial buds from two of the ultimate branchlets. As spores
mature they hang together in botryose clusters covering about
eight pairs of end branches (Fic. 3, A). At B in this figure the
conidia have been dislodged, exposing the extremities of the
conidiophore subdivisions, now collapsed. The length of the
conidiophore system and the number of times dichotomous
branching occurs depend of course upon the kind of nutrient
in the medium. On soil decoction agar the sporophores are very
short, and there is very little of the aerial type of hyphae. The
mycelial hyphae on the other hand branch more or less dichoto-
mously and in this respect also the fungus is like the L. cretea
studied by Fraser. The color of the aerial hyphae, conidiophores
and conidia in mass varies from pale ochraceous buff to vinaceous
buff (Ridgway), depending on age and vigor of growth.
t Mycologia 4: 218-222. 1912.
304 DopGE: LACHNEA WITH A BOTRYOSE CINIDIAL STAGE
The conidia are spherical, smooth, faintly colored, pale
ochraceous buff in mass, 7-9 v. in diameter. The ascogonia
begin to appear, as noted, in about ten days, and mature fruit
bodies will be formed within two or three weeks. Fraser lays
considerable stress on the branching of the trichogyne end of
the ascogonium of L. cretea, thinking such vegetative growth
the “progressive degeneration” of Atkinson, indicates that the
species is becoming apogamous. Many such abnormal or
aborted ascogonia are always found in cultures of these Dis-
comycetes and they should be carefully distinguished from those
ed
xO
MSY
cae
‘
Bee see aut
Thy ae sy
seis
iy wea
I Pa
Fic. 6. Hairs from the margin of the apothecium, ascus with spores and
paraphyses.
1G. 7. Part of a section of a small ashe showing the character of
the cells of the wall and of the margin
normal primoridia which develop into fruit bodies. Ascocarps of
L. abundans (Fic. 7) are 1-3 mm. in diameter and uniformly
ochraceous in color. The hairs (Fic. 6) are about 150 u long,
the asci 130-150 u, and the ascospores 7-8 x 12-14 pn. The
paraphyses, which are enlarged rather abruptly at the ends, are
4-5 wu broad.
It is certainly dangerous to place too much weight on the
asexual spore forms in determining relationships of their ascogen-
ous stages in the face of these apparently anomalous cases and
such others as we findin Ascobolus magnificus, whichis connected
DopGE: LACHNEA WITH A BOTRYOSE CONIDIAL STAGE 305
with a Papulospora* similar to those forms hitherto thought to
belong to Melanospora. If one, following Saccardo, for example,
seeks to find a description which might apply to the conidial
stage of Sclerotinia Geranii or of Lachnea abundans he turns
directly to the forms along with Botrytis cinerea on the basis
of the spore clusters. It seems to the writer, therefore, an im-
possible task to determine the real generic affinities of species
of form genera from a study of the conidial or pycnidial stages
alone. Brierly* has recently published a paper on Botrytis
cinerea. His argument is based entirely on the assumption that
this species has no perfect or ascogenous stage. It is certainly
unusual to state that a fungus has no ascocarpic stage simply
on the basis of large numbers of cultures that produce only
conidial fructifications. The recent work of Bensaude, Kniep
and others, on several of the well known Basidiomycetes em-
phasizes the importance of growing in pairs strains or races
derived from different spores, at least as a last resort, in attempts
to obtain the “‘perfect’’ stages. We are finding more and more
forms in which a strain, which is sterile when grown alone, still,
when grown together with sume other strain, at once takes
part in the development of a ‘‘perfect”’ stage.
There is no question that Lachnea abundans is homothallic,
a culture from a single conidium or a single ascospore being
capable under suitable conditions of giving rise to ascocarps.
Whatever may be the standing of the species now commonly
referred to the form genus Botrytis, after study has revealed
their ascomycetous connections, it is interesting to find an
“imperfect” fungus with a botryose conidial stage connected
with a little Lachnea which may be so easily cultured on ordinary
media.
BUREAU OF PLANT INDUSTRY,
U. S. DEPARTMENT OF AGRICULTURE,
WaAsHINGTON, D. C.
yo 12: 115-134. 1920
* Phil. Trans. Roy. Soc. B. 210. 83- II4. 1920.
INDEX TO AMERICAN BOTANICAL LITERATURE
1916-1922
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
America being used in the broadest sense.
Reviews, and papers that relate exclusively to foresty, agriculture, horti-
estes manufactured products of vegetable Orig or laboratory methods
f bacteriology.
An occasional ereption is made i in favor of some paper appearing in an Ameri-
can periodical w wholly to botany. Reprints are not mentioned
unless they differ from the original in some important particular. If users
of the Index will call =a ea: of the editor to errors or omissions, their
kindness will be appr
This Index is pees moran on cards, and furnished in this form to
subscribers at the rate of three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Rhoads, A. S. The pathology of Lupinus arboreus, with special
reference to the decay caused by two wound-parasites,
Collybia velutipes and Pleurotus ostreatus. Phytopathology
11: 389-404. pl. 18-zo. 16 F 1922.
Rigg, G. B. Identity of cetain yellow pigments in plants and
animals. Science II, 55: 101, 102. 27 Ja 1922.
Robbins, W. W. Mosaic diseases of sugar beets. Phytopathol-
ogy. 11: 349-365. f. 7-8. 27 Ja 1922.
Robertson, C. Flowers and insects, XXI. Data of anthecology.
Bot. Gaz. 73: 148-152. 15 F 1922.
Robertson, C. Flower seasons. Sci. Mo. 14: 201-203. F 1922.
Robertson, C. The sunflower and its insect visitors. Ecology
3: 17-21. (84 Mr 1092
Rock, J. F. Hunting the chaulmoogra tree. Nat. Geog. Mag.
41: 243-276. Mr 1922. [Illust.]
Taraktogenos Kurzit of Siam.
Rothgeb, B. E. Cultural experiments with grain sorghums in
the Texas panhandle. U.S. Dept. Agr. Bull. 976: 1-43.
f. I-11. 20 Ja 1922,
307
308 InDEX TO AMERICAN BoTANICAL LITERATURE
Rydberg, P.A. Phytogeographical notes on the Rocky Mountain
region, X. Grasslands and other open formations of the
Montane Zone of the Southern Rockies. Bull. Torrey
Club 48: 315-326. 28 F 1922.
Saccardo, P. A. Micetes Boreali-Americani. Giorn. Bot. Ital.
27: 75-88. 1920.
Sage, H. The royal palm (Oreodoxa regia). Am. For. 28: 85-
88. F 1922. [Illust.]
Sax, K., & Gowen, J. W. Productive and unproductive types
of apple trees. Jour. Hered. 12: 291-300. 20 F 1922.
[Illust.]
Schaffner, J. H. Additions to the catalog of Ohio vascular
plants for 1921. Ohio Jour. Sci. 22: 91-94. Ja 1922.
Schaffner, J. H. The classification of plants. KIT. Ohio Jour.
Sci. 22: 129-139. Mr 1922
Schaffner, J. H. Control of the sexual state in Arisaema tri-
phyllum and Arisaema Dracontium. Am. Jour. Bot.
72-78. 20 Mr 1922.
Schaffner, J. H. Progression of sexual evolution in the plant
kingdom. Ohio Jour. Sci. 22: 101-113. F 1922.
Schlechter, R., & Hoehne, F. C. Contribuicdes ao conheci-
mento das Orquidaceas do Brasil. Anex. Mem. Inst.
Butantan 1: 9-68. frontis.+pl. 1-14. Mr 1922.
Includes Fractiunguis and Pseudostelis, gen. nov., 38 new species and
2 new varietics.
Schneifer, C. Notes on American willows—XI. Jour. Arnold
Arbor. 2: 185-204. Ja 1922.
Setchell, W. A. Aboriginal tobaccos. Am. Anthropologist 23:
397-414. O-D i921.
Includes map.
Severin, H. H. P. Minimum incubation periods of causative
agent of curly leaf in beet leafhopper and Fels beet.
Phytopathology 11: 424-429. f. 1-4. 16 F 1
INDEX TO AMERICAN BOTANICAL LITERATURE 309
Shear, C. L. Frederik Georg Emil oe Phytopathology
12: 1-3. With portrait. 10 Mr 1922
Shull, G. H. Three new mutations in Oenothera Lamarckiana.
Jour. Hered. 12: 354-363. f. 8-16. O 1921,
Sinnott, E. W., & Blakeslee, A. F. Structural changes asso-
ciated with factor mutations and with chromosome, muta-
tions in Datura. Proc. Nat. Acad. Sci. 8:17-19. 15 F 1922.
Sismey, E.D. A contribution to the algal flora of the Ojanagan
[British Columbia]. Canad. Field Nat. 35: 112-114. 3
F 1922.
Small, J. K. Historic trails, by land and by water. Jour. New
York Bot. Gard. 22: 193-222. pl. 263-266. D 1921.
Record of exploration in Florida in Dec. 1919.
Small, J. K. Wild pumpkins. Jour. New York Bot. Gard.
23: 19-23. 1922.
Smiley, F. J., & others. Weeds of California and methods of
control. Monthly Bull. Dept. Agr. of California Mo. Bull.
11: 1-22, 1-360. f. 1-738. F-—Mr 10922.
Spaulding, P. Investigations of the white pine blister-rust.
U. S. Dept. Agr. Bull. 957: 1-100. f. 1-13. 6 F 1922.
Stephenson, H. T. Trees in winter. Am. For. 28: 79-84. F
1922. [Illust.]
Sterrett, W. D. A new oak from the Gulf States. Jour. Elisha
Mitchell Sci. Soc. 37: 178, 179. Mr 1922
Quercus ashet sp. nov.
Stout, A. B. Cyclic manifestation of sterility in Brassica peki-
nensis and B. chinensis. Bot. Gaz. 73: 110-132. f. I-7. 15
F 1922
Sutherland, J. American trees in Scotland. Jour. For. 20: 71-
74. Ja 1922
Sydow, H. Novae fungorum species—XVII. Ann. Mycol. 19:
304-309. 1921.
des new American species in Cryotosphaeria (1), Gibbera (1), and
Phaeocibosia (1).
410 InpeEx to American BoranicaL LitERATURE
Taylor, W. R. The embryogeny of Cystanthus parviflorus
Baker. Am. Jour. Bot. 8: 502-506. pl. 25,26. 15 F 1922.
Trelease, W. The peltate peperomias of North America.
Bot. Gaz. 73: 133-147. pl. 1-4: 15 F 1922.
Includes 10 new species from Mexico and Central America.
True, R. H. The significance of calcium for higher green plants.
Science IT. 55: 1-6. 6 Ja 1922.
Uphof, J. C. T. Ecological relations of plants in southeastern
Missouri. Am. Jour. Bot. 9: 1-17. pl. 1, 2+ f. 1-3. 21 F
1922.
Vaupel, F. Echinocactus Williamsit Lem. Monatsschr. Kak-
teenk. 31: 186. D 1921. [Illust.]
Vaupel, F. Fruchtender Cercus Strausit (H.) Vpl. Monatsschr.
Kakteenk. 32: 8. Ja 1922. [Illust.]
Vinall, H. N., & Cron, A. B. Improvement of sorghums by
hybridization. Jour. Hered. 12: 435-443. frontispiece+
jf. FO. D926.
Wann, F. B., & Muenscher, W. C. A preliminary list of the
Myxomycetes of the Cayuga Lake basin. Mycologia 14:
38-41. 6 Mr 1922.
Warburg, O. Die Pflanzenwelt. 3: 1-552. pl. 1-27. + f. 1-278.
Leipzig. 1922.
Weatherby, C. A. The climbing fern in the vicinity of Hartford.
Am. Fern Jour. 11: 109-113. pl. 2. 31 Mr 1922.
Webb, R. W. Studies in the physiology of the fungi. XV.
Germination of the spores of certain fungi in relation to
hydrogen ion concentration. Ann. Missouri Bot. Gard.
8: 283-341. f. 1-39. .S: 192%.
Weidman, R. H. Forest succession as a basis of the silvi-
culture of western yellow pine. Jour. For. 19: 877-885.
D 1921.
INDEX TO AMERICAN BOTANICAL LITERATURE 311
Weingart, W. Cereus geometrizans Mast. und Verwandte.
Monatsschr. Kakteenk. 32: 17-20. F 1922; 35-38. Mr
1922.
Weingart, W. Cereus princeps Hort. Wiesb. Monatsschr. Kak-
teenk. 32:20. F 1922. [Illust.]
From Guatemala.
Weir, J. R. Cenangium piniphilum n. sp., an undescribed
canker-forming fungus on Pinus ponderosa and P. contorta.
Phytopathology 11: 294-296. pl. 13 + f.1, 2. 15 N 1921.
Weston, W. H. A note relative to the reappearance of the
sugar cane downy mildew in the Philippines. Phytopath-
ology 11: 371-375. 27 Ja 1922.
Wherry E. T. Soil reaction in relation to plant growth. Am.
Jour. Pharm. 94: 110-114. 922.
Reported by Dr. H. Leffmann,
Wiggans, R. G. A classification of the cultivated varieties of
barley. Cornell Univ. Agr. Exp. Sta. 46: 369-456. pl. 34-
38. +f. t-72.. S 1921.
Williams, F. N. Critical notes on some species of Cerastium.
Jour. Bot. 59: 324-329. N 1921; 59: 349-353. D 1921.
Includes C. glutinosum glabratum, var nov., from Colombia.
Williams, R. S. Mosses from British Guiana and Dominica,
Lesser Antilles, collected by Miss E. F. Noel in 1914.
Bryologist 24: 65, 66. pl. 4. 15 F 1922.
Macromitrium trinitense sp. nov.
Willis, J.C. Endemic genera of pate in their relation to others.
Ann. Bot. 35: 493-512. O 19
Wilson, E. H. The cherries of Japan. Publ. Arnold Arb.
7. 1916.
Wilson, E.H. The conifers and taxads of Japan. Publ. Arnold
Arb. 8. 1916.
Wilson, E. H. Notes from Australasia. II. The — Zealand
forests. Jour. Arnold Arbor. 2: 232-236. Ja 19
Wilson, T. R. C. The effect of spiral grain on the strength of
wood. Jour. For. 19: 740-747. f. 1-4. N 1921.
312 INDEX TO AMERICAN BOTANICAL LITERATURE
Winston, J. R. Commercial control of citrus scab. U.S. Leer
Agr. 215; 3-8. 15 F 1922
Wolf, F. A., & Shunk, I.V. Tolerance to acids of certain bac-
terical plant pathogenes. Phytopathology 11: 244-250.
21 O-1621.
Woodcock, E. F., & Zeeuw, R.de. The anatomy of the haustor-
ial roots of Comandra. Michigan Acad. Sci. Rep. 22: 189-
192. pl. 15. Au 1g2t.
Wright, W. G. Research work of the Dominion Forest Service.
Jour. For. 20: 62-66. Ja 1922.
Young, H. C., & Bennett,C. W. Studies in parasitism I.- Toxic
substances produced by fungi. Michigan Acad. Sci. Rep.
22: 205-208. Au 1921.
Yuncker, T.G. A list of Indiana mosses. Proc. Indiana Acad.
Sci. 1920: 231-242. 1921.
Yuncker, T. G. A species of Cuscuta not hitherto reported
from Indiana. Proc. Indiana Acad. Sci. 1920: 229. 1921.
Soret erie Neue Flechten—IX. Myc. 19: 224-242. 1921.
udes new species in Opegrapha (2), Graphis (1), Graphina (3), En-
jeteates (1).
Zahn, K. H. Compositae—Hieracium. Pflanzenreich IV.
280. 1-864. 180 Ig2t.
Zavitz, E. J. Reforestation in Ontario. Jour. For. 20: 18-24.
Ja 1922.
Zeller, S. M., & Owens, C. E. European canker on the Pacific
slope. Phytopathology 11: 464-468. f. 1-3. 25 F 1922.
Zeman, V. Los hongos de la “Phalaris bulbosa."’ Revista
Facult. Agron. 14: 179-184. 1921.
Zeman, V. Bacteriosis del bananero (una nueva enfermedad).
Revista Facult. Agron. Univ. Nac. La Plata 14: 17-30. 1921.
[Lllust.}
Zundel. G. L. The effects of treatment for bunt on the ger-
mination of wheat. Phytopathology 11: 469-484. f. 1, 2
21 Mr 1922.
Vol. 49 3 a No. II
BULLETIN
OF THE
TORREY BOTANICAL CLUB
NOVEMBER, 1922
Effect of external and internal factors on the germination
| fungous spores
Ws. L. Doran
(WITH TWO TEXT FIGURES)
The greater part of the work ‘which has been done on the
germination of fungous spores has been incidental to studies of
the life histories of fungi. References to spore germination are
in. most cases isolated and few attempts have been made: to
draw any general conclusions from the results obtained. Since
all fungous diseases must begin with infection and since the
germination of the spore is the forerunner of infection, further
knowledge of the conditions which favor or inhibit spore germina-
tion is of great practical importance. In this paper are des-
cribed the results of the writer’s study of the effect of various
factors on the germination of fungous spores.
It is becoming increasingly common for field studies of the
efficiency of fungicides to be preceded by laboratory tests of
the toxicity of fungicides to fungi. If fungous spores are to be
germinated both in the presence and absence of fungicides, it
is important that all other conditions be identical and at or near
the optimum. It is therefore necessary that optimum conditions
for the germination of the spores be known before the laboratory
tests of fungicides are made.
Frequent references are made in the literature to the effect of
prevailing climatic conditions on the outbreak or severity of
Reine aa ee —The author gratefully acknowledges the assistance
and encouragement received from Dr. O. R. Butler in this wor
[The aa sgh October (49: 283~312. pl. 12-17) was seittand October
11, 1922.]
313
3i4- Doran: ON THE GERMINATION OF FUNGOUS SPORES - *
epiphytotics of fungous origin. Field experiments and. observa-
tions on the effect of weather on diseases of plants are affected
by a very complex set of factors. Conclusions cannot safely
be drawn until the various factors have-been separately studied
and such study is more easily effected in the laboratory than
in the field.
Factors influencing the germination of fungous spores may be
divided into two groups, internal and external. The internal
factors are maturity of the spore, longevity of the spore, anima-
tion, and a poorly understood factor which may be called vitality
of the spore. The external factors are oxygen, presence of
moisture either in the form of water vapor or water of conden-
sation (rain or dew), temperature, light, nutrient substances,
toxic substances, and duration of the time period during which
one or more than one of the factors acting jointly exert their
influence on the spore of the fungus.
EXPERIMENTAI METHODS
The germination of the following fungous spores was studied:
conidia of Venturia inaequalis (Cke.) Wint., conidia of Sclero-
tinia fructigena (Pers.) Schrt., spores of Alternaria Solani (E.
M.) Jones & Grout., spores of Botrytis cinerea Pers., spores of
‘Rhizofus nigricans Ehr., aeciospores and urediniospores of
Cronartium ribicola F. de Wal., aeciosopres of Gymnosporangium
clavipes C. & P., and biiesiictds of Puccinia Malvacearum Mont.
The spores of the fungi were obtained fresh from the living host
in all cases except Alternaria Solani. Spores of parasitic fungi
to be used for germination tests were found to be more depend-
‘able when obtained from this source than when obtained from
fungi growing on artificial media. A spore obtained from the
living host is presumably possessed of its full natural vigor.
A spore obtained from a fungus growing on artificial media may
have been subjected to a debilitating influence, or may, on the
other hand, have become unnaturally resistant to adverse con-
ditions. In this connection it should be noted that Young and
Cooper (1) found that when Glomerella rufomaculans was grown
on agar eight to fifteen days, the Spores were more resistant
‘to the toxic action of fungicides than when. the fungus had been
grown on agar only four. to eight days. If fresh spores from the
living host are not obtainable and spores from artificial culture
must be used, it is the experience of the writer that all spores
DoRAN: ON THE GERMINATION OF FUNGOUS SPORES 315
the germination of which is to be studied should be of the same
age. The germination of spores from a second culture transferred
from the first should not be compared with the germination of
spores from the first culture.
Distilled water was the medium in which the spores were
placed to germinate. As is subsequently described more in
detail, it was soon evident that the distilled water contained an
insufficient amount of air for spore germination. Consequently,
the distilled water was in all cases aerated before being used.
The aeration was accomplished by drawing air through the
water for about thirty minutes by means of an aspirator and an
Allihn gas washing bottle.
On culture plate benches in moist chambers were placed
glass slides and on them drops of water either containing spores
in suspension or the spores were shaken off into the drops. It
is important that all drops of water be approximately equal in
size especially when the toxic action of fungicides is to be
tested. Clark (2) studied spore fermination in hanging cultures.
Duggar (3) used that method also and a modification of it,
placing the cells in small Petri dishes. Mains (4) germinated
fungous spores in hanging drops on the cover of a Petri dish.
When the cardinal temperatures for the germination of the
spores being studied was unknown, the first step consisted in
the determination of the optimum temperature for germination,
and in all later tests the spores were germinated at their optimum
temperature. Constant temperatures were secured by the use
of a DeKhotinsky electric oven and a Hearson incubator.
The next point determined was the minimum length of time
in which all the viable spores of each fungus will germi ate
at the optimum temperature. At least six hours more than this
minimum time period was allowed to elapse before removing
the slides from the moist chambers aid determining the relative
number of spores germinated. If slightly less than sufficient
time is allowed a narrower range of optimum temperatures
will be found than if more time is allowed, resulting, when the
results are plotted, in a temperature-germination curve more
acute than flat at itssummit. If too long a time passes before
the germinating spores are counted, the germ tubes will have
grown and branched so as to make counting difficult and un-
certain. All data are based on relative number of spores germi-
nating.
316 DoRAN: ON THE GERMINATION OF FUNGOUS SPORES
The spores were counted by the use of a micrometer disc
ruled in squares and the help of a tallying register. The micro-
meter disc ruled in squares is placed in the ocular, the slide
bearing the drops of water containing spores is placed on the stage
of the microscope and moved by means of a mechanical stage
until the outer line on the micrometer disc appears as a tangent
to the circumference of the drop. The germinating spores
appearing between the first two lines of the micrometer disc
are counted, and then the ungerminated spores in this area are
counted. When one field as limited by the lines on the micro-
meter disc is counted, the slide is moved enough to bring a new
field into view for counting. This is continued until a total of
fifteen hundred to two thousand spores, both germinated and
ungerminated, have been counted. This necessitates the ex-
amination of several or many drops. From the count obtained,
the relative number of spores germinating is figured, germina-
tion elsewhere being raised proportionately. This accuracy of
counting is most necessary when the relation between tempera-
ture and spore germination is being determined. It is not as
necessary in the case of toxicity tests with fungicides except
when the differences in toxicity are small.
RELATION OF THE VIABILITY OF THE SPORE TO ITS AGE
Maturity—Until the spore has attained a certain age or
degree of maturity, it cannot germinate. Even before a spore is
really mature, it can germinate if other conditions are near
enough to the optimum. For example, the range between the
minimum and maximum temperature is considerably greater
for the germination of a mature urediniospore of Puccinia
Antirrhint than for the germination of an immature uredinio-
spore of this fungus, collected the first day it breaks through the
host surface. The immature spore may be prevented from
germinating by the action of fungicides not of sufficient con-
centration to prevent the germination of the mature spore.
A solution containing 0.257 per cent copper sulphate prevents
the germination of mature spores of Uromyces caryophyllinus..
A solution containing only half as much copper sulphate prevents
the germination of immature spores of this fungus. It should
be understood that mature and immature spores may not
differ morphologically. Spores here referred to as immature are -
so called because when collected young they did not germinate,
DoRAN: ON THE GERMINATION OF FUNGOUS SPORES 317
although older spores from the same fungus did germinate. All
conclusions as to germination should be based upon the behavior
of mature spores.
It is possible by exercising care in obtaining spores to separate
the mature from the immature to some extent at least. When a
fungus has matured its spores, they are usually freed in such
a way that they may be disseminated by the movement of air
or water. So far as possible, the natural method should be
duplicated in obtaining spores for experimental purposes. When
a scab lesion on the fruit or leaf of the apple is washed, it is
probable that only mature conidia and perhaps a few dead
conidia of Venturia inaequalis are detached. But when a similar
lesion on the fruit or leaf of the apple is brushed or scraped, it
is probable that many immature conidia are also detached. In
the case of all except the fungi having very short-lived spores
the immature spores on a fresh lesion greatly outnumbered the
dead spores. When the writer obtained conidia of Venturia
tnaequalis by brushing apple scab lesions the relative number of
conidia which were capable of germinating was only twenty-five
as compared with the relative number of one hundred germinat-
ing when the conidia were obtained by washing lesions with a
stream of water from a pipette. When fungous spores are to be
obtained for germination studies they should be freed from the ©
host as gently as possible and in as nearly as possible the same
way in which they would be removed in nature. In this connec-
tion it is interesting to note that Jones (5) found that the
ascospores of Pseudopeziza Trifolii and P. Medicaginis germinate
readily when they are discharged naturally but not when they
are crushed out of the ascus.
Some attention has been given to the later maturing of spores
which were detached from the fungus while still immature.
Schaffnit (6) is of the opinion that unless fungous spores are
internally mature before they are detached from the fungus,
they never gain power to germinate. But the results of Melhus
and Durrell (7) indicate that some urediniospores can mature
after being detached from the furigus.
With the exception of the spores of Botrytis cinerea, all the
fungous spores studied by the writer were —— killed if
detached when immature.
It is possible in some cases to distingiush between mature and
immature spores by their relative position in the fruiting parts
318 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
of the fungus. The only spores of Sphaerotheca mors-uvae which
Foreman (8) induced to germinate were the large spores found
at the tips of the chains of spores. Weimer (9) observed that
the teliospores of Gymnosporangium Juniperi-virginianae which
are situated at the apex of the spore horn germinate best and he
attributed this to their maturing earlier.
As the season advanced in the autumn the writer found it
increasingly difficult to secure good germination of the conidia
of Venturia inaequalis even when conidia were obtained from
fresh lesions. This is not attributed to the presence of a greater
number of spores too old to germinate, for they would probably
have been washed away earlier. It is believed that a larger
number of immature conidia of this fungus were present in the
autumn. It is not unlikely that the process of maturing of the
spore is slower when the temperature is lower
Very little work has been done on the relation of the viability
of fungous spores to the conditions under which the host plant
grew and in which the fungus developed. If certain conditions
may retard the maturing of the spores, it is quite possible that
other conditions during their development may entirely prevent
their ever germinating. On the other hand, certain conditions
during development of the spore may result in an extremely
- vigorous spore. Spores may be mistakenly designated as mature
and immature, on the basis of their behavior when placed under
optimum conditions for germination, when they should properly
be designated as spores of the same age, some vigorous and others
without vigor.
The inability to germinate any fungous spore under all possible
combinations of conditions may be temporarily explained by the
supposition that the spores are either dead, non-fu ictioning or
in a resting condition. There are perhaps many resting spores
not now recognized as such. Reed and Crabill (10) have ad-
vanced the theory that a rest period is necessary for the aecio-
spores of Gymnosporangium Juniperi-virgimianae and that these
spores do not germinate until the spring following their dispersal.
Longevity.—For all spores ‘there is a maximum time limit,
variable with the environmental conditions, beyond which the
spore cannot germinate. The spore may be alive and too old to
germinate or it may be dead. A live spore too old to germinate
under the range of conditions within which it formerly could
germinate is, according to the observation of the writer, able
DORAN: ON THE GERMINATION OF FUNGOUS SPORES 319
to germinate when all conditions approach the optimum. There
comes a time when even this is not stimulative and the spore is
then dead so far as we can tell. Neglect of the age factor may
render any study of spore germination undependable.
In studying the relation of the age of spores to their viability,
points to be considered are the date of collection, the conditions
in which the spores are placed after collecting, and the length
of time spores remain viable after their collection. Aeciospores
of Cronartium ribicola were found to germinate with diminishing
vigor as the season advanced. The aeciospores of this fungus
were collected on May 25th, June 4th and June 12th from the
same sorus. On the first date, the relative number of spores
germinating was one hundred, on the second date forty-nine
and on the third date twenty-nine. Observations with this.
and other fungi indicate that although there is at first a sharp
falling-off in viability, it is retained for a comparatively long
time and only gradually lost. In any spore germination studies
which involve comparisons, spores should be used which are
known to be of the same age.
Among the factors bearing upon the retention of viability
after the collection of spores are moisture, temperature, and
freedom from or connection with the parent. Detachment
from the parent was not found to have any effect on the length
of life of the aeciospores of Cronartium ribicola, viability de-
creasing the same in both cases. When collected, the relative
number of these spores germinating was one hundred, after
having been collected thirty days the germination fell to seventy-
three, after fifty days to five, and after a days no spores
germinated.
Freshly collected aeciospores of apenas ribicola were
stored indoors at temperatures of 7° C., 15° C., ae These
various temperatures had no effect upoa the peer of the
spores. But aeciospores stored in moist air retained their
: viability longer than those stored in dryer air. At least in the
case of this fungus moisture is of more importance than tempera-
ture in its effect on the length of life of the spore. This effect of
moisture is to be expected; for all spores lose water in dry and
moist air, and if other conditions are equal they should live
longer in moist air than ia dry air. These results agree with
those of Anderson (11) who found that in a humid atmosphere
320 DoRAN: ON THE GERMINATION OF FUNGOUS SPORES
the spores of Cylindrocladium scoparium live for several weeks
but that in a dry atmosphere they die in fifteen days.
Under no conditions did aeciospores of Cronartiwm ribicola
germinate when more than eight weeks old. few cases are
recorded in the literature of aeciospores of this fungus living
longer, but it is probable that in such cases germination is reduced
to a trace and that even that small germination occurs only
when all environmental conditions are very close to the optimum
for a time.
The conidia of Venturia inaequalis produced on the leaves
are somewhat shorter lived than those produced on the fruit.
According to Aderhold (12) the conidia of this fungus retain
their viability not more than eight weeks. The writer found
none of the conidia of this fungus to live more than six weeks on
the fruit and not more than four weeks on the leaf. With the
aging of the leaves in the fall, the life of the conidia becomes even
shorter. Difficulty was experienced in germinating conidia
from fruit in storage. The scab lesions, however, were con-
taminated with Cephalothecium roseum, the presence Of the
conidia of which in the drops of water may bave been partly
responsible for the failure of the conidia of Venturia inaequalis
to germinate. As will be described later, the presence in a drop
of water of the spores of several fungi prevents or retards the
germination of all the spores as compared with germination in
a drop of water containing only the spores of one fungus.
One of the factors bearing upon the retention of viability by
spores after their collection is their freedom from or connection
with the fungus or each other. Anderson and Rankin (13)
found that ascospores of Entothia parasitica which remained in
the perithecia in the bark germinated after being stored dry for
one year. But when ascospores were removed and separated
for only five months they lost the power to germinate. According
to Burrill (14) the spores of Glomerella rufomaculans remain
viable for a long time when their soluble protective coating in
which they are imbedded is present. But if it is washed away,
the spores soon lose the power of germination. Anderson and
Rankin (13) found that if the pycnospores of Entothia parasitica
are stored in a dry place they live at least one year. But if the
pycnospore horns are placed in water and the water allowed to |
evaporate so as to leave the spores separated, they do not retain
their viability more than one month.
DoRAN: ON THE GERMINATION OF FUNGOUS SPORES 321
It is possible that in most cases the mucilaginous covering
in which many spores are imbedded serves to prolong the life
of the spore. For this reason, whea studying the germination
of such spores, it is advisable that only those still adhering to-
gether in their protective coatings be used.
Exposure to light shortens the life of some spores, as shown
by the results of Hoerner (15) with the urediniospores of Puccinia
coronata.
Some attention has beea given to the longevity of the spores
of the Uredinales. The sporidia are the shortest-lived spores of
this group of fungi. According to Reed and Craybill (10) the
sporidia of Gymnosporangium Juniperi-virginianae do not live
more than six days in dry air. Spaulding (16) reports that the
sporidia of Cronartium ribicela live less than ten minutes at
room temperature with the humidity at ninety. Duff (17)
found that two weeks after the collection of the urediniospores of
Cronartium ribicola their germination was so reduced as to be-
come practically negligible. The life of a teliospore measured
from the time of its formation to its death may be long, but the
life of a teliospore if measured from the close of its normal resting
period until its death is not long; it is apparently as short in
some cases as the life of many non-resting spores. Melhus,
Durrell, and Kirby (18) report that the teliospores of Puccinia
graminis, which mature in April, cannot germinate after the end of
June following.
The experience of the writer is to the effect that in most cases
aeciospores are longer lived than urediniospores. The average
life of urediniospores is between thirty and sixty days. The
average life of the aeciospore is about 50 per cent greater. The
several spore forms of the Uredinales behave alike in this, that
as they grow older their range between maximum and minimum
conditions for germination becomes narrower.
EXTERNAL FACTORS
Temperature relation—One of the most important external
factors bearing upon the germination of fungous spores is the
temperature relation. In order to draw from the scattered data
in the literature some more general conclusions than we now have
as to the cardinal temperatures for the germination of fungous
spores, TABLE I has been prepared. In it are given the cardinal
322 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
temperatures and the range of temperature for germination of
thirty fungous spores.
If the optimum temperature for the germination of the spores
of Plasmodiophora Brassicae, which Chupp (19) found to be
27°-30° C., is typical for the Myxomycetes, then the temperature
requirements for the germination of the spores of the slime moulds
must be higher than those of fungi.
A study of TABLE I shows for the germination of the spores of
several groups of fungi the following average cardinal temperatures
and average range between minimum and maximum temperatures.
For the germination of the Phycomycetes the average cardinal
temperatures are 1.2° C., 18° C., and 26.1° C., and the range is
25.0 degrees Centigrade. For the germination of the aeciospores
of the Uredinales, the average cardinal temperatures are 6.5
C., 13.0° C. and 22.0° C. and the average range is 15.5 degrees
Centigrade. The average cardinal temperatures for the ger-
mination of the urediniospores are 5° C., 16.2° C., and 28.9° C.,
and the range through which these spores can germinate is
23.9 degrees Centigrade. In the case of teliospores the cardinal
temperatures for germination are 7.4° C, 19.9° C., and 28.1° C.,
and the range for germination is 20.7 degrees Centigrade.
Temperature limits are narrower for the production of sporidia
by germinating teliospores than for the production of germ tubes
only. This is illustrated by the results obtained by the writer
with the teliospores of Puccinia Malvacearum. At temperatures
as low as 5° C. promycelia are produced but no sporidia. Above
23° C. promycelia are produced but sporidia are rare.
These results agree with those of Dietel (29) who found that
above 23° C. the teliospores of Puccinia graminis produce no
sporidia but only germ tubes. Reed and Craybill (10) noticed
that the teliospores of Gymnosporangium Arai sping entering
do not germinate when the temperature rises to 24° C. But
above that temperature they sometimes develop promycelia.
When the Fungi Imperfecti and the one representative of the
imperfect stage of the Ascomycetes named in Taste I are con-
sidered collectively, it is seen that for the germination of these
spores the cardinal temperatures are 7.3° C., 23.1° C., and
39.6° C. Between the minimum and maximum temperature
limits for germination ‘here is an average range of 32. 3 degrees
Centigrade.
DoRAN: ON THE GERMINATION OF FUNGOUS SPORES
TABLE I
323
Cardinal temperatures and range of temperature for the germination
of the spores of representative fun
Author-| | Cardinal Faeaenmina
ity* | oe | Min. Opt: Max. (Range
18 Plasmodiophora Brassicae...... | — |27°-30°C —_— —_—
20 mopora viticola, conidia . — |25°-30°C — —
24° 1P. essa comidia Hoos ret cS — (25°-35°C — —
22 =| Cystopus candidus, conidia... .. a Os 10°C, [25. © an
23 (Phytophthora infestans, conidia | 2°-3° C. |12°-13°C. a4 225° G.23
24 =| Peronospora parasitica, conidia a 8°-12° C.| 29° C.
25 gti lem clavipes, aeci- |
Pita t OS AS ee is Se ta CG. 25°C. 17
25 ticsrkida ribicola, aeciospores | 5° C. mC. 19°C. 14
4 (Puccinia Phlei-pratensis, ure-
MEDIOBOOVER 6 pe 18°C. 30° C —_
4 |P. coronata, urediniospores... .. —- 18°C. 300 —
26 | P. coronata, urediniospores Ye — 130°C 23
7 |P. coronata, urediniospores..... sak 17°-22° C.'35° C. 34
26 =| P. rubtgo-vera, urediniospores . .| 2° C. — |31°C. 29
26 =| P. graminis, urediniospores..... °C, — 31°C. 20
27 =| P. dispersa, urediniospores. . .. . 10°-12° C.'18°-20° C.|25°-27° C.| 17
25 | P. Antirrhini, urediniospores...| 5° C. ro C. a Bee 5
28 |P. Sorghi, urediniospores....... ec. 14°C. 25°C ae
25 Saertegs ribicola, uredinio-
Rs) ae ae va ib ta: 25°C +7
25 Uromyces caryophyllinus, ure-
‘ hes ee OS ra". 29°C 25
2 oe Trifolit uredinio-
_- Spores eerste ee — 116°C. 44°C —
10 Cymbdésorenieee Juni peri-vir-
inianae, teliospores......... 13 ign SS et: 29°C. 18
9 G. Juniperi-virginianae, telio-
BOOED 6 oh ace ierace's + Be 23°-24° C.'29° C. 22
29 (Puccinia graminis, oribegggges et. oa’ C; as". 14
18 P. graminis, teliospores........ tobe 20° C. a5-h. 20
29 _|P. Larici, teliospores......--.-|. ot. — —
25 | P. Malvacearum, ~en meg asco (eC 6 C. 27
18 |P. graminis, basidiospores. ..... — {15*20°C. ~— ~—
30 easel mgenriam Phicas Pi 22°-27° C — _~
31 | Melanconium sp...........+++- — |23°-27°C =. as
32 Phyllosticte bashes cea — es oo an or
we Le ARPMNE,. «ise: °C. — '47°C. : 29
84 |Alternaria Solani...........-- 1°-3° C. |26°-28° C. 37°-45° C.| 44
35 re ge — _ |24°-28°C. 34°C. “Gis
*
Numbers refer to aa literature nose
324 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
For these several groups of fungi the minimum temperatures
for spore germination occur between 1° and 7.4° C. The opti-
mum temperatures are all between 13° and 23.1° C. The maxi-
mum temperatures for the germination of the spores occur
between 22° and 39.6° C. The Phycomycetes have the lowest
minimum temperatures, followed in order by the urediniospores,
the aeciospores and the teliospores of the Uredinales. Teliospores
and the sporesof the Fungi Imperfecti have the highest, and about
the same minimum temperature for germination.
Of these five groups of fungous spores, aeciospores have the
lowest optimum temperature for germination followed in tura
13 616.2 18 a i 23.1
100 x
/\, 7 \ s aN
99) PER CENT Bat Aes PHYCOMYCETES
st : : AECIOSPORES —---——-—
oy RRR UREDINIOSPORES +++
70 ELIOSPORES —*—x—*—
FUNGI IMPERFECTI-~~~-~
60
50 .
40
™ ae
20 ~
* \ x
; fs ‘ <
e / f oa tet tae \.
© A23Z45 6574 is 162 18 20-22-24 26 wiss 396
Fic. 1. Curves showing the rin temperatures for the germination
of the spores of representative fung
by urediniospores, conidia of Phycomycetes, teliospores, and
then by the spores of the Fungi Imperfecti, which seem to have
the highest optimum temperature for germination.
Aeciospores have the lowest maximum temperature for
germination, and the spores of the imperfect fungi have the
highest. About midway between these two come urediniospores.
Their maximum temperature for germination is higher than that
of the conidia of the Phycomycetes.
The spores of the Fungi Imperfecti can, it would appear,
germinate through the greatest range of temperature. As com-
pared with this, the range of the Phycomycetes is 77 per cent
DorAN: ON THE GERMINATION OF FUNGOUS SPORES 325
as great, that of the urediniospores is 74 per cent as great, that
of the teliospores is 64 per cent as great, and the range of the
aeciospores is only 48 per cent.
IG. I shows the ranges and the cardinal temperatures for
- these groups, and their -relative positions.
The results obtained by the writer and others indicate that any
departure from optimal conditions tends to narrow the tempera-
ture limits within which fungous spores will germinate. The
literature contains a very few references to the relation which
PER CENT
GERMINATION
DEGREES CENTIGRADE
er Soke 6.7.8 940 12 13 14 15 16 1718 19 20 21 22 23 24 25 26 27 2629 30 Ht 32,
Fic. 2. Curve showing singel limits for the germination of the
conidia of Venturia inae
exists between the temperatures required for the process of spore
germination and the other environmental conditions prevailing
during that period. Gardner (30) found that the minimum
temperature for the germination of the spores of Colletotrichum
lagenarium is 7° C. when the spores are in exposed drops of water,
but it is 14° C. when the spores are in hanging drops of water.
When these spores are in exposed drops of prune decoction, the
minimum temperature for their germination is 4° C. It would
appear that the presence of nutrient substance and the availa-
bility of sufficient we hci both tend to lower the minimum
temperature.
- The work of the writer included the determination of the
cardinal temperatures for the germination of the conidia of
Venturia inaequalis, the spores of Botrytis cinerea, and those of
326 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
Rhizopus nigricans. Cardinal temperatures for the germination
of the other spores used had been previously determined (Doran,
The results obtained with the conidia of Venturia inaequalis are
shown in TABLE II and by the curve in Fic. 2. For the germina-
tion of the conidia of Venturia inaequalis, the minimum tem-
. perature is 3° C., the optimum temperature is 14°-15° C., and
the maximum temperature is 31° C. The conidia can germinate
through a range of twenty-eight degrees.
This is an unusually long range. Since the causal fungus of
apple scab is not especially exacting as regards temperature for
the germination of the conidia the indications are that the
distribution or severity of this disease, following the primary
infection by the ascospores is dependent more upon rainfall
than upon temperature. The conidia of Venturia inaequalis are
disseminated most freely during June, July and August. As
shown in Fic. 2, warm nights do not prevent their germination.
A study of the temperature-germination curve in F1G. 2 shows
that the falling off in germination of the conidia is more rapid
from the optimum to the minimum temperature than from the
optimum to the maximum temperature. Over 40 per cent of the
number of spores germinating at the optimum (14°-15° C.)
germinate at 9° C. and 24° C., respectively.
TABLE II
Effect of temperature on the germination of the conidia of
Ventur ria inaequalis
Relative germination at different temperatures measured in degrees Centi-
grade
2°1.3°.16 |S. {to°. (12°, [4°jz5°1z8° [20° [24° [28° 130° -i31° 132°
2 42
© | 4 | 18 | 60 | 80 |r100}100/ 68 | 58 | 46 | 24 | 10
ae
ris
oo Oo ©
N
we
on
wt
8
8
“I
°
on
on
w
[one]
to
°
-_
Ny
So oN &
oo °o 9
Mean} o 0.75} 3 |21.5)/56 2|76.5/100'100|77 2
56.5/41.5|22.2| 11 21.0
2
In these studies of the relation of spore germination to tem-
perature, as to all other external conditions, the relative number
of spores germinating was taken as an indicator of the reaction
DoRAN: ON THE GERMINATION OF FUNGOUS SPORES 327
of temperature. Johnson (26) and Melhus and Durrell (7)
preferred to use the growth or length of the germ tubes as an
indicator of the temperature reaction. To test the relative
merits of these indicators, both were used by the writer in the
case of germinating conidia of Venturia inaequalis. When the
optimum temperature for germination was based upon length
of germ tube rather than upon number of spores germinating
the optimum was less sharply defined. The writer believes that
to consider length of germ tube rather than number of spores
germinating involves growth rather than germination and so
should be studied separately, for there is no reason to believe
that the optimum temperature for the growth of a fungus is the
same as the optimum temperatures for spore germination.
Melhus (22) found the optimum temperature for the germination
of the conidia of Phytophthora infestans to be 12°-13° C. Jones,
Giddings and Lutman (30) found that the optimum temperature
for the growth of this fungus is between 16° and 19° C. The
cardinal temperatures for the germination of the conidia of
Botrytis cinerea were found to be minimum 7° C. to 17° C., an
maximum 26° C. For the germination of the spores of Rhizopus
nigricans, the minimum temperature was found to be 10° C.
and the optimum 19° C. to 20° C. All later studies of the germ-
ination of Botrytis cinerea were made at 15° C. and of the germ-
ination of Rhizopus nigricans at 19° C.
Time relation—The length of time required for the process
of spore germination is in itself an important factor. If before
the elapse of the necessary time period the environmental
conditions depart from the optimum, the germination process is
impeded or stopped and the spore may even be killed. In all
laboratory studies of spore germination, if conditions in nature
are to be duplicated, no more time need be allowed for germina-
tion than that in which nearly optimal conditions are likely to
continue in nature. Probably most fungous spores begin and
complete germination between darkness and daybreak. One of
the principal natural protections of plants from pathogenic
fungi is the failure of environmental conditions to remain be-
tween minimum and maximum limits long enough for spore
germination. Infection does not occur if during the protrusion
and growth of the germ tube the necessary precipitated moisture
evaporates or the air falls below a certain degree of humidity
or the temperature passes beyond minimum or maximum limits.
328 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
According to the observation of the writer, extremes of tempera-
ture at this time, if not too long continued, have no more serious
effect than to temporarily stop the growth of the germ tube.
This is resumed when the temperature again approaches the
optimum. But when the humidity of the air falls, the results
to the spore are more serious. In the case of all spores worked —
with, and especially with conidia of Venturia inaequalis, it was
found that a germinating spore can live but a short time in a dry
environment, which to an ungerminated spore would be in no
way detrimental. A return to optimal conditions proved such
spores to be not merely inhibited but dead.
Wallace (37) considered the length of time apple trees remain
wet so as to allow spore germination an important factor in
determining whether or not a certain rain would permit infection
by Venturia inaequalis. He found that the ascospores of this
fungus can germinate in four hours but he believed that apple
trees must remain wet eight or ten hours to be abundantly in-
fected. The writer will here comment that the further the
temperature is from the optimum for germination, the longer must
the surface of any plant remain wet in order for the spores on
it to germinate and infect the host.
Duggar (3) found that after drying the germinating spores
of Botrytis would grow no more. But the germinating spores of
Aspergillus showed new growth after being dried several days.
Duggar’s results with the parasitic form mentioned are in agree-
ment with the results secured by the writer, who worked only
with parasitic forms.
he relation of environmental factors to the time required
for spore germination thas occasionally been noted. The amount
of moisture present may affect the length of time in which the
spore germinates. Taubenhaus (38) reports that the spores of
Glomerella rufomaculans from the sweet pea germinate in from
six to twenty-four hours, depending on whether there is much
or little moisture present in the atmosphere.
The writer has noticed that spores germinating in moist air
require a relatively longer time for the process than do the
same spores germinating in or on precipitated moistqre.
Anderson (11) observed that at 25° C., which is within optimal
limits for the germination of the spores of Cylindrocladium
scoparium, the spores of this fungus begin to germinate in two
to three hours, but at lower temperatures a longer time is neces-
DorAN: ON THE GERMINATION OF FUNGOUS SPORES 329
sary for their germination, five hours at 12° C. and twenty-four
hours at 8° C. Tisdale (39) found that at 4° C., which is their
minimum temperature for germination, the spores of Didymellina
wridis germinate in twenty-one hours, but at their optimum
temperature of 20° C. to 26° C. these spores germinate in two
and one half hours, and at their maximum temperature which
is 30° C. their germination requires eleven hours.
The presence of a nutrient solution may hasten the germina-
tion of some spores. Gardner (30) found that the spores of
Colletotrichum lagenarium germinate in five hours in agar but
require ten to twenty-four hours for their germination in dis-
tilled water at the same temperature.
It appears that immature spores and aged spores may germin-
ate more slowly than mature spores. Melhus, Durrell, and
Kirby (18) found that both before and after the teliospores of
Puccinia graminis have completed their rest period, the time
required for their germination is longer than that required just
as they end their rest period in the spring.
The results mentioned above indicate that all fungous spores
germinate most rapidly when all environmental conditions are
nearest to the optimum.
TABLE III shows the time necessary for the germination of
twenty-three representative fungous spores. After a study of
this table it is possible to draw certain conclusions as follows:
The average time required for the germination of fungous spores
is about twelve hours; for the germination (indirect) of the conidia
of the Phycomycetes it is four hours; for the ascospores of the
Ascomycetes it is thirteen hours; for the germination of the
chlamydospores of the Ustilaginales it is twenty-one hours; for
the germination of the aeciospores of the Uredinales it is ten
hours; for the germinations of the urediniospores of the Uredin-
ales it is five hours; for the germination of the teliospores of the
Uredinales it is three hours; and the average time required for the
germination of the spores of the Fungi Imperfecti is seventeen
hours.
The writer has found that the length of time necessary for the
germination of the fungous spores named in TABLE IV is: aecio-
spores of Cronartium ribicola twelve hours; urediniospores of
Cronartium ribicola five and one-half hours; for the production
of basidia by the teliospores of Puccinia Malvacearum two hours;
one hour more, a total of three hours are necessary for the
330 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
production of sporidia by the teliospores of this fungus; aecio-
spores of Gymnosporangium clavipes require four hours; conidia
of Venturia inaequalis about twenty-four hours; and conidia of
Sclerotinia fructigena five hours. These tests were all made at
TABLE III
Length of time required for the phe ie of the spores of
representative fung
| Hours necessa
Authority* Fungus \for spcediiieibet:
23 | Phytophthora teas COnIGie oe eee eee i
22 Cystopus candidus, conidiasc. i. 6... eres es 2-10
40 Plasmopara viticola, niet pe ae oe) ate es 3-12
37 Venturia inaequalis, ascospores................ 4
41 Guignardia Bidwellii, ascospores............... 36
13 Endothia parasitica, ascospores...............-. 6-12
39 Didymellina Iridis, sheen eintegemigea eS 18-36
SCOSPOLeS. 256 FS SEE Cees 6
5 Pseudopeziza Trifolii ‘asain See Pe re. aca 12
42 Diplocarpon Rosae, conidia. ........ 6.252. 05 ess 24
38 Glomerella rufomaculans, — fg et as ee 6-24
43 Tilletia foetens, chlamydospores................ 48
43 Ustilago Hordei, chia paidonpores A > aera gieseael 6.5
43 Ustilago Tritici, chlamydospores............... 14-15
25 Cronarti bicola, aectospores .<.. 5220025... 8-10
25 Puccinia Antirrhini, urediniospores............. 5-8
46 P. Phliet-pratensis, uri ‘sees ODORCS es pre as hoa, 3
35 P. Malvacearum, teliospores................... 3
9 ae Peaibet died telio-
Pr Orc brs OF eee Pee Pir ak Te Pees 3-4
30 Calettrich SCRA oi BS 10-24
45 MNF oi bok ik cemkn im ee pe ee E 16-24
32 Phllostct ANIME. oo ois coos ee es ees 16-30
33 BNE sb veo Gen ies ce ee 18-24
11 Cylindrocladium scoparium............02.0000% 2-3.5
46 Depronna GOIN: be a ee 18
sd Numbers refer to the literature cited.
the respective optimum temperatures for germination. Other
tests were made at temperatures approaching the minimum
and maximum temperatures for the germination of the spores of
these fungi. Under these conditions, in every case, a longer
time v 1 for g tion than at optimum temperatures.
Oxygen relation.—It was observed that spores in the interior
of a drop of distilled water never germinated as well as those on
DORAN: ON THE GERMINATION OF FUNGOUS SPORES 331
the surface. The conidia of Venturia inaequalis germinated
only when they were on or near the surface of the water. Telio-
spores of Puccinia Malvacearum did not produce sporidia when
deeply submerged in the drop. But when distilled water was
used, which had previously been aerated, equally good germina-
tion was secured whether or not the spores were submerged
in the drops.
DeBary (47) described the behavior of spores in water be-
tween a cover glass and a glass slide. In this case, the spores
TABLE IV
Time period necessary for the germination of representative
fungous spores — optimum conditions
Fungus and spores No. of hours|Spores germinating (relative numbers)
Cronartium ribicola, IO. °o
Aeciospores 12 100
Cronartium ribicola, 5-5 100
Urediniospores
Gymnosporangium clavipes ,| 4.0 100
Aeciospores :
Puccinia Malvacearum, 2.0 |Producing basidia too. Producing
Teliospores basidiospores o
3.0 |Producing basidiospores 100
Sclerotinia fructigena, 5.0 100
Conidia
Venturia inaequalis, 16.0 25
Conidia 27.0 100
near the periphery of the cover glass germinate better than those
near the center due‘to the relative amounts of air available.
Duggar (3) found that a reduced oxygen supply retards spore
germination. Blackman (48) noticed that if the germ tube of
Phragmidium violaceum does not grow through the water and
so reach the air it develops abnormally. Weimer (9) observed
that when the teliospores of Gymnosporangium Juniperi-
virginianae are covered with water they produce only long tubes
instead of the normal promycelia and basidiospores. Melhus
332 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
and Durrell (7) found that when the urediniospores of Puccinia
coronata are submerged only a small percentage of them germinate
as compared with the number germinating when they float on
a drop of water.
When spore germination tests were made in non-aerated
distilled water, the writer secured best results by sowing the
spores on the surface of the water rather than immersing them.
Some spores, however, will not float. According to Duggar (3)
the spores of many of the Phycomycetes and Hymenomycetes
usually sink. The specific gravity of the spores of several
Hymenomycetes determined by Buller (49) were between 1.02
and 1.21. Since some spores sink, all distilled water for germina-
- tion studies should be aerated.
At this point reference should be made to the injurious effect
of the presence of fungous spores on the germination of the spores
of other fungi. Although the injurious effect of this competition
tween spores might be attributed to the secretion of toxic
substances, there is little likelihood that this is the case. There
is an indication that it is due to there being insufficient oxygen
for all. The effect is much more marked in non-aerated than in
aerated distilled water and is more marked in the interior of a
rop of non-aerated distilled water than on its surface. All
spores of Uredinales which the writer has observed germinate
best when the drop of water in which they are contained is not
contaminated by the presence of other fungous spores. When
spores of Alternaria, Colletotrichum, or Cladosporium are present
in a drop of water with the teliospores of Puccinia Malvacearum,
the spores of the first three fungi may germinate but the telio-
spores of Puccinia Malvacearum do not germinate, although
check teliospores in uncontaminated drops germinate perfectly.
When in place of Puccinia Malvacearum the conidia of Venturia
inaequalis were used a similar result was obtained; they germin-
ated far better when no other spores were present with them in
the drop of water. When there is lack of sufficient oxygen for
all, only those spores which require the least will germinate.
It was also observed that when relatively few spores of one
species were present in a drop they germinated better than
when the drop was crowded. This, too, may be attributed to
insufficient oxygen. In all spore germination studies where
optimum conditions are desired; an effort should be made to
exclude from the culture drop the spores of all fungi except the
Doran: ON THE GERMINATION OF FUNGOUS SPORES 333.
one under consideration, and there should not be present an
excessive number of spores of that one fungus.
There are a few references which have come to the attention
of the writer on the deleterious effect of competition or crowding
on germinating spores. Edgerton (50) noticed that when more
than twelve or fifteen spores of Colletotrichum Lindemuthianum
are present in one cubic millimeter of water, they germinate
more poorly than when a smaller number of spores are present.
Taubenhaus (51) found it advisable to wash hollyhock leaves
in water to remove the spores of saprophytic fungi preparatory
to studying the germination of the teliospores of Puccinia
Malvacearum. He does not indicate in what manner the pres-
ence of the spores of other fungi interfere with the germination
tests.
Light relation.—Opinion in the literature is not unanimous
as to the effect of light on spore germination. According to
DeBary (52) and Farlow (53) light inhibits the germination
of the spores of the Oomycetes. Cuboni (54) concluded that
intense light interferes with the germination of the conidia of
Plasmopara viticola. Ward (55) found that the spores of the
brome rust germinate as readily in light as in darkness. Melhus
(22) observed no difference in the percentage of conidia of
‘ystopus canididus germinating, nor in the time required for
their germination, whether germination took place in light or
darkness. Melhus (23) found that light does not interfere
with the germination of the conidia of Phytophthora infestans
if the optimum temperature for their germination is not exceeded.
Duff (17) found that exposure to glass filtered sunlight is not
injurious to the urediniospores of Cronartium ribicola, providing
the temperature does not rise too high. He found that exposure
to the ultra-violet rays from an electric arc completely inhibited
spore germination. Lauritzen (56) concluded from his invest-
igations that light is not a limited factor in the infection of
plants by fungi.
The writer tested the effect of sunlight on the germination of
the spores of Aliernaria Solani and the conidia of Sclerotinia.
fructigena. The spores of both of these fungi germinate quite as
well in sunlight, whether it be direct, diffuse, glass filtered or
not glass filtered, as they do in darkness, provided that the
conditions of temperature and moisture meanwhile remain near
the optimum. The indications are that the spores of fungi
334 Doran: ON THE GERMINATION OF FUNGOUS SPORES
germinate in either light or darkness, when the other environ-
mental conditions are between maximum and minimum limits.
It is, however, probable that the germination of fungous spores
in nature usually takes place in darkness, since it is in the night
that conditions of temperatures and moisture nearer the opti-
mum for germination usually prevail for a longer time.
Water relation—An examination of the literature indicates
that the moisture requirement of germinating spores is not
uniform. Tulasne (57) was able to germinate teliospores of
the Uredinales as well in a saturated atmosphere as in a drop
of water. Patrigeon (20) found that for the germination of the
conidia of Plasmopara viticola precipitated moisture is necessary
and merely damp air is insufficient. According to DeBary
(47) the spores of the Uredinales germinate better when they are
merely in a moist atmosphere than when they are in water.
Lesage (58) found that the spores of Penicillium glaucum can
germinate in damp air so long as the humidity does not fall
below 82 or 84 per cent. Taubenhaus (51) readily germinated
the teliospores of Puccinia Malvacearum in a humid atmosphere.
Levin (59) was able to infect tomatoes with Septoria Lycopersict
by merely applying dry spores. Weimer (9) found the telio-
spores of Gymnosporangium Juniperi-virginianae unable to
germinate when they were in contact with no moisture except
that in the atmosphere. These spores did not germinate until
the air became super-saturated, so that small drops of water
collected on the slides in contact with the spores. Anderson
(11) states that the spores of Cylindrocladium scoparium never
germinate except when they are directly in water, a moist
atmosphere being insufficient. According to Melhus and Durrell
(7) the urediniospores of Puccinia coronata germinate only when |
they are in direct contact with water. Smiley (33) found that
in the absence of a film of water, the spores of Phyllosticta
Antirrhini do not germinate. Melhus, Durrell and Kirby
found that the sporidia of Puccinia graminis germinate pro-
fusely in drops of water, and but very poorly when their only
moisture is obtained from the atmosphere.
The writer tested the effect of moist air as compared with
precipitated moisture on the germination of the spores named in
TABLE V. The aeciospores of Gymnosporangium clavipes ger-
minate perfectly i in moist air, as they do in water. The spores of
Alternaria Solani and the conidia of Venturia inaequalis germinate
DORAN: ON THE GERMINATION OF FUNGOUS SPORES 335
only 4 per cent in moist air as compared with 100 per cent
in water. This means that in nature the relative number
of spores of these two last named fungi germinating in moist
air is very small, but undoubtedly sufficient to cause some in-
fection. The conidia of Sclerotinia fructigena will not germinate
in moist air but require precipitated moisture. The conidia of
Peronospora pygmaea were also found to require precipitated
moisture rather than moist air for their germination.
Of the spores studied by the writer and mentioned as above in
the literature, in only five cases can they germinate well when
their only source of moisture is the atmosphere. Three germinate
very poorly in moist air. Six germinated only in precipitated
moisture. The evidence is insufficient to warrant the conclusion
that any group of fungi characteristically obtain their water for
germination principally in the form of vapor. When optimum
conditions for laboratory experiments on spore germination are
TABLE V
Effect of moist air as compared with precipitated moisture
n the germination of fungous spores
Spore germinating (relative numbers)
G. clavi- |S. fructi-\P. pyg-
pes gena
|V. inae-
Moisture conditions
la 4 qualis
maea
100 100
0 oO
100
4
Ina drop of distilled water
On dry slide in moist chamber
100 100
4
to be obtained, water in the form of precipitated moisture rather
than water vapor should be supplied. The question of too much
water is probably closely related with the question of too little
oxygen.
SUMMARY
I. The germination of the following spores was studied;
conidia of Venturia inaequalis (Cke.) Wint., conidia of Sclerotinia
fructigena (Pers.) Schrt., spores of Alternaria Solani M.)
Jones & Grout, spores of Botrytis cinerea Pers., spores of
Rhizopus nigricans F. de Wal., aeciospores of Gymnosporangium
clavipes C. & P., and teliospores of Puccinia Malvacearum Mont.
2. Spores of parasitic fungi germinate better when obtained
from the living host than when obtained from artificial media.
336 DorAN: ON THE GERMINATION OF FUNGOUS SPORES
3. Mature spores can germinate through a broader range of
environmental conditions than can immature spores.
4. Freshly mature spores can germinate through a broader
range of environmental conditions than can old spores. As the
spores age, viability at first decreases sharply after which it is
only gradually lost. Longevity of spores is dependent on condi-
tions of storage after detachment from the host. Moisture is
of more importance than temperature in its effect on the length
_of life of the spore.
5. Spores of the Phycomycetes can germinate at the lowest
minimum temperatures, followed in order by the urediniospores,
the aeciospores, and the teliospores of the Uredinales. Aecio-
spores have the lowest optimum temperatures for germination
followed in order by urediniospores, conidia of Phycomycetes,
teliospores, and the spores of the Imperfect Fungi.
6. The nearer all conditions are to the optimum, the shorter
the time required for spore germination.
7. Competition or crowding inhibits spore germination.
This is attributed to insufficient oxygen.
8. Spores of the fungi studied germinated in either light or
darkness.
9. Precipitated moisture is unnecessary for the germination
of some spores if water vapor is available.
New HAmpsuire AGRICULTURAL EXPERIMENT STATION,
DurHAM, New HAMPSHIRE
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4
DorAN: ON THE GERMINATION OF FUNGOUS SPORES
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INDEX TO AMERICAN BOTANICAL LITERATURE
1920-1922
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or any upon American material; the word
America being used in the broadest s
Reviews, and papers that relate Chaioae to foresty, agriculture, horti-
canoes spss aS apt ig of vegetable ee or - oratory methods
nd f bacteriology.
An occasional siete is made i in favor of some paper appearing in an Ameri-
can 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 three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
Anderson, R. J., & Kulp, W. L. Analysis and composition of
corn pollen. Jour. Biol. Chem. 50: 433-453. F 1922.
Anderson, P. J. Development and pathogensis of the onion
smut fungus. Massachusetts Agr. Exp. Sta. 4: 99-133. f.
i~3.° N 1921.
Andrews, A. L. Further bryophytes from North Carolina (and
Tennessee). Bryologist 24: 49-58. 9 Ja 1921.
Andrews, A. L. Notes on North American Sphagnum, IX.
Bryologist 24: 81-86. 27 F 1922.
Armstrong, G. M. Studies in the physiology of the fungi.
XIV. Sulphur nutrition: the use of thiosulphate as in-
fluenced by hydrogen-ion concentration. Ann. Missouri
Bot.: Gard. 8: 237-281. f. 1-27. 3 1921.
Arndt, C. H. The growth of field corn as affected by iron and
- aluminum salts. Am. Jour. Bot 9: 47-71. pl. 4 + f. 1-6.
20 Mr. 1922.
Arthur, J.C. Uredinales collected by Fred J. Seaver in Trinidad.
- Mycologia 14: 12-24. 6 Mr 1922.
Includes 5 new species.
342 INDEX TO AMERICAN BOTANICAL LITERATURE
Acthur, J. C. Uredinales collected by R. Thaxter and J. B.
Rorer in Trinidad. Bot.Gaz.73:58-69. f. 7-4. 18Ja1g22.
Ashe, W. W. The eastern shrubby species of Robinia. Jour.
Elisha Mitchell Sci. Soc. 37: 175-177. Mr 1922.
R. grandiflora sp, nov.
Ashe, W. W. Forest types of the Appalachians and White Moun-
tains. Jour. Elisha Mitchell Sci. Soc. 37: 183-198. Mr 1922.
Avten, J. T. The organic phosphorus content of some Iowa
soils. Soil Sci. 13: 119-124. F 1922.
Awschalom, M. Datos sobre la influencia del selenio en la
vegetacién cuando sustituye al ion sulfurico en el liquido
nutritivo de Knop. Revista. Facult. Agron. 14: 122-162.
1921.
Baas-Becking, L. G. M. The origin of the vascular structure
in the genus Botrichium; with notes on the general anatomy.
Rec. Trav. Bot. Neerlandais 18: 333-372. pl. 5,6 +f. I-40.
1921.
Bailey, J.W. Adventures in mossland.—A hunt for Desmatodon
latifolius (Hedw.) Brid. Bryologist 24: 86-88. 27 F 1922.
Bailey, L. W. Diatoms from the Quill Lakes, Saskatchewan,
and from Airchie, Alberta. Contr. Canadian soos if:
457-108." PF. a; o> AGeT:
Barker, E. E. The architecture of the coco palm. Nature-
study Rev. 18: 44-50. f. 1-4. Ja-F 1922.
Benoist, R. Contribution a 1’étude de la flore des Guyanes.
Bull. Soc. Bot. France 68: 311-323. 2 Au 1921.
Berry, E. W. Additional occurrences of Pleistocene plants.
Torreya 22: 10, 11. F 1922.
Berry, E. W. Carboniferous - a from. Peru. Am. Jour.
Sci. 203: 189-194. Mrs
Berry, E. W. The flora of the Woodbine sand at Arthur’s
Bluff, Texas. U.S. Geol. Survey 129: 153-180. pl. 36-40
+ f. 12. 23 Mir. 1685.
Includes Trochodendroides, gen. nov.
INDEX TO AMERICAN BOTANICAL LITERATURE 343
Birge, E. A., & Juday, C. Further limnological observations on
the Pinger Lakes of New York. Bull U. S. Bur. Fisheries
37: 209-252. 1922.
Includes notes on phytoplankton.
Blake, S. F. New Plants from Guatemala and Honduras.
Contrib. U.S. Nat. Herb. 24: 1-32. pl. 1-10+ f. T+. 1922.
Includes 46 new species.
Blake, S. F., & Sprague, T. A. On the generic name Wikstroemia.
Jour. Bot. 60: 52-54. F 1922.
Blakeslee, A. F. Variations in Datura due to changes in chro-
mosome number. Am. Nat. 56: 16~31. f. 1-7. F 1922.
Blomquist, H. L. Vascular anatomy of Angiopteris evecta.
Bot. Gaz. 73: 181-199. pl. 5-8 + f. 1-8. M 1922.
Botto, A. Una nueva forrajera para las regiones del norte.—El
pasto de Rhodes. Revista Facult. Agron. 14: 170-178.
f. 1-3. 1921.
(Chloris Gayana Kunth.)
Bouyoucos, G. A study of the fertility of the pols of Greece.
Soil Sci. 13: 63-79. F 1922.
Brand, A. Borraginaceae-Borraginoideae epee Pflan-
zenreich IV. 252: 1-183. f. I-22. 22 N 1921.
Broadway, W. E. Botanical collecting in French Guiana. Jour.
New York Bot. Gard. 22: 177-183. O 1921.
Burns, G. P. Eccentric growth and the formation of redwood
in the main stem of conifers. Vermont Agr. Exp. Sta.
Bull. 219: 1-16. pl. 1-q4 + f. I-10. Je 1920.
Butler, E. J. Phytopathology in the United States. Kew Bull.
Misc. Inform. 1922: 85-91. 1922.
Canela, P. F. Sobre la presencia de una plaga en los frutales
en San Luis. Bol. Minis. Agric. de la Nacion 26: 253-256.
1921.
Cannon, W. A. Plant habits and habitats in the arid portions of
South Australia. Carnegie Inst. Washington Publ. 308:
1-139. pl. 1-34 + f. I-31. 1921.
344 INDEX TO AMERICAN BOTANICAL LITERATURE
Caverhill, P. C. Forestry problems in British Columbia. Jour.
For. 20: 44-53. Ja 1922.
Chardon, C. E. Contribution to our knowledge of the Pyre-
nomycetes of Porto Rico. Mycologia 13: 279-300. pl. 13-
15. + f. 1-4. 21 Ja 1922.
Includes 9 spp. nov. and 1 comb. nov.
Churchill, J. R. Cimicifuga racemosa in Massachusetts.
Rhodora 23: 201-203. 9 Ja 1922.
Clements, F. E. Aeration and air-content. The role of oxygen
in root activity. Carnegie Inst. Washington Publ. 315:
3-183; 1921:
Clokey, I. W. Carex notes. Am. Jour. Sci. 3: 88-91. pl. 2.
Ja 1922.
Includes Carex apoda and C. Paysonis, spp. nov.
Coates, L. The “peach-almond” hybrid. Jour. Hered. 12:
328, 329. 20 F 1922. [Illust.]
Cocks, R. S. A list of the trees of Louisiana. Jour. Arnold
Arbor. 2: 204-216. Ja 1922.
Cockerell, T. D. A. Flora of Porto Santo. Torreya 22: 4-10.
fo2, 2 F 1922.
Coker, W. C., & Grant, F. A. A new genus of water mold
related to Blastocladia. Jour. Elisha Mitchell Sci. Soc.
37: 180-182. pl. 32.. Mr 1922.
Septocladia gen. nov. and S. dichotoma sp. nov.
Collins, G. N. Teosinte in Mexico. Jour. Hered. 12: 339-350.
24-9; 62s,
Conti, M. El campo experimental de hidrologia agricola.
Revista Facult. Agron. 14: 97-121. 1921. [Illust.]
Cooper, W. S. The ecological life history of certain species of
Ribes and its application to the control of the white pine
blister rust. Ecology 3: 7-16. f. 1-3. 25 F 1921.
Cundall, F., & Fawcett, W. Dr. Anthony Robinson of Jamaica.
Jour. Bot. 60: 49-52. F 1922.
Davis, B. M. Species, pure and impure. Science II. 55: 107-
Tig. 3 P1622. ;
INDEX TO AMERICAN BOTANICAL LITERATURE 345
Demerec, M. Heritable characters of maize. X. Zebra striped
leaves. Jour. Hered. 20: 406, 407. f. 12. N 1921.
Denison, I. A. The nature of certain aluminum salts in the
soil and their influence on ammonification and nitrification.
Soil Sci. 13: 81-106. F 1922.
Denny, F. E. Formulas for calculating number of fruits re-
quired for adequate sample for analysis. Bot. Gaz. 73:
44-57...18 Ja 1922.
Detwiler, S. B. Blister rust appears in the Puget Sound region.
Am. For. 28: 97, 98. F 1922.
Dietel, P. Zur Umgrenzung der Gattung Pileolaria Cast. Ann.
‘Mycol. 19: 300-303. 1921.
Dodge, C.K. Miscellaneous papers on the botany of Michigan.
Michigan Geol. & Biol. Surv. 31: 15-234. 1921.
Plant lists.of various sections of state. Includes biography, portrait
and map.
Dunbar, J. How to grow ee Jour. New York
Bot. Gard. 22: 184-190. 1921
Durand, E.J. The genus Catinella. Bull. Torrey Club 49: 15-21.
21 Mr 1922.
Dwight, T. W. Timber administration of the Dominion Forest
Service. Jour. Forestry 20: 4-9. Ja 1922.
Elliott, C. Sterility of oats. U.S. Dept. Agr. Bull. 1058: 1-8.
pl. I-4. 6 Mr 1922.
Emerson, R. A. The nature of bud variations as indicated by
their mode of inheritance. Am. Nat. 56: 64-79. F 1922.
Emerson, R. A., & Hutchinson, C. B. The relative frequency
of crossing over in microspore and in megaspore develop-
ment in maize. Genetics 6: 417-432. S 1921. —
Eriksson, I. The life of Puccinia Malvacearum Mont. within
the host plant and on its surface. Phytopathology 11: 459-
463. 25 F 1922.
Ezekiel, W. N. Some factors affecting the production of
apothecia of Sclerotinia cinerea. Phytopathology 11: 495-
499. 21 Mr 1922.
346 INDEX TO AMERICAN BOTANICAL LITERATURE
Faris, J. A. Violet root tip (Rhizoctonia crocorum DC.) in the
United States. Phytopathology 11: 412-423. 16 F 1922.
Farr, C. A. Quadripartition by furrowing in Sisyrinchium. Bull.
Torrey Club 49: 51-61. pl. 2. Mr 1922.
Farwell, O. A. Corrections in botanical nomenclature. Am.
Midl. Nat. 8: 33-35. Ja 1922.
Includes new combinations under Dilepyrum (1) and Gardenia (3).
Faull, J. H. Some problems of forest pathology in Ontario.
Needle blight of white pine. Jour. For.20:67-70. Ja1922.
Fernald, M. L. The Gray Herbarium expedition to Nova
Scotia. Rhodora 23: 89-111. pl. 130. 19 My 1921; 23:
130-152. 26 Au 1921; 23: 153-171. 24 O 1921; 23: 184-—
195. 15 D1921; 23: 223-245. 26 Ja 1922; 23: 257-278.
$. 133. 27 F 1922; 23: 264-300. 22 Ap 1922."
Includes new species in Lophiola (1) and Antennaria (1), 5 12 new varieties
and 2 new forms.
Fink, B. An addition to the distribution of a rare fungus.
Mycologia 14: 49, 50. 6 Mr 1922.
Tylostema verrucosum in Ohio.
Fitzpatrick, H. M., Thomas, H. E., & Kirby, R.S. The Ophio-
bolus causing take-all of wheat. Mycologia 14: 30-37.
pl. 10 + f. r. 6 Mr 1922.
Fromme, F. D. Incidence of loose smut in wheat varieties.
Phytopathology 11: 507-510. 21 Mr 1922
Gail, F. W. Factors controlling distribution of Douglas fir in
semi-arid regions of the Northwest. Ecology 2: 281-291. ff.
I-32. -20 Ja 1622.
Gaudron, J. El peso de las raices del Manihot utilissima en
relacion con la fasciacion de los tallos. Arch. Asoc. Peru
1: 20-37. pl. 1, 2. 1921.
Gericke, W. F. Certain relations between the protein content
of wheat and the length of the growing period of the head-
bearing stalks. Soil Sci. 13: 135-138. F 1922.
Gerould, J. H. The dawn of the cell theory. Sci. Monthly 14:
268-277. Mr 1922.
INDEX TO AMERICAN BOTANICAL. LITERATURE 347
Girola, Cc; D. Agallas de corona del duraznero. Bol. Minis.
Agric. 26: 257-259. pl. z,.2 (1 colored), Jl-S 1921,
Pseudomonas tumefaciens.
Girola, C. D.. Cultivo de la Yerba mate Torrefaceion y pre-
Pataqion del producto. Bol. Minis Agric. 26: 447-488. /f.
I- 1921.
Girola, C. D. Sobre algunas enfermedades de la papa. Bol.
Minis. Agric. 26: 260-264. pl. z, 2. JI-S 1921.
Gleason, H. A.. The witch-hazels. Jour. New York Bot.
Gard. 23: 17-19. F 1922.
Goebel, K. Erdwurzeln mit Velamen. Flora 11821-26../f.4;52
1922.
Graff, P. W. Philippine Basidiomycetes—IV. Bull. Torrey
Club 48: 285-295. 19 Ja 1922.
Gress, E. M. Annotated list of fossil plants of the Dakota
formation (Cretaceous) in the collections at the Carnegie
Museum. Ann. — Mus. 13: 274-332. pl. 14, 15.
1921.
Includes 3 spp. nov.
Greaves, J. E. Influence of salts on bacterial activities “of soil.
ot. Gaz. 73: 161-180. Mr 1922
Greenman, J. M. ‘Two new senecios from the West Indies.
Ann. Missouri Bot. Gard. 8: 97-102. pl. 1-2. 9 Ja 1922.
Includes S. subsquarrosus and S. Freemanti, spp. nov.
Grout, A. J. Mosses of a Staten Island house and lot. Bryolo-
gist 24: 64. 9 Ja 1922
Guilliermond, M. A. Nouvelles observations sur l’origine des
plastides dans les phanérogames. Rev. Gén. Bot. 33: 401-
419, 449-466. f. 1-8. 1921.
Haasis, F. W. ~ Relations between soil-type and root form of
western yellow pine a Ecology 2: 292-303. f. 1-3.
20 Ja 1922.
Hall, H. M., & Long, F. L. Rubber content of North American
plants. Carnegie. Inst.: Washington: Publ. 3133 I-65. pl.
2-3. (202.
348 INDEX TO AMERICAN BOTANICAL LITERATURE
Hallermeier, M. Ist das Hangen der Bliiten eine Schutzein-
richtung? Flora 115: 75-101. 1922.
Harms, H. Uber einige Carica-arten aus Siidamerika mit
besondere Beriicksichtigung der peruanischen Arten. No-
tizbl. Bot. Gart. u. Mus. Berlin-Dahlem 8: g1-100. 1 Ap
1922.
Includes 4 spp. nov.
Harper, R. M. A botanical bonanza in Tuscaloosa County,
Alabama. Jour. Elisha Mitchell Sci. Soc. 37: 153-160.
pl. 28. Mr 1922.
Harper, R. M. Geography of central Florida. Florida Geol.
Surv. Ann. Rep. 13: 71-307. f. 7-43. 1921.
Harshberger, J. W. Geographic names and terms of signi-
ficance in plant geology and ecology. Bull. Geog. Soc.
Philadelphia 20: 32-46. Ja 1922.
Harvey, L.H. Yellow-white pine formation at Little Manister,
Michigan. Bot. Gaz. 73: 26-43. f. I-6. 18 Ja 1922.
Haskell, R. J. Phytophthora infestans on egg plant in the United
Stutes. Phytopathology 11: 504, 505. 21 Mr 1922.
Hayes, H. K., & Stakman, E.C. Resistance of barley to Hel-
minthosporium sativum P. K. B. ee ey 11: 405-
411. 16 F 1922
eo Se. coe genera of the Clostridiaceae. Jour. Bact.
1-38. Ja
Includes 25 new genera.
Hibbard, P. L. Some experiments on reclamation of infertile
alkali soils by means of gypsum and other treatments.
Soil Sci. 13: 125-134. F 1922.
Higgins, B. B. Notes on the morphology and systematic re-
lationship of Sclerotium Rolfsii Sacc. get Elisha Mitchell
Sci. Soc. 37: 167-172. pl. zo. Mr
Hoerner, G. R. Infection capabilities a: crown rust of oats.
Phytopathology 12: 4-15. 10 Mr 1
Hollick, A. Local flora notes—Staten Island. Torreya 22:
1-3. Ja-F 1922.
Vol. 49. No. 12
BULLETIN
OF THE
TORREY BOTANICAL CLUB
DECEMBER, 1922
Miscellaneous notes on plants of Southern California—II*
Puitiep A. Munz AnD Ivan M. JOHNSTON
LEMNA TRISULCA L.
Lemna trisulca L. Sp. Pl. 970. 1753.
This plant, reported by Abrams (Fl. Los Angeles 78. 1904)
from Bear Valley in the San Bernardino Mountairis and by
Torrey (Pacific R. R. Rep. 4: 142. 1856) from San Gabriel
Creek, has been collected by Miss G. Corwin at the mouth of
Santa Ana Canyon near Mentone, where it was found in a
ditch at about 2000 feet altitude. It is, therefore, apparently
to be looked for at various points along the Santa Ana River
MICROSTYLIS MONOPHYLLOs (L.) Lindl.
Ophrys monophyllos L. Sp. Pl. 947. 1753.
Microstylis monophyllos Lindl. Gen. Sp. Orch. 19. 1830.
A few typical specimens of this species, F. W. Petrson 2271,
were collected in marshy ground at 8000 feet elevation on the
South Fork of the Santa Ana River in the San Bernardino
Mountains, where they grew with such plants as Gentiana
simplex, G. humilis, Parnassia cirrata, and Luzula comosa. This
is the first collection west of the Rocky Mountains.
“ Phoradendron californicum var. distans forma leucocarpum
Trelease - forma nova
Like the variety distans, but having white berries.
[The BuLLetin for November (49: 313-348) was issued November 9,
1922.
e first paper of this series was published in the BuLLETIN for Feb-
ruary, 1922 (49: 31-44)
349
350 | Munz AND JOHNSTON: PLANTS OF CALIFORNIA—IT
Type: Twenty-nine Palms, southern edge of Mohave
Desert, May 1, 1921, Munz 455c¢, (Baker Herb. 9547).
Very conspicuous as white-fruited clumps growing in bushes
of mesquite with the typical red-fruited form. Specimens of
this collection were sent to Dr. Wm. Trelease who considered
it worth recognition and permitted us to publish it. A second
collection, Munz 4703, likewise on mesquite, from Andreas
Canyon near Palm Springs, can also be reported.
. Eriogonum nodosum var. Jaegeri var. nov.
With habit, inflorescence, and floral structures of the typical
form of the species but with stems tomentose only below on the
leaf-bearing CoeLiral, the upper parts of the stem and the in-
florescence being glabrous with, at most, a little wool in the
axils. rie iaaes ‘plabrous or slightly floccose, the subtending
bracts woolly on their inner edges.
Tyre: Dry Morongo Wash, near Hole-in-the-Wall Springs,
seed ere sete November 15, 1921, E. C. Jaeger 251 (Baker
Herb.
This ak form of E. nodosum Small (Bull. Torrey Club
25: 49. 1898) was collected at the eastern base of the San
Bernardino Mountains, between 2500 and 3000 feet altitude,
and well deserves recognition because of the totally different
aspect as compared with the typical form. It is reported as
very plentiful by its finder, Mr. Edmund C. Jaeger, one of the
most active collectors of the desert flora.
. Eriogonum fasciculatum var. flavoviride var. nov.
Vith capitate inflorescence and general habits of ie var.
‘kthen T. & G., but lower in stature, about A foot high;
light green in color instead of ashy; leaves glabrous and green
above, but somew what tomentose and pallid nate awe glabrous
without and hairy within
Type: Pinyon Wells, southern edge of Mohave Desert,
April 30, 1921, Munz 4505, (Baker Herb. 9425).
This peculiar yellowish green glabrate plant was found to
be abundant in a sandy desert canyon in high Lower Sonoran
Zone above Pinyon Wells. It was associated all along the
canyon with var. polifolium, but was very distinct from this in
size and color. We have seen also material from Cottonwood
Pass, Hall 6093, about fifteen miles to the southeast and in the
same general mountain range; here too it grew with var. polifol-
MUNZ AND JOHNSTON: PLANTS OF CALIFORNIA—II 351
zum but was smaller and earlier in anthesis. The relation of this
glabrous variety to var. polifolium is analogous to that existing
between typical E. fasciculatum and the pubescent var. foliolosum.
ALLIONIA PUMILA Standley
Allionia pumila Standley, Contr. U. S. Nat. Herb. 12: 345.
1909.
Allionia Brandeget Standley, I. c. 346.
The only Allionia hitherto reported from California is A.
Brandegei, the type of which came from the Providence Mountains
in the eastern part of the Mohave Desert. Two collections
recently made by F. W. Peirson in the San Bernardino Mountains
are of this species, which has recently been included in A.
pumila by Standley (No. Am. Flora 21: 226. 1918). These
collections not only extend the range of this species to the western
part of the desert but even into the coastal drainage. One
(Peirson 2257) from the Santa Ana River, at 4500 feet elevation,
is twice as tall as the type of A. Brandegei and has the leaves
broadly ovate and cordate, rather than oblong and scarcely
cordate. The other plant (Peirson 1850), from Cushenberry
Grade, is more like Brandegee’s type collection, being about
ten inches high and with leaves ovate to oblong.
ALLIONIA NYCTAGINEA Michx.
Allionia nyctaginea Michx. Fl. Bor. Am. 1: 100. 1803.
Allionia ovata Pursh, Fl. Am. Sept. 97.
The ovate-leaved form of the above a dites was found in
June, 1919, to be locally established along a siding of the Santa
Fe Railroad about a mile west of Upland, Johnston 2169. Ex-
amination of the locality in January, 1922, showed the colony
to have been destroyed by the oil spray used by the railroad
for weed destruction. This is apparently the first record for
_ the state.
/Scopulophila Rixfordii (Brandegee) comb. nov.
Achyronychia Rixfordii Brandegee, Zoe 1: 230. 1890.
Scopulophila nitrophiloides Jones, Contr. W. Bot. 12: 5. 1908.
Eremolithia Rixfordii Jepson, Fl. California 499. f. 100. 1914.
This odd desert plant seems to be generically distinct from
Achyronychia, but Jones’s Scopulophila has precedence over
the much later generic name Eremolithia of Jepson,
352 MUNZ AND JOHNSTON: PLANTs oF CALIFORNIA—II
/ Claytonia lanceolata var. Peirsonii var. nov.
As in the typical form of the species, but with peduncle and
primary axis of a pmloae ae shortened so as to make the flowers
appear umbellat
TYPE: near ae s Cabin on Ontario Ridge, San Gabriel
Mountains, at 8200 feet altitude, May 29, 1920, Wa. &.
Mabel Peirson (Baker Herb. 10524).
Known only from the eastern portion of the San Gabriel
Mountains where, beside the type, we have seen a collection,
. W. Peirson 2151, from the Devil’s Backbone at 7000 feet
altitude. It is probable that this variety will necessitate
a es ee 4 the status of C. umbellata Wats. (Bot.
King Exp. 43. pl. 6, f. 4-5. 1871) of the northeastern part
of California, vee re principal character of that species is the
same umbellate inflorescence that characterizes our variety.
The new variety, however, may be told from C. umbellata
by its shorter-petioled, broader leaves, which are widest below
the middle (instead of above), and by its shorter stem, larger
flowers, and isolated southern range.
THLASPI ARVENSE L.
Thlaspt arvense L. Sp. Pl. 646. 1753.
This crucifer, common in the Rocky Mountain region, can
now be reported from Southern California (F. W. Peirson 66)
at Henniger’s Flats near Pasadena. The only other collection
we have seen from California is by Chase from Adin, in Modoc
County.
’ Draba corrugata var. saxosa (Davidson) comb. nov.
Draba saxosa Davidson, Bull. So. Cal. Acad. 19: 11. 1920.
The phase of Draba corrugata that is found on San Jacinto
Peak may well be deserving of a varietal name, but scarcely
of a specific one. As pointed out in another paper (Johnston,
Pl. World 22: 88. 1919) the San Jacinto plant differs only in
matters of degree and is closely simulated by some collections
from the San Bernardino Mountains,
MUNZ AND JOHNSTON: PLANTS OF CALIFORNIA—II = 353
Sphaeralcea rosacea sp. nov.
A shrubby perennial forming loose clumps 10-15 dm. hich:
stems ashy, stellate-tomentose, slender; ascending, virgate:
leaves remote, ovate- or oblong-cordate and somewhat three-
lobed, doubly crenate, stellate-pubescent but greener above,
— usually 2-2.8 cm. long and 1.5-2.5 cm. wide, petioles
~—20 mm. long; inflorescence a very loose open —— =
m. long, the ascending branches remote and o
Deed: ‘calyx stellate-tomentose, at anthesis Hen mim. high
and 5~7 mm. wide, sins lanceolate or oblong-lanceolate lobes,
at fruiting 10-14 mm. long, bractlets setaceous, deciduous,
mm. long; 3-20 long: petals pinkish when
fons h but drying a rose-violet, 17-21 mm. long, claw densely
villous-ciliate; fruit much depressed, etallnteteabeorea: carpels
one-seeded, about 3. 5 mm. high, oblong in outline, sides smooth
or at least not reticulate, seminiferous only below, the upper
half thinner and scarious.
Type: Palm Springs, April 2, 1917, Alice B. Chittenden
(Herb. Cal. Acad.).
A very ornamental species that is known to us only from
the vicinity of Palm Springs, Riverside County, where it is a
common plant in rocky ground in the canyons opening into the
Colorado Desert. Besides the type we have seen a collection
by Parish, No. 4109, and one by Johnston. This plant is the
“‘purple-flowered form of S. ambigua,’’ which is mentioned by
Robinson (Synop. Fl. 1: 315. 1897) in his review of the genus.
Though it has long passed as a form of S. ambigua, it is amply
distinct and probably not even an immediate relative of that
species, differing from it in habit and stature; in size, color
and disposition of flowers; and in shape and texture of leaves.
When we first saw the two species growing together, as they
commonly do about Palm Springs, we never once thought
of associating the two plants under one name; in fact, basing
determination on superficial resemblances, the new species was
taken to be a species of Malvastrum. The proposed species
bears a considerable resemblance to certain of the shrubby mal-
vastrums, having more or less woody stems and similarly colored
flowers, but the technical characters of the carpels definitely
ally it with Sphaeralcea.
VIOLA SHELTONI Torr.
Viola Sheltoni Torr. Pac. R. R. Rep. 4:67. pl. 2. 1856.
354 Munzanp JounsTon: PLANts of CALIFoRNIA—II -
Locally common on moist, shaded, oak-covered canyon-sides
in the upper chaparral belt on the south slope of the San Antonio
Mountains, where specimens have been collected in Charcoal
Fork of Cucamonga Canyon, at 4200-4500 feet altitude, Johns-
ton 1303, and near Brown’s Flats, at 4200 feet, Johnston 1759.
The specimens appear the same, character for character, as the
common plant of Mendocino and Siskiyou Counties. The plant
was known to range as far south as Mariposa County, where it
was collected by Congdon, but between that station and the
one in Southern California, which we now report, there lies a
gap of over 280 miles in which the plant is not known to occur.
This station for V. Sheltont was announced as V. lobata by
Johnston (Bull. So. Calif. Acad. 17: 65. 1918).
OENOTHERA SPECIOSA Nutt.
Oenothera speciosa Nutt. Jour. Acad. Phila. 2: 119. 1821.
Hartmanntia speciosa Small, Bull. Torrey Club 23: 181. 1896.
Well established along Chino Creek, from Pomona to near
the confluence with the Santa Ana River. First collected
several years ago by Johnston. Recently the following specimens
have been collected near Pomona, all in moist soil along the
stream: Mrs. Leningion, Munz 4680, 4560. These discoveries
add a new species and subgenus of Oenothera to the flora of Cal-
ifornia. Our material is atypical in its rounded instead of ob-
cordate petals and in being tallet, some of the specimens attaining
a height of four feet.
’Oenothera cardiophylla var. splendens var. nov.
A large flowered variation with hypanthium two-thirds as
long as the capsule and 20-25 mm. long; petals 15 mm. long.
Type: high among rocks at the foot of The Needles, Colorado
River, March 7, 1910, Grinnell (Univ. Calif. Herb. 196590).
A very remarkable variation of the species and, because of
its large flowers, probably of ornamental value. We know of
but two collections, the one from The Needles by Grinnell and
the other from the Chocolate Mountains by Childs. The
foliage and habit of the plant are those characteristic of the
species. No doubt some will consider this distinct enough for
a species, but we are deterred from publishing it as such because
of the immense amount of variation in the floral structures of
Muwnz ANnp JouNsTON: PLANTS or CatirorntA—II 355
typical O. cardiophylla, of which this seems only to be a remark-
able extreme.
CHIMAPHILA UMBELLATA (L.) Nutt.
Pyrola umbellata L. Sp. Pl. 396. 1753.
Chimaphila umbellata Nutt. Gen. 1: 274. 1818.
Chimaphila occidentalis Rydb. No. Am. Flora 29: 30. 1914.
This species can now be reported from Southern California,
having been collected in the Dollar Lake region of the San
Bernardino Mountains at 8750 feet altitude, F. W. Peirson 820.
This discovery and that of Pyrola picta Smith, Johnston 2860,
add two species of Pyrolaceae to the list prepared by Parish
(Pl. World 20: 247. 1917) for the San Bernardino mountains.
“Asclepias eriocarpa var. microcarpa var. nov.
Follicles 3-4 cm. long; seeds 5 mm. long; coma 15 mm. long;
pedicels 2—3.5 cm . lo OnE in flowers and vegetative parts like the
typical form of the species
Type: North sits of Deep Creek, San Bernardino Moun-
tains, on a sunny hillside at 6000 feet altitude, July 17, 1921,
I. M. Johnston 2833 (Baker Herb. 9563.)
This variety is the form of the species frequent through the
pine belt of the San Bernardino Mountains and differs from the
typical form, which is not uncommon on the plains at the foot
of the mountains, in its much smaller follicles. Many plants
of the valley-inhabiting A. eriocarpa Benth. have been seen by
the authors, but none have been noted which failed to have
fruit much larger than in the variety here described. A. erto-
carpa has follicles 8-10 cm. long on pedicels 4—6 cm. long, while
the seeds measure about 8 mm. in length and have a coma 24
mm. long.
The small follicles can not be attributed to the blighting
action of frost, for all produce perfect seeds and in all colonies seen
the fruit was uniform in size and as here described. Further
exploration may reveal the fact that the pine-belt form of A.
eriocarpa found in the San Jacinto and San Gabriel Mountains
is also referable to this variety; fruiting material from these
ranges is lacking.
356 Munz AND JOHNSTON: PLANTS oF CaLIFoRNIA—II
‘ Phlox bernardina sp--nov:
_ Phlox dolichantha Brand; Engler, Pflanzenreich 4*°°: 67. f.
18, A-D. 1907; Nels. W. Am. Phloxes 28. 1898; Milliken,
Univ. Cal. Pub. Bot. 2: 65. 1904. Not P. dolichantha Gray,
Proc. Am. Acad. 22: 310. 1887.
TYPE: in open pine forest, 6000 feet altitude, Bear Valley,
June 1886, Parish 1818, (Univ. Calif. Herb. 194015)
The new species indicated here is the plant from the San
Bernardino Mountains that has been known as P. dolicantha.
The type of P. dolichantha Gray, however, came from the Pahran-
gat Mountains in southern Nevada and is evidently a form of
what has been called P. Stansburyi brevifolia and P. superba.
In the recent treatments of Phlox by E. Nelson, Milliken, and
Brand, the plant from the San Bernardino Mountains has
usurped the name of P. dolichantha to the exclusion of the
original Nevada plants. P. bernardina differs from P. dolichantha
and its immediate relatives in its very elongated corolla-tube,
which is over four, instead of less than three, times the length
of the calyx; in having its stems not shrubby below but her-
baceous to the ground; and in its habitat, which is not at low
altitudes in desert mountains but in the open pine forests of
the San Bernardino Mountains. The new species is separated
from its nearest relative by over 100 miles. It is known only
from San Bernardino Mountains and appears to have been
collected there in Bear Valley only.
NAMA HUMIFUSUM Brand
Nama humifusum Brand, Beitrage Hydrophyll. 9. 1911.
In Southern California this species has been known only
from the three localities (Soldiers’ Home, Los Angeles County,
and Diamond Lake and Sweetwater Valley in San Diego County)
given by Brand in the Pflanzenreich (4%: 146. 1913); the
plant, however, may be reported from Laguna Canyon in Orange
County, where it was collected in May, 1919, Munz, Street,
& Williams 268r.
Lycrum SPENCERAE Macbride
Lycium Spencerae Macbride, Contr. Gray Herb. 53: 18. 1918.
The type of this species proves to be a flowering specimen of
Prunus fasciculata (Torr.) Gray (Proc. Am. Acad. 10: 70.
MUNZ AND JOHNSTON: PLANTS OF CALIFORNIA—II 357
1874). The identifying of this lycioid almond with L. Spencerae
adjusts for local botanists the problem of explaining how so
distinct a species, as this Lyciwm seemed to be, could so long
go undetected in a locality so frequently visited as Cajon Pass.
We are under obligations to Mr. F. Macbride for making at
the Gray Herbarium the comparisons necessary for the establish-
ing of the identity of the two species.
’Penstemon Clevelandi var. connatus var. nov.
With habit of growth-and flowers of the species, but with
leaves jagged-serrate, usually aps and with at least the
our upper pairs connate-perfoliat
Type: near Van Deventers, Be a base of the San
Jacinto Mountains, June 1901, H. M. Hall 2149 (Univ. Calif.
Herb. 54999).
A very well marked variation that is apparently restricted
to the eastern (Colorado Desert) base of the San Jacinto Moun-
tains. Referable to this variety are the following: Hall r160
from the type locality, Parish Bros. 1216 and Johnston, April,
1917, from near Palm Springs. The most evident relationship
is with the variety Stephensi of the eastern Mohave Desert,
the new variety agreeing in its foliar characters but differing
in its somewhat smaller corolla and beardless sterile stamens.
In our recent discussion of the var. Stephensi (Bull. Torrey
Club 46: 41. 1922) the specimens of P. Clevelandi Gray mentioned
as simulating that variety in habit are in fact referable to var.
connatus.
P. Clevelandi in its typical form occurs along the desert
borders to the south of the range of the var. connatus. The
northernmost locality seen for it is Coyote Canyon, Hall 2766.
In the typical form the leaves are entire or rarely remotely
dentate, never glaucous and are entirely free or, at most, only
partially joined and not completely fused, with the suture line
obliterated as in the variety.
/ Galium angustifolium var. pinetorum var. nov.
A low herbaceous perennial, stems 2-4 dm. high, simple,
erect, tufted or are from along short, prostrate, pean
subterranean stem
Type: Sierra aii Mountains, ee Angeles County, 1893,
A. Davidson (Univ. Calif. Herb. 28169).
358 Munz AND Jonnston: PLANts oF CaLirorniA—II
A variety of the pine-belt and differing from the typical
form of the species only in habit of growth. Additional speci-
mens referable to this variety are the following: Santa Lucia
Mountains, K. Brandegee, on the north, and Cuyamaca Moun-
tains, T. S. Brandegee, July 7, 1894, on the south. Plants from
the San Jacinto Mountains, Hall 2509, have the growth-habit
of the variety, but are strictly branched and become 5-7 dm.
high. The variety has been mentioned without a formal name
by Johnston (Pl. World 22: 118. 1919).
¥Galium Hallii sp. nov.
Dioecious ie eke at the base, with relatively
few long (4-6 dm. mbent quadrangular stems, you ipa
stems short-hispid, older stems glabrate and with a shin
foliating papery bark; leaves in fours, hirsute like the tenes: Se
light green, ovate-elliptical to elliptical, 5-11 mm. long, sessile,
margins strongly revolute, midvein strong, lateral veins weak
or absent; flowers borne in loose leafy clusters on the con-
spicuously and gracefully recurved tips of the branchlets, yellow-
ish, about 2 mm. in diameter, bearing a few stiff hairs; fruit
black, slightly juicy, ned. 3 mm. in diameter, covered with
dense villous hairs of 2 mm. length.
Type: Coldwater Fork of Lytle Creek, San Gabriel Moun-
tains, in gravelly ground at 5200-5700 feet altitude, July 7,
1918, Johnston 2067 (Univ. Calif. Herb.).
Specimens examined: Coldwater Fork of Lytle Creek, San
Gabriel Mountains, 5500 ft. alt. Johnston 1667; Lytle Creek,
5750 ft. alt., Hall 1227; Rock Creek Canyon, San Gabriel Moun-
tains, Abrams & McGregor 598; Seymour Creek, Mt. Pinos
region, 6700 ft. alt., Hall 6505; Tehachepi, 1889, Greene; Erskin
Creek, 4—5000 ft. alt., Purpus 5340,
This is a remarkably clear-cut undescribed species of the
G. multiflorum-alliance that has been confused with G. multt-
florum, G. stellatum, and G. occidentale; from all of these it
certainly differs and can be distinguished by its nodding flower-
clusters. The plant has been recognized as distinct for several
years, and the increasing material at hand seems to justify its
description as a new species. It has been treated as ‘‘Galium
sp.”’ by Johnston (Pl. World 22: 118. 1919).
The range is eminently a natural one, for the plant occurs
at and just below the lower relies of the _— belt i in the rocky
canyons that open into th of the Mohave
Munz AND JOHNSTON: PLANTS OF CALIFORNIA—II 359
Desert. The type locality, the southernmost known station, is
almost on the Los Angeles—San Bernardino County line;
from this point the range extends northward to Erskin Creek
in the Sierra Nevada of Kern County.
It might be mentioned here that G. occidentale McClatchie
(Erythea 2: 124. 1894) is a trivial form of G. californicum
H. & A., and that G. grande McClatchie (J. c.) is indistinguishable
from G. pubens Gray. These identities are established by a
study of authentic specimens.
BAHIA DISSECTA (Gray) Britton
Amauria dissecta Gray, Mem. Am. Acad. II. 4: 104. 1849.
Bahia dissecta Britton, Trans. N. Y. Acad. Sci. 8: 68. 1888.
Villanova dissecta Rydb. Bull. Torrey Club 37: 333. 1910.
Amauriopsis dissecta Rydb. No. Am. Flora34:37. 1914.
_ The first collection of this species to be reported from the
state was made in the San Bernardino Mountains, F. W. Peir-
son 2245, along the Santa Ana River, at 6500 feet elevation,
August 24, 1920.
PoMoNA COLLEGE
CLAREMONT, CALIFORNIA
sh
Dei ics,
Notes on Carex—XI]
KENNETH KENT MACKENZIE
1. AN INLAND RELATIVE OF A SALT MARSH SEDGE
The only representative of the Ovales which is an inhabitant
of salt marshes, as far as I am aware. is Carex hormathodes
Fernald, which is an abundant and characteristic species along
the North American coasts from western Newfoundland to
southeastern Virginia.* In 1902 attention was first drawn by
Professor Fernald to an allied pla 1t found in fresh water swamps
and reported by him as growing from Massachusetts to the
District of Columbia. This was first treated by him as Carex
tenera Dewey var. Richii Fernald, and later on as Carex hor-
mathodes var. Richii Fernald.
In northern New Jersey this variety Richii is an abundant
plant in places in swampy meadows, and with its graceful
nodding heads is one of the most pleasing and attractive of our
sedges. It is in every way a much more handsome plant than
is Carex hormathodes. But not only does it have an entirely
different habitat and a different appearance, but it has technical
characters which are absolutely good. The perigynia have
suborbicular bodies abruptly contracted into the beaks; the
perigynia are loosely ascending or spreading in the spikes;
the terminal spike is abruptly contracted at base below the
perigynia, and the staminate portion is conspicuous; and the
spikes are scattered in a very flexuous head. In Carex hormatho-
des on the other hand the perigynia have oblong-ovate bodies
much less abruptly contracted into the beaks; the perigynia
are appressed ; the terminal spike tapers at base, and the staminate
portion is short; and the upper spikes are approximate. The
* The inland reports for this species are all erroneous, as also are the re-
ports of its occurrence from further south than southeastern Virginia. It
however is to be expected at least as far south as North Carolina, The var.
invisa (Boott) Fernald represents nothing but stunted specimens.
361
362 MAcKENziE: Notes ON CAREX—XII
differences between the two species are well brought out in the
figures (No. 356-8) in Gray’s Manual.
In proposing specific rank for this plant I am glad to say
that I have the cordial support of Professor Fernald and Mr.
Bayard Long, both of whom are well acquinted with it in the
field.
’ Carex Richii (Fernald) Mackenzie sp. nov.
Carex tenera Dewey var. major Olney, Exsicc. fasc. 2, No. 15
(in part). 1870.
Carex tenera var. Richit Fernald, Proc. Am. Acad. 37: 475.
1902
Carex hormathodes Fernald var. Richii Fernald, Rhodora 8: 166.
1906
Densely cespitose, the rootstocks very short-prolonged,
black, fibrillose, the culms 6-10 dm. high, erect, but very slender
to base, nodding, sharply triangular, very rough beneath head,
exceeding the leaves, dark brownish at base and clothed with
leaves of the year two to four to a culm, on lower fourth, but
widely separate, the lower nodes ten exposed, the blades flat,
LBP 2.5 mm. wide, usually 1-3 dm. long, long-attenuate, the
eaths green striate ventrally nearly to mouth, the ligule
eucroundtae the culm, short-prolonged beyond base of blades
spikes four to eight, strongly separate in a moniliform, flexuous
t
the pistillate part drbicular to ovoid-orbiculur, 6-12 mm. long,
6-9 mm. wide, rounded at apex, and with some twenty i ‘thirt rty
oe. or spreading-ascending perigynia with conspicuous
tips; r bracts setaceous, prolonged, the upper scale-like;
scales eel ie and osiieneis brown tinged, acuminate
or aristate from the s arp midvein, shorter and mu wi
than the perigynia; perigynia very flat and thin except where
distended by achene, membranceous, See or in age straw-
colored, 4-5 ie long, 2.5 mm. wide, t he body suborbicular,
and sessile at base, abruptly it ala hehe the flat, serrulate,
red-tipped, obliquely cut dorsally at length bidentate beak,
more than half lengt - y; ice lenticular, stipitate,
oblong-obovoid, 1.5 mm. long, 0.75 mm. wide, yellowish-brown,
apiculate; style slender, obweurel jointed with lalene: stigmas
two, reddish, slender, short
MACKENZIE: Notes ON CaREX—XII 363
SPECIMENS EXAMINED*
MASSACHUSETTS: Middleboro, Murdoch, June, 1900 (N.E.)
Canton, Churchill, June 17, 1900 (N.E.); Dover, Winslow,
June 21, 1912 (N.E.); Dedham, Forbes, June 10, 1912 (N.E.);
Stony Brook Reservation, Kidder, June 14, 1920 (N.E.); Sud-
bury, Rich et al. 99 (Type), June 17, 1902 (N.E.; FPN: Y..
K.M.); Bedford, Murdoch, June 7, 1902 (N.E.); Weston,
Weatherby, June 20, 1909 (N.E.); West Cambridge, Pease 3084,
May 27, 1903 (N.E.);South Sudbury, Miss Carr, June 30, 1897
(N.E.); Cambridge, Beal (H.), also Hubbard, June 2, 1896 (N.
E.), also Deane, June 8, 1887 (H.), also Fernald, June 10, 1891
(Col.); Stoneham, Rich, June 5, 1894 (N.E.); Reading, Perkins,
June 14, 1883 (N.E.), also Manning, June 8, 1882 (N.E.);
Arlington, Wm. Boott, June 5, 1870 (H.); Mystic Pond, Wm.
Boott. June 5, 1870 (H.); Amesbury, Eaton, June 1, 1903 (H.);
Sherborn, Loomis 1033, June 19, 1912 (K.M.).
RHODE IsLanp: Providence, Olney (N.Y.).
Connecticut: Waterbury, Blewitt, June 15, 1911 (N.E.),
also 1287, June 10, 1912 (N.E.); South Windsor, Woodward,
June 28, 1909 (N.E.); Columbia, Weatherby gogo, June 26,
1917 (N.E.); Thompson, Weatherby 4114, July 1, 1917 (N.E.);
Pomfret, June 5, 1871 (N.Y.); Newington, Wright, May, 1879
(H.); Sprague, Graves, June 13, 1902 (H.); Griswold, Graves,
June 16, 1899 (H.); Killingly, Bissell, June 22, 1902 (H.);
Thompson, Bissell, June 22, 1902 (H.); Oxford, Harger, June 5,
1904 (H.), also 4262, June 11, 1905 (P.); East Hartford, Weather-
by, May 25, 1903 (H.); Milford, Eames 5120, June 24, 1904
th M.; 3):
NEw York: Junius, Sartwell (H.).
NEw JerRSEY: Closter, Austin (H.); Newark, Livingston 35,
May 27, 1894 (H.); Crosswick’s Creek, Britton, May 29, 1904
(N. Y.); Passaic River, Bergen County (N.Y.); Oradell, Bergen
County, Mackenzie 759, June 19, 1904 (K.M.); Stirlington,
Morris County, Mackenzie 1267, May 28, 1905 (K.M.); Mon-
*Specimens are cited from herbaria as follows: Columbia University
(Col.); Gray Herbarium, Harvard University (H.); K. K. Mackenzie (K.M);
New England Botanical Club (N.E); New. York Botanical Garden (N.Y.);
New York State Museum (Albany); Philadelphia Academy of Natural
Sciences (P.).
364 MACKENZIE: NOTES ON CAREX—XII
mouth Junction, Mackenzie 4963, May 26, 1912 (K.M.); Succa-
sunna, Mackenzie 4128, June 13, 1909 (K.M.); Delanco, Burling-
ton County, Long 3474, May 19, 1910 (P.); Centre Square,
Brinton, May 20, 1892 (P.); eight miles southwest of Mickleton,
Heritage, May 25, 1892 (P.); Budd’s Lake, Morris County,
Heritage, May 28, 1895 (P.).
PENNSYLVANIA: Easton, Garber, 1868 (P.), also Porter,
June 5, 1890 (P.); Lehigh Gap, Lehigh County, Preiz 1831,
May 20, 1909 (P.); Slatington, Lehigh County, Pretz razé2,
June 6, 1920 (P.), also Bachman 2132, June 1, 1911 (P.); Strass-
burg, Lehigh County, Pretz 3355, May 20, 1911 (P.); Cementon,
Lehigh County, Pretz 10131, May 30, 1920 (P.); Rockhill,
MacElwee, May 31, 1903 (P.); Quakertown, Bucks County,
Martindale (P.); Sellersville, Prez, May 25, 1884, June 1, 1888,
and May 26, 1896 (P.), also Crawford, June 1, 1989 (P).; Penn
Valley, Bucks County, Crawford, June 1, 1899 (P.); Tully-
town, Crawford & Brown, 1902 (P.); Byberry, Martindale,
June, 1864 (P.); Wissahickon, A. H. Smith (P.); Sugartown,
Chester County, Pennell & Long 7628, June 26, 1912 (P.);
Nottingham, Chester County, Pennell & Long 7483, June 22,
1912 (P.); Mercersburg, Franklin County, ex herb. Porter, May
25, 1847 (P.); Wyoming Valley, Thurston, 1892 (P.); Elk
County, McMinn, 1868 (P.).
DELAWARE: Wilmington, poe June 30, 1896 (P.);
Vandyke, Newcastle County, Long May 24, 1909 (P.); Town-
send, Commons, May 17, 1883 (P.).
INDIANA: Palmyra, Harrison County, Deam 23417, June 4,
1917 (K.M., H).
2. A SEGREGATE FROM CAREX ALBOLUTESCENS
Among the varieties of Carex straminea Willd. recognized
by Professor Bailey was var. cumulata Bailey, based on speci-
mens collected in Pennsylvania by Porter, in New Brunswick
_by Fowler and on the Saskatchewan Plains by Macoun. This
variety, proposed in 1889, was maintained by Bailey in 1890
in the 6th Edition of Gray’s Manual (p. 622), where its dis-
tribution was given as from Pennsylvania to New England and
northward. It was said to be rare and to grow in dry grounds.
Three years later in 1893 Professor Bailey placed the plant
under Carex albolutescens Schw., as a variety, but added noth-
MACKENZIE: NoTEs ON CAREX—XII 365
~ ing to his previous publications concerning its distinguishing
characters or range; and this treatment is followed by Kiikenthal
in the Pflanzenreich (4%°: 209). By other authors Professor
Bailey’s name has been treated as a synonym of Carex albo-
lutescens.*
As shown however by the large number of specimens of
Bailey’s variety which have been collected in recent years, it
not only has an entirely different range from Carex albolutescens
but it has several constant and well-marked points of difference,
and is clearly entitled to specific rank. It is a northern species
ranging from Saskatchewan to New Brunswick and south to
Massachusetts, Connecticut and northern Pennsylvania. It
occurs in a great variety of habitats varying from wet to dry, but
seems able to thrive in much drier places than can Carex
albolutescens. In the southern part of its range it seems only
to be recorded from the higher elevations. It is not a coastal
plain plant like Carex albolutescens, but like that species is not
a species of limestone areas.
As above stated Carex albolutescens is typically a coastal
plant. It reaches its northern limit in eastern Massachusetts
but has a very extensive range to the south, being found as
far south as northern South America. In the more southern
part of its range it gets into the mountains.
The best technical character to separate the northern plant
from Carex albolutescens is that in it the perigynia are nerveless
ventrally, while in Carex albolutescens they are strongly several-
nerved. But in addition to this the spikes are numerous (five
to thirty) and densely aggregated; the green perigynia are more
conspicuous than in Carex albolutescens, thus giving the heads
a characteristic heavy, green, or in age brownish, appearance;
the sheaths are looser, and the leaf-blades of the sterile culms
are usually wider, being 4-5 mm. in width; in developed plants
the lateral spikes also are truncate-rounded at base.
Carex albolutescens has fewer spikes (three to ten), which
are less densely aggregated; and the heads have a very character-
istic silvery-green appearance; its sheaths are tighter than in
the northern plant and the leaf-blades of the sterile culms are
*The perigynium and the heavily spiked head, figured in the seventh
edition of Gray’s Manual (f. 351) as Carex albolutescens, represent Bailey’s
plant, and the head with five spikes represents genuine Carex albolutescens.
366 MACKENZIE: Notes ON CAREX—XII
normally narrower and but 2.5-3 mm. in width (although
occasionally more); and the lateral spikes are round-tapering
at base.
The northern plant is evidently abundantly distinct and it
should be treated as
‘Carex cumulata (Bailey) Mackenzie sp. nov.
Carex alata Torr. var. pulchra Olney (mostly), Exsicc. fasc. 2,
No. 23 (also Bailey Proc. Am. Acad. 22: 150 as synonym).
1871.
Carex straminea Willd. var. cumulata Bailey, Mem. Torrey
Club 1: 23. 1889.
Carex albolutescens Schw. var. cumulata Bailey, Bull. Torrey
Club 20: 422. 1893.
Very densely cespitose, the rootstock not prolonged, the
culms 3-9 dm. high, stiff, sharply triangular, roughened beneath
head, brownish at base and clothed with the dried-up leaves of
ed, the lower nodes sometimes expo lades
flat, light-green, thickish, erect or ascending, 3-5 mm. wide, -
7-25 cm. long, roughen ed towards apex, the sheaths loose,
strongly green striate easel and conspicuously prolonged be-
yond base of blade and continuous with the ligule; sterile shoots
elongat e cig usually 4-5 mm. wide, occasi hate nar-
rower, erect, the s s very loose; alas five to thirty
candrous, very coeele aggregated into an ovoid or oblong hed: a
cm. long, 1-2 cm. thick, the spikes ovoid, 6-10 mm. long, 5-8
mm. thick, short-pointed at apex, the lateral truncate or rounded
at base, the terminal clavate, — Bas ring or in age
brownish, closely many-flower ia ascending with
ot
oa
popepicucs: ae ovate, obtuse or acute, much narrower and
shorter e perigynia, white-hyaline with one- to three-
nerv een center ; peri in, pia es hse wea m
te middle, ct ea me ight babar a dorsally,
> . 2 ¥3
obovoid, at maturity yellowish brown, very short substipitate,
2 mm. long, 1.25 mm, wide, bent-apiculate; style slender,
MACKENZIE: NoTES ON CAREX—XII_ 367
jointed with achene, at length deciduous; stigmas two, slender,
light-reddish, short
SPECIMENS EXAMINED
New Brunswick: Bass River, Fowler, August 9, 1872 (H,
N.Y.); Miramichi, Fowler, August 24, 1894 (H.), also Williams
& Fernald, July 25, 1902 (H.); Kent County, Fowler (Type),
July 1870 (H., N.Y., Col. Albany, P.).
Nova SCOTIA: Bell’ s Island, Lahave River, Macoun 82518,
August 10, 1910 (H.); Bridgewater, Macoun 825109, July 18,
1910 (H.); Springhill Junction, Pease & Long 20300, July 18,
1920 (K.M.); Port Mouton, Fernald & Graves 20312, August 18,
1920 (K.M.); Middleton, Fernald et al. 20325, July 20, 1920 (K.
M.); Broad River, Fernald & Bissell 20311, August 16, 1920
(K.M.).
MAINE: Somesville, Fernald, September 22, 1892 (N.E.,H.);
Bangor, Knight, August 7, 1904 (H.); Pembroke, Fernald 14309,
July 17, 1909, (N.E., H.); Southport, Fernald, August 1 and 2,
1894, (N.E., H); Kennebunk, Parlin 867, August 26, 1897
(H., N.E.); Mt. Desert Island, Greenleaf, August 3, 1883 (N.E.),
also Rand, Redfield, Faxon, Fernald, Williams, numerous col-
lections 1890-1898 (H., N.E., Col., N.Y.); Milford, Fernald &
Long 12928, August 25, 1916 (N.E.); Lovell, Pease 17307,
August 30, 1918 (N.E.); Matinicus, C. A. E. Long, August 28,
1919 (N. E.); summit of Mt. Battie, 1325 feet, Furbish, July 14,
1903 (N.E.), also Bissell, August 13, 1913 (N.E.); Durham,
Dinsmore, August, 1898 (N.E.); Bear Mountain, Livermore,
Furbish, July 24 and 25, 1896 (N.E.); Kennebunk, Deane,
July 5, 1894 (N.E.); Berwick, Parlin 1a62, July 24, 1898 (N.E.);
North Berwick, Fernald, July, 1897 (N.E.).
New Hampsuire: Nottingham, Eaton, June 20, 1900 (H.);
Mt. Monadnock, 3000 ft., ace July 3, 1899 (N.E.), also “A.
W. D.,”” August 7, 1916 (N.
MAssacauserts: Mt. ee Ashby, Knowlton, August
9, 1908 (H.); Medford, W. Boott, June 30, 1861, (N.E.); July
25, 1863 (H.); Edgewith, W. Boott, July, 1853 (H.); Ashland,
Morong, July 10, 1879 (H.); West Manchester, Hubbard 195,
October 11, 1911 (N.E.); Middlesex Falls, Kidder, June 28,
1920 (N.E.); Wilmington, Collins, June 25, 1887 (N.E.); Mt.
Washington, Berkshire County, Hoffman, August 23, 1914
368 ’ Mackenzie: NoTEs oN CAREX—XII
(N.E.); Lake Undine Dome, Berkshire county, Hoffman, August
27, 1902 (N. E.).
ConneEcTiIcuT: East Windsor Hill, ‘A. W. D.”’, August 26,
1915 (N.E.); South Windsor, Bissell, July 30, 1916 (N.E.); also
Blewitt 1502, July 10, 1912 (N.E.), also 3470, August 8, 1917
2 ee 2
New York: Taghkanick Range, Hoysradt, July-August
(H.); Karner, Albany County, House, 6410, July 7, 1919 (Albany) ;
Whitestown, Oneida County, Haberer 3760, August 22, 1912
(Albany); Ft. Ann, Washington County, Burnham 4g, June 19,
1892 (N.Y.); Sams Point, alt. 2600 ft., Rusby, August 20, 1896
(Col.); Mt. Riga, Hoysradt, July 17, 1878 (Col.).
PENNSYLVANIA: Naomi Pines, Porter, July 1, 1893 (P.),
also Britton, July 2, 1893 (Col.); Pocono Plateau, T. Greene,
September 4, 1861 (P.), also Porter, September 11, 1869 (P.),
July 25, 1881 (P.), August 6, 1891 (Col., H.), and July 1, 1893 (P.).
Micuican: Alger, Wheeler, July 2, 1895 (H.); Omer, Arenac
County, Wheeler, July 24, 1902 (H.); Duck Lake, Arenac
County, Wheeler, t900 (H.);Township 18, n. 4 E., Section 24,
Wheeler, August 8, 1900 (H.); Township 19, n. 6 E., Section 16,
Wheeler, August 9, 1900 (H.); Emerson, Chippewa County,
Dodge, August 28 and September 4, 1914, (K.M.); Manistique,
Schoolcraft County, Dodge, September 22, 1915 (K.M.); Alpena,
Wheeler, July 8, 1895 (H., Col.).
Ontario: Lake of the Woods, Macoun 33, July 28, 1872
7h).
SASKATCHEWAN: Fort Ellice, Macoun 219, 1872 (H.).
3. A NORTHERN RELATIVE OF CAREX BREVIOR
In the northern part of the United States from Maine to
Michigan and in southern Canada from Ontario to Manitoba,
there is found a large, robust and handsome member of the
Ovales. It has five to ten spikes, which normally are well sep-
arated, but plants with approximate or subapproximate spikes
are not uncommon. It is the plant treated by Professor Fernald
as Carex festucacea Scik. (Proc. Am. Acad. 37: 477. pl. 3, f.
47-48), and also so treated in Gray’s Manual (7th ed., p. 221. f.
363). To it is added in the treatments referred to a variety
brevior (Dewey) Fernald, distinguished by having but three to
six approximate or subapproximate spikes.
Mackenzie: Notes on CarEXx—XII 369
When I prepared the article on Carex for the second edition
of Britton & Brown’s Illustrated Flora, I treated these two
plants together, because the number of heads and their degree
of separation was not a constant character. Later, in a paper on
Carex straminea and its allies (Bull. Torrey Club 42: 603. 1916),
it was pointed out that the genuine Carex festucacea was a differ-
ent plant than either of the above, and the name Carex brevior
(Dewey) Mackenzie was applied to what I had treated as Carex
festucacea Schk. in the Illustrated Flora.
In the meantime Kiikenthal (Pflanzenreich 4°: 207), notic-
ing the close general resemblance of Fernald’s Carex festucacea
to Carex Bicknellit Britton (Carex straminea var. Crawei
Boott), cited it under his treatment of that plant.
A detailed study has disclosed that the large northern plant
here under discussion has characters which are constant in a
very considerable series of specimens. As compared with Carex
brevior it has the perigynia appressed-erect instead of spreading-
ascending. The mature perigynia are membranaceous and
strongly pellucid-puncticulate. They entirely lack the firm
coriaceous texture so characteristic of Carex brevior and are
also more nerved than in that species. These differences are
well brought out in Fernald’s figures above referred to, even
the difference in the texture of the perigynia coming out.
The culms of the northern plant are much developed the
first year of their growth, and the old dried leaves towards the
base of the culm are therefore very marked in the flowering year.
In Carex brevior on the other hand the development the first
year is much less, and the dried leaves are short and rather
inconspicuous in the flowering year. The sheaths too in the
northern plant are white-hyaline ventrally for but a short
distance, while in Carex brevior this feature is very strongly
developed, and they also differ from those of the latter plant in
being strongly papillate dorsally. As stated by Fernald the
northern plant has five to ten spikes which normally are distinct
and well separate, and Carex brevior usually has fewer spikes
which are approximate or sub-approximate. However, this is
but a well-marked tendency and specimens of the northern
plant with a congested head or Carex brevior with spikes as
widely separated as in the northern plant are not uncommon.
Carex Bicknellii can be readily distinguished by its larger
370 Mackenzie: Notes oN CarEx—XIi
(5.5-7-5 mm. long), nearly translucent perigynia, which are
strongly about ten-nerved ventrally, and very strongly winged.
It is, 1 would say, more closely related to the northern plant
than is Carex brevior.
A real understanding of the members of the Ovales found
in the northeastern part of North America and the characters
which separate them dates from Professor Fernald’s revision
of the group in 1902. The plant which I have been here dis-
cussing is one which he has collected on a number of occasions.
It is therefore one to which it is highly fitting to give his name.
Unfortunately there is a poorly understood Japanese species
to which his name has been given. Under the circumstances I
trust that I will be pardoned in proposing for the plant I have
been discussing a longer name than would ordinarily be de-
sirable.
Carex Merritt-Fernaldii Mackenzie sp. nov.
“Carex festucacea Schkuhr’’ Fernald, Proc. Am. Acad. 37: 477.
pl. 3, f. 47-8. 1902.
Cespitose from short-prolonged, lignescent, black-fibrillose
rootstocks, the culms 3-10 dm. high, slender but stiff, sharply
triangular above, terete towards base, strongly bi sa diceedi ng
the leaves, smooth or very slightly di oe beneath head,
conspicuously clothed at base with the s -bladed Aeaves of
the previous year, the lower bladeless; Coe git well-developed
blades three to six to a culm, on the lower third but not bunched,
the blades erect-ascending, dull- or yellowish-green, firm, 1.5—-3
mm. wide, usually 1-2 dm. long, strongly roughened towards
ventrally for a short distance below mouth only, strongly
papillate dorsally, extending up beyond point of insertion of
continuous with ligule, the latter longer than wide;
spikes four to ten (usually six to eight), aggregated to separate,
the head 1.5-8 cm. long, 7-15 mm. wide, the spikes gynaecan-
drous, subglobose to ovoid, 7-15 mm. long, 5-9 mm. wide, with
posi to thirty a ppressed-ascending g perigynia above (the beaks
appressed-ascending), blunt at apex, abruptly contracted into
the short clavate basal staminate portion; bracts scale-like, the
lowest often prolonged, 1-4 cm. long, the upper merely acuminate
or short-awned ; scales ovate, obtuse to short-cuspidate, yellow-
ish brown with three-nerved green center and narrow hyaline
margins, shorter and much oncggird ‘goat the perigynia above;
perigynia suborbicular, 4-5 mm 2.5-3.5 mm. wide, con-
cave-convex, thickish over the aig but the walls membran-
MACKENZIE: NoTEs ON CaREX—XII 371
aceous, strongly pellucid-puncticulate, yellowish-green or in age
yellowish-white, strongly winged to base, serrulate to below
middle, truncate rounded at base, strongly several to many
nerved dorsally, nerveless to three to five-nerved ventrally,
often only towards base, the wings one- to two-nerved, abruptly
narrowed into the flat, serrulate, reddish-brown tipped, obliquely
cut dorsally, bidentate beak, about 1 mm. long and less than
one-third length of body, the margins of the orifice reddish-
brown; achenes lenticular, 1.75 mm. long, 1.5 mm. wide, sub-
stipitate, apiculate, brown, shining; style slender, straight,
jointed with achene, deciduous; stigmas two, reddish rown,
slender, long.
TYPE LOCALITY: Orono, Maine, Fernald, July 3, 1897
(TyPE in the Gray Herbarium of Harvard University).
SPECIMENS EXAMINED
MAINE: Seal Harbor, Redfield, July 14, 1890 (H., N.E., P.),
also Williams July 25, 1890 (N.E); Canton, Parlin 2021, August
8, 1906 (H.);. North Berwick, Parlin 798, July 15, 1897, also
May 1892 (H.); Mt. Desert Island, Rand, July 9, 1893, also
July 15, 1894 (H.); Basin Mills, Knight, July 15, 1904 (H.);
Orono, Fernald, June 1, 1899 (N.E.), July 14, 1890 (N.E.),
July 12, 1897 (N.E.), July 2 and 3, 1897 (H.); Bumford Point,
Parlin, July 4, 1890 (H.); Canton, Parlin 1972, July 10, 1906
(H.), also 2275, 1907 (H.); Tatnic, Parlin, July 11, 1891, also
June 1892, (N. E., H.); Orono, Fernald & Long 12953, July 22,
1916; (N.E., P.); Milford, Fernald & Long 12955, August 25,
1916 (N.E., P.); Farmington, Knowlton, July 9, 1911 (N.E., P.);
Woodstock, Pease 17126, July 11, 1918 (N.E.); Canton, Parlin
2094, June 25, 1907 (N.E.); Dixfield, Parlin 2461, June 28, 1908
(N.E.); Gilead, Pease 16013, July 5, 1914 (N.E.); Rumford
Point, Parlin, July 4, 1890 (N.E.); Dennysville, Fernald 1444,
July 19, 1909 (N.E.); Columbia, Knowlton, August 4, 1916
(N.E., P.); West Pembroke, Fernald 1452, July 6, 1909 (N.E.);
Big Hill, Fernald 1438, July 17, 1909 (N.E.); Leeds, Knowlton,
July 23, 1916, (N.E., P.); New Gloucester, Knowlton, July 25,
1920 (N.E.); Baldwin, Fernald et al. 12956, August 30, 1916
(N.E., P.); Kennebunkport, Deane, July 12, 1894 (N. E.);
North Berwick, Parlin 798, July 15, 1897 (N.E.); Alfred,
Fernald & Long, 12054, August 10, 1916 (N.E., P.); Wells,
Fernald & Long 12952, August 8, 1916 (N.E,, P.); North Ber-
372 Mackenzie: Notes oN CAREX—XII
wick, Parlin & Fernald, June 13-14, 1896 (N.E.); Tatnic Hill,
Parlin & Fernald, June 13-14, 1896 (N.E.); Cape Porpoise,
Furbish, July, 1901 (N.E.); South Berwick, Parlin, July 31, 1896
(N.E.); Hermon, Knight 5035, June 30, 1906 (K.M.); Veazie,
Knight, July 15, 1905 (K.M
New HAMPSHIRE: Soon Mand & Robinson 507, June 13,
1898 (H.); Shelburne, Deane, July 8, 1914, also July 30, 1914
(N.E., P.); Plymouth, Fernald, 11589, July 30, 1915 (N.E.);
Durham, Knowlton, July 1, 1917 (P.).
VERMONT: Mason Hill, Pownal, Churchill, June 18, 1901
(N.E).
MassacHusetTts: Amherst, B. P. Clark, June 6, 1881 (P.).
New York: Wellesley Island, Jefferson County, Robinson
& Maxon 1, June 28, 1902 (H.); Thousand Island Park, Ball,
July 1904 (K.M.); Middle Grove, Saratoga County, Peck
(Albany) ; East Galway, Saratoga County, E. A. Burt (Albany) ;
West Ft. Ann, Washington County, Burnham, June 27, 1918
(Albany).
OntTaRIO: Lake Nipigon, John Macoun, July 11, 1884 (H.);
Kingston, Klugh, June 30, 1906 (H.); ‘‘Portage River, Lake
Superior”’, ex herb. C. E. Smith, August 3, 1865 (P.).
MICHIGAN: Washington, ex herb. Dewey; Herb. (H.); Kew-
enaw County, Farwell 762, August 1890(H.); Muskegon, Mc-
Clatchie 48 (H.); Douglas Lake, Emmet County, Ehlers, 296,
299, July 17, 1920, also L. E. Smith 1, July 3, 1917 and 159,
Aug. 20, 1917(K.M.); Clifton, Farwell, July 18, 1890 (Albany);
Isle Royale Williamson, 2276 August 15, 1912 (P.).
MANITOBA: Lake Winnipeg Valley, Bourgeau 1857 (H.).
oe ee ee ee
ADDENDUM
After the above was written, and when on a visit to the Phil-
adelphia Academy of Sciences, Mr. Bayard Long asked me to
look at the type specimen of Carex albolutescens Schw. preserved
there. It was apparent that it was a young and immature
specimen of the genuine Carex siraminea Schk. as interpreted by
me (Bull. Torrey Club 42: 605-6), aconclusion which Mr. Long
had himself arrived at. The previous identification of the plant
MACKENZIE: Notes oN CAREX—XII 374
of Schweinitz with the plant treated in this article and generally
as Carex albolutescens can be readily understood. The two
species really stand next to each other, and as long as genuine
Carex straminea was not understood and kept separate, it was
a natural thing to do to identify the plant of Schweinitz in the
way that was done. The name, however, has always been
inappropriate, and one is glad to be able to reduce it to synonymy.
t now appears that the plant treated in this article and gen-
erally as Carex albolutescens (‘‘ Carex albolutescens Schw.”’ Kiik-
enth. in Engler, Pflanzenreich 42°: 208; Mackenzie in Britton
& Brown, Ill. Fl. (2d.. ed.) 1: 385. f. g4z [excluding var. cumulata
Bailey]; Robinson & Fernald in A. Gray Man. (7th. ed.) 219.
pl. 351 [left hand] excluding var. cumulata Bailey; ‘‘ Carex foenea
Willd.’’ Boott, Ill. Carex 3: 118, pl. 375) is withoutaname. It
therefore gives me much pleasure to be able to name it for that
excellent field man and acute student, Mr. Bayard Long, as Carex
Longii. A specimen collected July 24, 1907, by Mr. Long at Cold
Spring, Cape May County, New Jersey, and preserved in the
herbarium of the Philadelphia Academy of Natural Sciences is
designated as the type. _
Stimulation of plants by carbon disulphide
CARRIE OLDENBUSCH
The problem of stimulation of plants by minute doses of
chemicals, which are poisonous in higher concentrations, has
long been an interesting one. Raulin (1) in 1869 was probably
the first to undertake work of this kind, using a nutrient solution
made up of a large number of substances as a culture medium.
As the stimulant, Raulin added to his nutrient solution small
amounts of metallic salts, such as silver nitrate, platinum di-
chloride, or copper sulphate, and found that cultures of Asper-
gillus niger, grown in these solutions produced a more luxuriant
mycelium than cultures to which nothing had been added.
Pfeffer (2) in 1895, by means of comprehensive experiments, de-
termined that such a complicated nutrient solution was unnec-
essary and originated a number of solutions which have been
and are still extensively used in work of this kind. Benecke
(3), in the same year, published a paper in which he also gave a
much simpler normal solution than Raulin’s but slightly different
from Pfeffer’s.
Richards (4), using the latter’s solution with the addition
of traces of zinc, iron, sodium, lithium, and other salts, obtained
a heavier growth of Aspergillus niger and Penicillium glaucum
than in cultures to which none of the above were added, demon-
strating that Raulin’s solutions really resulted in stimulation.
Ono in 1900 corroborated Richards’s results and also worked
out the effect of these salts on certain algae. He found that
algae as a rule have their point of greatest stimulation at a much
lower concentration of the salt but in other respects are similar
to fungi. Richards (5) and Ono (6) determined that the econ-
omic coefficient of sugar is considerably lower in the cultures
to which salts had been added than in the controls, proving that
stimulated fungi require a smaller amount of carbohydrate
food in proportion to the quantity of waste acid produced.
Since this time many other investigators have worked on
this problem, obtaining similar results. Kahlenberg and True
(7) formulated a law which they gave as an explanation of this
stimulation phenomenon. They claim that since the chemical
and physical properties of solutions are due to the properties of
375
376 OLDENBUSCH: STIMULATION OF PLANTS
the ions dissociated in the solution and of the undissociated
molecules, then the physiological properties of a solution are due
to the same factors. The authors worked out this theory with
lupine seeds, the germination of which was accelerated by dilute
solutions of electrolytic salts. They claimed that this stimu-
lation was due to the hydrogen and metallic ions in the solutions.
Heald (8) working with seeds of other plants obtained similar
results and also attributed them to the electrolytic dissociation.
Stevens (9) working with fungus spores applies this same theory.
Iwanoff (10) claims that metallic salts are poisonous to plants in
proportion to the atomic weight of the metal, the heavier metals
being more poisonous than those of smaller atomic weight. In
his opinion, the same is true of the alcohols, the lower alcohols
(methyl and ethyl) being less poisonous than those of higher
molecular content, such as propyl and butyl. Steinberg (11),
in a recent paper on the stimulation of Aspergillus, attributes the
stimulation to the increase in H ion content when zinc salts are
added to the nutrient solution. It is not obvious, however,
that the action of such oligodynamic substances is due to the
hydrogen ion concentration alone, especially since we find non-
electrolytes, such as ether, chloroform, and other anaesthetics,
exerting a stimulatory influence. Carbon disulphide, being a
very volatile liquid, which is non-dissociable in aqueous solution,
~ also comes in this category.
Various authors have worked with ether and chloroform,
although few have done anything with carbon disulphide. Town-
send (12) in his experiments with Avena seeds found that if
dormant seeds were exposed to a moist concentrated atmosphere
of ether for a short time, or to a weaker atmosphere of ether for a
longer period, the seeds germinated sooner and grew more rapidly
than under normal conditions. This acceleration lasts for some
time and gradually disappears. Ina later paper (13) Townsend
verified his results with other seeds: Zea Mays, Cucurbita,
Phaseolus, etc. He also found that dry seeds, exposed to an
atmosphere of HCN gas (14), germinated more rapidly than
those not exposed although this advantage did not last for any
appreciable time. Miss Latham (15), working with Sterigmato-
cystis nigra, determined that cultures of the fungus grown on
nutrient media and exposed to chloroform vapor, produced a
more luxuriant growth with a greater dry weight than similar
OLDENBUSCH: STIMULATION OF PLANTS a75
cultures unexposed. Furthermore the oxalic acid formation was
lower per unit dry weight in the stimulated cultures.
Johannsen (16) was the first to discover that exposing dor-
mant buds to an atmosphere impregnated with ether forced the
buds to open earlier than normal. Chloroform has the same
effect but is more powerful, less being needed to bring about the
same result.
No attempt has been made to cover the whole field of the
very voluminous literature on this subject. A comprehensive
account may be found in Czapek’s ‘‘ Biochemie der Pflanzen,”
second edition, Vol. 1, page 147 et seq., 1913.
Among the investigators who experimented with carbon
disulphide, none used it in gaseous form but either in pure liquid
state or in aqueous solution. Sirker (17) in Japan found that
mulberry bushes, grown in soil previously treated with CS., pro-
duced more branches, higher plants and more leaves with a
greater dry weight for each individual leaf than bushes grown in
untreated soil. Koch (18) in 1912 found the same thing with
other plants. Koch asserts that the carbon disulphide is not
used as a carbon source, nor does it act as an insecticide in
killing off the harmful lower organisms in the soil, since under
such circumstances a larger amount of the CS. would be more
beneficial. This, however, is not the case, a larger amount of
the CS. being harmful, and the stimulation takes place only when
the compound is added in minute quantities. Fred (19) in the
same year stated that carbon disulphide added to the soil in
dilute solutions stimulated the growth of soil bacteria, the nitro-
gen-fixing forms in particular, so that the quantity of nitrogen
built up into higher compounds was measurably increased over
normal. In a later paper Fred (21) corroborated his earlier
results and concluded that while after treatment with carbon
disulphide the soil showed a reduction in the number of micro-
organisms, it was followed by a great increase and an increase
also in the by-products of their action. With relatively strong
applications of carbondisulphide to the soil, corn and oats
seemed deleteriously affected, while mustard and buckwheat
were benefitted, as shown by increase in dry weight. In acid
soil, clover is also stimulated.
e experiments described below, which further test the
effect of carbon disulphide on plants, were performed in the
378 OLDENBUSCH: STIMULATION OF PLANTS
Botanical Laboratories of Barnard College, under the super-
vision of Professor H. M. Richards, for whose advice and assis-
tance the writer is greatly indebted.
- In this work, while no attempt was made to cover every type
or condition of plant tissues, a number of widely differing kinds
of material were employed. The experiments fall into three
groups, each of which could profitably be elaborated beyond
what was attempted here. These three categories are as given
below and will be considered as separate topics in the following
discussion.
1. Stimulation of seeds and seedlings.
2. Stimulation of buds
3. Stimulation of certain fungi.
Carbon disulphide is a clear, practically colorless liquid when
pure, very inflammable and volatile. It has a disagreeable odor,
especially when impure, because of the decomposition into other
sulphur compounds, hydrogen sulphide in particular (see Graham,
20). It is soluble in all proportions in alcohol, ether, and chlo-
roform, but only slightly soluble in water. The solubility in
water, according to Graham, varies inversely as the temperature,
as follows:
At iz -13° €. 2.0 gms. CS dissolve a I liter ot water.
I
27° “ce 168 “ec 6é “cc oc I ce “c “ce
30° 33° “ 1.45 “c “ Toe se c tc
From these data, it was estimated by interpolation that at
20°-22° C. (average room temperature) about 1.8 gms. CS:
dissolves in 1 liter of water.
In the following paper, the concentrations of the solutions
used were determined in terms of molecular solutions. The
molecular weight of carbon disulphide is 76.125, and if 1.8 grams
dissolve in 1 liter of water at approximately room temperature,
then a saturated aqueous solution is 0.0235 of a molecular solu-
tion. More dilute solutions were prepared from this, in fractions
varying by 1/2000M. No attempt was made to determine or
maintain the exact concentration closer than this, because carbon
disulphide is so easily vaporized and a certain amount of that in
solution is continually volatilizing out. In cases where the ex-
periments lasted over a few days, the pean were renewed at
intervals of two or three days.
OLDENBUSCH: STIMULATION OF PLANTS 379
I. STIMULATION OF SEEDS AND SEEDLINGS
The first phase taken up was to determine if there was any
stimulation of young seedlings by dilute solutions of carbon
disulphide. Townsend (12, 13, 14) tested the effect of ether
vapor and hydrogen cyanide gas on different kinds of seeds, by
soaking seeds in water for twenty-four hours and then putting in
an airtight dampchamber containing ether vapor. The exposed
seeds germinated more quickly than unexposed controls, but
whether left in ether chamber or removed immediately after
germination, the stimulated seedlings soon lost their advantage,
and five days after germination the control seeds equaled the
stimulated in growth. Likewise dry seeds exposed to HCN gas
for a short time and then soaked in water germinated more
rapidly than seeds not treated with the hydrogen cyanide.
However in every case, growth was equal after a week.
In the following experiments, wheat grains (Triticum sativum)
were used. One hundred grains were soaked in the CS; solution
for twenty-four hours. Seed pans were filled with moist sphag-
num, covered with damp cheese cloth, and grains planted out on
top of this. They were then covered with a second layer of
cheese cloth, with a pad of moist sphagnum on top of the whole.
Pans were kept under a shelf in the greenhouse for three or four
days, taking care not to allow the sphagnum to dry out. The
seedlings were then uncovered, epicotyls measured and the
average taken of those germinated. It was found that the
saturated solution completely inhibited growth, while the seed-
lings in the other solutions grew as shown in TABLE I.
TABLE I
AVERAGE GROWTH IN MILLIMETERS OF EPICOTYLS OF WHEAT SEEDLINGS AFTER
BEING SOAKED IN FOLLOWING CS: SOLUTIONS FOR 24 HOURS
EFT TO GERMINATE FOR 3 OR 4 D
Series 1. Germinated 4 days
B
Exp. A. . be Average Avg. per day
M/2,000 37-7 37-7 9.4
M/4,000 39.7 35-2 38.5 37-8 9.4
M/6,000 40.6 29.2 34-9 8.7
M/8,000 34.8 31.1 41.3 39-1 9.8
M/12,000 38.3 34.1 36.2 9.1
M/16,000 42.7 33-2 37-6 37.8 9.5
M/22,000 41.2 28.9 43-4 37:8 9-5
M/32,000 40.0 36.6 39.4 38.7 9.7
M/40,000 41.5 41.5 10.4
380 OLDENBUSCH: STIMULATION OF PLANTS
TABLE I (con:)
Serizs IJ. Germinated 3 days
Exp. D. E. Average Avg. per day
M/2,000 13.6 10.0 11.8 3.9
M/4,000 13.3 13.8 13.6 4.5
M/8,000 16.8 14.5 15.7 5.2
M/16,000 14.7 T27 14.2 4.7
M/22,000 14.9 14.2 14.6 4.9
M/32,000 18.7 14.4 16.6 5.5
M/40,000 12.7 15.3 14.0 4.7
Control II.1 13.6 13.3 4.4
Although these results show some irregularities due to the
comparatively small number of cases, they also reveal a marked
stimulation of growth especially in the solutions between
M/22,000 and M/40,000 in CS, content. The stimulation in
the three day experiments is proportionally greater than in the
four day instances, showing that the stimulation tends to
diminish, and growth tends to approach the normal after an
interval, just as Townsend found.
An experiment with Vicia Faba seedlings was carried out in
the following manner. Theseeds were planted and allowed to
germinate until the hypocotyls reached a convenient length to
measure. A number 400 cc. bottles were almost filled with
Knop’s nutrient solution and enough CS, solution was added to
make of desired concentration. Thebottles were then fitted with
corks, each cork having three holes bored in it, which were just
small enough to prevent the seeds from falling through. The
seedlings grown as above, were then washed in distilled water,
hypocotylsm easured, and arranged in culture bottles so that
seed rested on top of cork and hypocotyl extended through the
hole into the solution. The bottles were placed in a dark
chamber in the green house (temperature varying from 18° to
25° C.), the hypocotyls were measured every twenty-four hours
and the daily growth computed.
OLDENBUSCH: STIMULATION OF PLANTS 381
TABLE II
AVERAGE GROWTH IN MILLIMETERS OF THREE HYPOCOTYLS OF Vicia Faba
SEE ee, PLACED IN NUTRIENT SOLUTIONS CONTAINING TO
F FOLLOWING CONCENTRATION, EACH EXPERIMENT
COMPRISING TWELVE SEEDLINGS
M/2,000 M/4,000 M/6,000 M/8,000 M/20,000 M/40,000 Control
Exp. A
Ist day 58 6.5 13.0 13.7 7.0
2nd 2:2 5.4 11.7 16.1 9.5
Total 74 11.9 24.7 29.8 16.5
Vv! 3.8 6.0 12.4 14.9 8.3
Exp. B
Ist day 18.2 18.2 22.7 22.9 17.8 18.9
and,“ 19.4. 23.4 21.6 21.0 16.2 19.6
ad 21.4 19.0 19:7" 15.8 8.1 14.4
4th “ 19.2 19.2 23.5 17.0 17.7 16.3
Total 78.2 79.8 87.5 76.7 59.8 69.2
Avg 19.6 20.0 21.9 19.2 15.0 1.4
Bap ©
Istday 16.6 14.6 13.6 73.7. 147 16.6 19.3
Hid: 10.5 16.0 18.7 21.8 21.6 16.0 17.5
grat 14.1 18.4 18.0 29.0 26.7 15.5 19.7
Total 41.2 49-0 50.3 64.5 63.0 48.1 56.5
vg £3.77 16.3 16.8 21.5 21.0 16.0 18.8
Exp. D
Ist day 9.5 16.8 10.8 20.0 39.0 1.5 19.8
2nd ‘ 7.0 11.5 II.0 18.0 12.5 11.0 16.5
aa: * 8.0 14.0 12.9 24.5 18.5 13.2 17.0
Total 24.5 42.3 34.7 62.5 70.0 25.7 53-3
Avg. 8.2 14.1 11.6 20.8 23.3 6 17.8
Total average of the four experiments (i.e. twelve examples).
19.6 13.2 15.6
TABLE II shows a distinct stimulation in the growth of
hypocotyls of Vicia seedlings, in the M/6,o00 to M/20,000 CS,
solutions, the average greatest stimulation being in the M/8,000
dilution, although the M/20,000 shows a greater acceleration in
some instances. In the M/40,000 concentration, the CS, content
has become so dilute that it no longer has any effect on growth.
In the higher concentrations, the M/6,o00 solution caused a
marked stimulation in Exp. A and B, stained in other cases
382 OLDENBUSCH: STIMULATION OF PLANTS
growth is not as great as normal in these dilutions. Although
the M/4,000 and M/2,000 solutions do not entirely inhibit
growth, they do retard it below normal. Complete inhibition
was found in the saturated CS, solution, but no dilutions were
tried between that and the above.
Koch (18) tried similar experiments with Helianthus annuus
and Cucurbita Pepo seeds, letting the seeds germinate and then
placing them inthe dark, with their hypocotyls in solutions
containing 0.1 and 0.05 cc. CS, to 1000 cc. water for thirteen
to fourteen hours. In one case only, in the solution containing
0.05 cc. CS2 to the liter(about M/150 solution), did he obtain
growth higher than normal. This being only a single instance
was probably an accident due to the solution being weaker
than intended and cannot in any sense be called a stimulation.
Koch did not try any solutions weaker than this and therefore
did not reach the stimulatory region but really only determined
that carbon disulphide acted as an inhibitory agent.
2. STIMULATION OF DORMANT TWIGS
Johannsen (16), investigating the effect of ether on dormant
twigs, found that they were stimulated so that the buds opened
sooner and developed more rapidly when exposed to ether vapor
for twenty-four to forty-eight hours, than when unexposed.
When using chloroform a much smaller amount was_neces-
sary to produce the same result (9 grams of chloroform giv-
ing the same acceleration as 40 grams of ether).
With these results in view, experiments were carried out to
see if vaporized carbon disulphide would stimulate buds to open.
Willow twigs were cut from trees outdoors which had been
subjected to winter conditions and were probably frozen in
spite of the general mildness of the winter. The lower three or
four inches of the stem were placed in a bottle of water and a
large bell-jar put over them. Pieces of filter paper, moistened
with 10 cc. of carbon disulphide were quickly inserted under the
bell-jar. The whole was placed in a pan of water making a
water seal so that the carbon disulphide, which vaporizes almost
immediately, could not escape from the bell-jar. The twigs
left in this atmosphere (approximately saturated with CS;
gas) for periods ranging from ten minutes to eighteen hours.
They were then removed, the stems were put into fresh water
and with a control placed in the greenhouse.
OLDENBUSCH: STIMULATION OF PLANTS 383
The results of a number of experiments can be summarized
as follows. After three days, the twigs exposed ten and twenty
minutes began to develop, the buds showing patches of green
at the tips. After seven days, the buds on the twigs exposed
thirty minutes were open considerably. One of the hour experi-
' ments showed slight greening of the upper buds but other speci-
mens gave no evidence of development. After nine days, the
control (not exposed to CS, at all) had two buds developing.
Twigs exposed ten, twenty and thirty minutes continued to show
better bud development for two to three weeks longer, after which
the condition of all tended to become the same. Twigs exposed
for one and a half hours or for two hours started to open but
after a day or so became brown and withered, while those
exposed for four hours or more showed no signs of development
at any time, indicating that the protoplasm itself must have
been killed.
3. STIMULATION OF CERTAIN FUNGI
As stated above, there has been a vast amount of work done
on stimulation of fungi by metallic salts and other chemicals.
Because of this, a number of experiments were set up to test the
effect of carbon disulphide on fungi.
To determine the effect of dilute solutions of carbon disulphide
on the germination of spores of Rhizopus nigricans and Sterig-
matocystis nigra, hanging drop slides were used. Small glass
rings were attached to microscope slides by means of a solution
of dextrin in water (vaseline is not practicable since carbon
disulphide is very soluble init). The medium used for the ger-
mination of the spores was prepared according to the following
formula (see Richards, 4
Potassium phosphate : 0.5 gm.
Magnesium ‘oulghate 0.25 gm
Ammonium nitrate 1.0 gm
Ferrous sulphate trace
Sugar 5.0 gms
Distilled water. ; 100 cc.
To this was added 5 gms. of gelatine, and the mixture was
then heated in water bath. When the gelatine was completely
solved, the medium was filtered tubed (10 cc. to the tube) and
sterilized. Before the gelatine eon the CS, solution was
added to make of desired concentrati
384° OLDENBUSCH: STIMULATION OF PLANTS
A small drop of this gelatine was put on a clean cover glass
and inoculated with fungus spores. After inverting the cover
glass, and sealing on to the glass ring the slides were left standing
at room temperature under a bell-jar containing moist filter
paper to prevent slides from drying up. After some hours,
slides were examined microscopically for germination, and where
possible the germination-tubes were measured by means of a
standardized micrometer eye-piece.
According to TaBLe III. solutions of carbon disulphide in
nutrient gelatine varying from M/16,000 to M/40,000 in con-
centration, stimulate spores of Rhizopus nigricans and Sterig-
matocystis nigra to germinate sooner than in the same media
but without the CS,. The point of greatest stimulation lies
between the M/32,000 and the M/40,000 solutions.
In another type of experiment with Sterigmatocystis spores,
the stimulation of growth was measured by the increase in dry
TABLE III
AVERAGE GROWTH IN MICRONS OF IO GERMINATION TUBES OF FUNGOUS
SPORES, INOCULATED ON NUTRIENT MEDIA TO WHICH CS, HAD BEEN ADDED
Ruizorus NIGRICANS
Exp. A.
No. - M/800 M/8,000 M/16,000 M/22,000 M/32,000 M/40,000 Control
hou
‘Si 10.6 11.9 59-9 Over 300 oOver300 Over300 250.
Exp. B.
13% no no no no 6.7 52.6 no
15 no no no 13.9 16.4 83.2 no
18 5.0 no no 70.0 50.4 Over 300 45.9
20144 = 26.9 no 9.5 103.9 76.7 over300 59.5
Exp. C.
13 29.7 57-4 over300 139.2 over300 over 300 no
14 59.6 over ae Over 300 oOver300 over300 over 300 no
17 over 300 ce ae “c a“ at ae “ es ae 98.6
Exp. D.
13 129.6 . 202.6 mca 13.2 8.1 10.1 no
313.0 — 22.7 24.1 46.5 14.8
18 over300 over300 —— 44.3 73-6 79.3 44-5
Exp. E
14 67.5 68.3 —— 20.4 39.2 21.0 10.9
17 101.6 148.1 — 62.2 84.3 43-4 25-5
—_— 91.6 96.9 88.2 52.6
19 127.4 over 300
OLDENBUSCH: STIMULATION OF PLANTS 385
TABLE III (con.)
STERIGMATOCYSTIS NIGRA
Exp. A
No. of
hours M/800 M/8,000 M/16,000 M/22,000 abe 000 M/40,000 Control
29% = 44.5 _ $00 0.7 — 6.5 - 20. n
48 126.2 Over 300 — rb 300 over300 224.9
Exp. B
I5 no no no no no 6.4 no
20 no no no no no 75.6 no
35 no no no no no over 300 no
65 no — Over 300 over300 over300 “ “ start
Exp. C
19 no no no no 117.0 124.9 no
25 no no no 137-5 oOver300 oOver300 no
27 no no no over 300 * no
68 no no no Pe ide tceeee no
gI no no OEE COO Oe ae a a ee
Exp. D
19 no no 78.4 57-1 over 300 = =152.3 no
25 no no Over 300 over300 ‘ ** over 300 no
74 no no 4c sa “ee as ac oe ‘i “ee start
weight of the germinating hyphae. In their experiments 150cc.
Erlenmeyer flasks were used as culture bottles and, with pipettes
and other apparatus, were cleansed with cleaning solution, rinsed
in tap water a number of times, then in distilled water and finally
in double distilled water. The same solution as in the former
experiment, but without the gelatine and with double distilled
water was employed as a culturemedium. ‘The trace of ferrous
sulphate was introduced by adding one drop of a saturated
solution of ferrous sulphate in double distilled water.
After the nutrient solution was made up, it was inoculated
in bulk with the Sterigmatocystis spores, and 50 cc. of the inocu-
lated solution pipetted into each flask. The aqueous CS,
solution (also prepared with double distilled water) was added
last. The flasks were then corked by means of a mercury air
trap, which would allow air to come into the flask, but would not
allow any gas to escape from the flask and thus change the CS,
ratio.
386 OLDENBUSCH: STIMULATION OF PLANTS
After being stoppered, the flasks were left in a dark chamber
at room temperature and, as a preventive measure, the controls
were put in a different compartment from those to which carbon
disulphide had been added. After five days, the flasks were
removed, pads filtered off on weighed filter paper and dried in
hot air oven at 70° to dry weight. No attempt was made to
determine the CO, or acid production, although this would have
been an interesting point to investigate.
Although growth was not wholly normal in the controls,
presumably due to lack of adequate circulation which may have
caused carbon dioxide to accumulate, nevertheless the gross
appearance of the stimulated was strikingly different from that of
control specimens. The solutions containing CS, in the most
optimum concentrations bore tough pads, thicker and con-
siderably heavier than normal. All the stimulated pads were
wrinkled and curled, and slightly yellow on the under surface.
Practically no spores were produced, although in some cases a
few yellow sporangia were observed. The control felts on the
other hand were thin and tender in texture, and more or less
covered with, black spores. However, if further experiments
were to be carried out, a more perfect system of maintaining the
concentration of CS, in the flasks would have to be devised in
order to obtain absolutely normal controls.
TABLE IV
DrY WEIGHT IN MILLIGRAMS OF CULTURES OF STERIGMATOCYSTIS NIGRA,
GROWN IN NUTRIENT SOLUTION TO WHICH CS, HAD BEEN ADDED
Exp. Control. M/80,000 M/40,000 M/20,000 M/10,000 M/5,000
2
No CS
A 439 838 800 740 585 270
B 442 1028 815 748 675 370
Cc 530 765 1035 495 490 oe
D 578 788 740 810 540 casa
Avg 497 855 848 698 573 320
The results of TaBLE IV are sufficient to indicate that there:
is a marked stimulation in cultures to which carbon disulphide
has been added. The region of greatest stimulation is in the
flasks M/80,000 and M/40,000 CS, solutions, the more dilute
- concentration being slightly more advantageous than the other.
OLDENBUSCH: STIMULATION OF PLANTS 387
Complete inhibition was not found in any of the experiments
tried, although the flasks containing M/5,000 solution only pro-
duced felts of approximately three-fourths of the normal weight.
In some instances the M/10,000 acted slightly as an inhibitor so
that growth below normal was produced.
SUMMARY
All things considered, the results of this investigation lead
one to conclude that carbon disulphide acts on plants as many
other chemical stimulants do. When applied either in aqueous
solution or in vaporized condition it inhibits growth, if given in
too concentrated a solution or if vapor is used for too long a time.
When used in dilute amounts, carbon disulphide has the
ability to stimulate:
(1) Dormant protoplasm, as in the case where twigs exposed
to CS, vapor for a short period of time quickly resumed their
growth; :
(2) Active protoplasm, as in case where the hypocotyls of
Vicia seedlings elongated more antes upon the addition of
small amounts of CS,;
(3) Conidial spores of certain fungi which germinated sooner
and produced germination tubes elongating more rapidly, the
stimulation causing also causes an increase in dry weight.
Carbon disulphide acts also as other stimulants do, in that the
acceleration of growth lasts only for a certain period of time,
and then growth tends to reach normal again. This fact was
shown in the twig experiments and those with Triticum grains.
The same would probably be true of the other experiments if
left running for a sufficient length of time.
BoTANICAL DEPARTMENT,
BARNARD COLLEGE
COLUMBIA UNIVERSITY,
New York City
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INDEX TO AMERICAN BOTANICAL LITERATURE
1920-1922
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or — upon American material; the word
America being used in the broadest s
Reviews, and papers that relate ec idieiy to foresty, agriculture, horti-
culture, manufactured products of vegetable origin, . or ranean —
riology.
An occasional peaks is made i in favor of some paper appearing in an hee
can 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 three cents for each card. Selections of cards are
not permitted; each subscriber must take all cards published during the term
of his subscription. Correspondence relating to the card issue should be
addressed to the Treasurer of the Torrey Botanical Club.
-Hollick, A. Paulownia in winter. Jour. New York Bot. Gard.
23: 1-3. pl. 267, 268. Ja 1922.
Holm, T. Contributions to the morphology, synonymy, and
geographical distribution of arctic plants. Rep. Canad.
Arctic Exped. 1913-18. 5°: 1-139. 10 F 1922 [Illust.].
Holm, T. Seasonal dimorphism in Arisaema triphyllum. Am.
Midl. Nat. 8: 42-48. f. 1-5. Mr 1922.
Holm. T. Studies in the Cyperaceae XXXIII. Am. Jour. Sci.
3: 138-144. f. r-I2. F 1922.
House, H. D. Nomenclatorial notes on certain American plants
—II. Am. Midl. Nat. 8: 61-64. Mr 1922.
Howe, M. A. The Collins er eptioN of Algae. Jour. New York
Bot. Gard. 23: 23-24. F 1
Hughes, D. K. The “‘serrato-ciliata” group of Tropaeolum.
Kew Bull. Misc. Inform. 1922: 63-85. f. 7-4. 1922.
Includes 16 spp. nov.
Hunter, M. R. The present status of Scolopendrium in New
York State. Am. Jour. Bot. 9: 28-36. f. 7,2. 21 F 1922.
391
392 INDEX TO AMERICAN BOTANICAL LITERATURE
Illick, J. S. The maples. Am. For. 28: 12-19. Ja 1922.
Illick, J. S. What our Christmas trees are. Am. For. 27: 739-
749. D 1ogai. [Illust.]
Illick, J. S. The sycamores. Am. For. 28: 145-150. Mr 1922.
{Ilust.]
Illick, J. S. Woody plants of Franklin County, Pennsylvania.
Pennsylvania Dept. For. Bull. 21: 5-42. 1 Ja 1921.
Jackson, B. D. Thomas Nuttall (1786-1859). Jour. Bot. 60:
59. -- 1922,
Note, supplied by Mr. F. R. Dixon-Nuttall, as to late residence and
burial place of Thomas Nuttall.
Jepson, W. L. The giant Sequoia. In Hall, A. F., Handbook
of Yosemite National Park [California], 235-246. pl. 22, 23.
New York. 1921.
Joffe, J. S. Studies of sulfur oxidation in sulfur-floats-soil
mixtures. Soil Sci. 13: 107-118. F 1922.
Johnson, J. The relation of air temperature to certain plant
diseases. Phytopathology 11: 446-458. pl. 21-23. 25
1922.
Jones, D. F. Hybridization in plant and animal improvement.
Sci. Mo. 14: 5-23. Ja 1922. [[llust.]
Jones, D. F. The indeterminate growth factor in tobacco and
its effect upon development. Genetics 6: 433-444. f. I-5.
5. FOat.
Jones, D. F. Selective fertilization as an indication of germinal
differences. Science II. 55: 348, 349. 31 Mr 1922.
Jones, F. R., & Vaughan, R. E. Anthracnose of the garden pea.
Phytovatholosy 11: 500-503. pl. 25+f. 1-2. 21 Mr 1922.
Judd, C. = Kilanea National Park trees. Hawaiian For. &
Agr. 18: 255-260. D ig21. [Illust.]
Kaiser, G. B. Little journeys into mossland. V. The old genus
Hypnum. Bryologist 24: 58-60. 9 Ja 1922.
INDEX TO AMERICAN BOTANICAL LITERATURE 393
Kauffman, C. H. Studies in the genus Inocybe I. The species
of Inocybe in Peck’s collections. New York State Mus.
Bull. 1919: 43-60. My-Je 1920.
Kelly, J. P. The synthesis of full coloration in Phlox. Science
LH S83 aay og Mr 25a2:
Kindle, E. M. Notes on the forests of southeastern Labrador.
Geog. Rev. 12: 57-71. f. 1-11. Ja 1922.
Klugh, A. B. A correction concerning the life zones of Canada.
Biol. Bull. 41: 272-275. N 1gatr.
Knowlton, C. H., Ripley, W. S., & Weatherby, C. A. Third
report of the committee on floral areas. Rhodora 23: 209-
220. 9 Ja 1922.
Preliminary lists of New England Plants, XXVIII.
Knowlton, H. E. Studies in pollen, with special seference to
longevity. Cornell Univ. Agr. Exp. Sta. 52: 751-793.
Ja 1922.
Knudson, L. Non-symbiotic germination of orchid seeds. Bot.
Gaz. 73: 1-25. f. 1-3... Ja 1922
Korstian, C. F., & Fetherolf, N. J. Control of steam girdle of
spruce transplants caused by excessive heat. Phytopath-
ology 11: 485-490. f. 7-3. 21 Mr 1922.
Krantz, F. A., & Bisby, G. R. Relation of mosaic to running-
out of potatoes in Minnesota. Univ. Minnesota Bull. 197:
1~31. f. 1-18. S$ 1921.
Krause, K. Rubiaceae peruvianae novae. Notizbl. Bot. Gart.
u. Mus. Berlin-Dahlem 8: 101-103. 1 Ap 1922.
Includes spp. nov. in Palicourea (1), Psychotria (2), Cephaelis (1),
Richardsonia (1).
Krause, K., & Hoehne, F. C. Contribugdes conhecimento das
Rubiaceas do Brasil meridional. Anex. Mem. Inst. Butan-
tan 1: 9-33. Pl. I-6. 1922. ;
Includes 6 new species.
Kremers, R. E. Biogenesis = oo of peppermint. Jour. Biol.
Chem. 50: 31-34. Ja 1922.
394 INDEX TO AMERICAN BOTANICAL LITERATURE
Krieger, L.C.C. Schizophyllum commune with a stipe. Myco
logia 14: 47, 48. 6 Mr 1922.
La Rue, C. D., & Bartlett, H. H. A demonstration of numerous
distinct strains within the nominal species Pestalozzia
Guepini Desm. Am. Jour. Bot. 9: 79-92. 20 Mr 1922.
Lesdain, M. B. de. Lichenes prope Habanam in insula Cuba
anno 1914 a cl. fratre Arséne Brouard lecti. Bryologist 24:
68, 69. 15 F 1922.
Levine, M. Studies on plant cancers. III. The nature of the
soil as a determining factor in the health of the beet,
Beta vulgaris, and its relation to the size and weight of the
crown gall produced by inoculation with Bacterium tume-
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Lewis, I. F., & Taylor, W. R. Notes from the Woods Hole
Laboratory, 1921. Rhodora 23: 249-256. pl. 133+f. I, 2.
ay ¥ ‘TO22.
Includes 1 comb. nov.
Loeb, J. Quantitative laws in regeneration. III. The quanti-
tative basis of polarity in regeneration. Jour. Gen. Physiol.
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Experiments with Bryophyllum calycinum.
Laftfield, J. V. G. The behavior of stomata. Carnegie Inst.
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Ludwig, C. A. The control of angular leaf spot of cotton.
Phytopathology 12: 20-22. 10 Mr 1922.
Macbride, T. H. Some of the ways of the slime mould. Mycolo-
gia 13: 329-334. 21 Jia 1922.
Mains, E.B. Heteroecism of Puccinia montanensis, P. Koeleriae
and P. apocrypta. Mycologia 13: 315-322. f. 1-4. 21 Ja
1922.
Martin, G. W. Rhizophidium polysiphoniae in the United
States. Bot. Gaz. 73: 236-238. f. -10. Mr 1922.
Maxon, W.R. Notes on American ferns—XVIII. Am. Fern
Jour. 11: 105-107. 31 Mr 1922.
INDEX TO AMERICAN BOTANICAL LITERITURE 395
McAlpine, D. Bitter pit in apples and pears: the latest results
in preventive measures. Phytopathology 11: 366-370.
a7 Ja 1922,
McDougall, W. B. Biller my! : Drastic trees. Jour. For.
20: 255-260. f. 1-3. Mr
McDougal, W. B. Symbiosis in a deciduous forest. I. Bot.
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McKinney, H. H., & Johnson, A. G. Wojnowicia graminis (Mc-
Alp.) Sacc. & D. Sacc. on wheat in the United States.
Phytopathology 11: 505, 506. 21 Mr 1922.
Meade, R. M. Positions and movements of cotton leaves.
Jour. Hered. 12: 444-448. f. 7-9. D 1921.
Meister, G. K. Natural hybridization of wheat and rye in
Russia. Jour. Hered. 12: 467-470. D 1921
Merl, E. M. Biologische Studien uber die Utriculariablase.
Flora 115: 59-74. f. 1-3. 1922. ;
Millar, W. N. Status of forestry in Western Canada. Jour.
For. 20: 10-17. Ja 1922.
Moore, B. Humus and root systems in certain northeastern
forests in relation to reproduction and competition. Jour.
For. 20: 233-254. Mr 1922.
Moore, B. Influence of certain soil factors on the growth of
tree seedlings and wheat. Ecology 3: 65-83. II f.1-6. Ja
1922.
Mulford, F. L. Broad-leaved evergreen shrubs for the South.
Am. For. 28: 99-104. F 1922. [Illust.]
Murphy, P. A. The sources of infection of potato tubers with
the blight fungus, Phytophthora infestans. Sci. Proc.
Royal Dublin Soc. 16: 353-368. pl. 11-14. Au 1921.
Experimentss conducted in Canada.
Murrill, W. A. A fragrant polypore. Mycologia 14: 46, 47.
6 Mr 1922.
Trametes suaveolens.
396 INDEX TO AMERICAN BOTANICAL LITERATURE
Murrill, W. A. Hygrophorus caprinus. Mycologia 14: 48, 49.
6 Mr 1922.
Murrill, W. A. Illustrations of fungi—X XXIII. Mycologia
14: 25-29. pl. z—9. 6 Mr 1922.
Oliveira, A. A. Métodos practicos para el cultivo de la yerba
mate en el territorio de misiones. Bol. Minis. Agric. 26:
395-402. O-D 1921.
Overholts, L.O. Diagnoses of American porias.—I. Mycologia
14; 1-11. pl. 1+f. 1-6. 6 Mr 1922.
Palm, B. T. The false mildew of tobacco introduced into the
United States from the Dutch East Indies. Phytopathology
11: 430-432. 16 F 1922.
Palmer, E. J. The forest flora of the Ozark region. Jour.
. Arnold Arbor. 2: 216-232. Ja 1922.
Parks, H. E. California hypogaeous Fungi—Tuberaceae.
Mycologia 13: 301-314. 21 Ja 1922.
Paton, J. B. Pollen and pollen enzymes. Am. Jour. Bot. 8:
471-501. 15 F 1922.
Payson, E. B. Monograph of the genus Lesquerella. Ann.
Missouri Bot. Gard. 8: 103-236. f. 1-34. 9 Ja 1922.
Pearson, W. H. Ricciocarpus natans (L.) Corda, from Africa.
Bryologist 24: 69, 70. S 1921. 15 F 1922.
Pessin, L. J. Epiphyllous plants of certain regions in Jamaica.
Bull. Torrey Club 49: 1-14. pl. 1 + f.7. 21 Mr 1922.
Peters, R. Moisture requirements of germinating seeds.
Kansas Univ. Sci. Bull. 13: 23-37. f. 1, 2- My 1920.
Petzke, E. La Fourcroya gigantea (una _ textil). Bol.
Minis. Agric. 26: 403-407. O-D 1921
Pfeiffer, H. Conspectus Cyperacearum in America meridionali
nascentium. I. Genus Heleocharis. Herbarium 58: 85-88.
1922.
Piche, G. C. The forests of Quebec. Jour. For. 20: 25-43. Ja
1922
INDEX TO AMERICAN BOTANICAL LITERATURE 397
Piper, C. V. The identification of Berberis Aquifolium and
Berberis repens. Contrib. U. S. Nat. Herb. 20: 437-451.
pl. 24. 1922
Piper, C. V. An unusual type of “Ou get in ere
cristatum. Jour. Hered. 20: 423. f. zo. Nt
Pittier, H. New or noteworthy plants from Colombia and
Central America—8. Contrib. U. S. Nat. Herb. 20: 453-
490. pl. 27~30. 1922.
Includes new species in Virola (1), Dialyanthera (1), Pithecollobium (10),
Machaerium (9), Protium (2), Zanthoxylum (3), Eurya (1), Lucuma (2),
Sideroxylon (1), Vitex (4), Calycophysum (2).
Pittier, H. On the species of Dalbergia of Mexico and Central
America. Jour. Washington Acad. Sci. 12:54-64. 4 F 1922.
Includes 8 new species.
Plitt, C. C. A preliminary report, with notes, on the lichens
found near the Cinchona Botanical Station, Jamaica,
British West Indies. Bryologist 24: 60-64. 9 Ja 1922; 24:
70-74. TEE 1929.
Poole, R. F. Some recent investigations on the control of
Sclerotinia Libertiana on the green house on the muck
forms of Bergen County, New Jersey. Phytopathology
12: 16-20. f. 1-3. 10 Mr 1922.
Popenoe, W. The Andes berry. Jour. Hered. 12: 387~393.
frontispiece + f. 1-3. N 1921.
Rubus glaucus.
Popenoe, W. The frutilla or Chilean strawberry. Jour. Hered.
12: 457-466. f. 13-17. D I9g21.
Popenoe, W. The native home of the cherimoya. Jour. Hered.
12: 331-336. 20 F 1922. [Illust.]
Prince, G. H. Forestry administration in New Brunswick.
Jour. For. 20: 54-61. Ja 1922
Pritchard, F. & & Porte, W.S. Effect of fertilizers and lime on
control of tomato leaf spot (Septoria Lycopersici). Phyto-
pathology 11: 433-441. f. 1-10. 25 F 1922,
398 INDEX TO AMERICAN BOTANICAL LITERATURE
Pulling, H. E. Biophysics as a point of view in plant physiology.
Am. Jour. Bot. 9: 37-46. 21 F 1922.
Ramsey, F. T. The Swazey barberry. Jour. Hered. 20: 426,
497. °f. o4...N 1921.
Berberis Swazeyt
Rayner, M. C. Nitrogen fixation in Ericaceae. Bot. Gaz. 73:
226-235. f. I-4. 1922.
Record, S. J. Boxwoods of commerce. Bull Torrey Club 47:
297-300, f.. I... 31 D.1922.
Record, S. J. Notes on woods. Science II. 55: 267-269; 10
Mr 1922.
Rehder, A. New species, varieties, and combinations from the
herbarium and the collections of the Arnold Arboretum.
Jour. Arnold Arbor. 3: 11-51. Jl 1921.
Includes 1 sp. nov. in Prunus.
Reiche, C. Rasgos biologicos del abrojo. Mem. y Rev. Soc.
Cien. 40: 41-44. pl. zr. O 1921.
Opuntia tunicata.
Renacco, R. Sobre algunas enfermedades parasitarias de las
plantas cultivadas. Anal. Soc. Cien. Argentina 81: 62-70.
Ja—F 1916.
Round, E. M. A modern plant fossil. Bull. Torrey Club 49:
Os, O4. 7. t. t2 Ap 1922.
Sargent, C. S. Notes on North American trees—IX. Jour.
Arnold Arbor. 3: 1-11. Jl 1921.
Includes 11 spp. nov. in Crataegus.
Sears, P. B. Variations in cytology and gross morphology of
Taraxacum. Bot. Gaz. 73: 308-325. pl. 9, 10. Ap 1922.
Sears, P. B. Variation in cytology and gross morphology of
Taraxacum. Il. Senescence, rejuvenescence,; and leaf var-.
iation in Taraxacum. Bot. Gaz. 73: 425-446. f. 1-0.
Je 1922. é
Seaver, F. J. Phyllostictales. oo (pars). North
American Flora 6: 1-84. 5 Ap 19:
INDEX TO VOLUME 49
New names and the final members of new combinations are in bold face type.
ia constricta, on li
Acerates lanuginosa, 98; viridiflora, 98
se egundo, 2 rubrum, 165;
sien
chyron nychia, 351; Rixfordii, 351
x42. equinus, wer mil 7 nats
papilionaceus, a Sajor-Caju, 230;
striatus, 231; 232
— 32: f aenc fe, 242; utahense,
2
fay de 104; dasystachum, 105;
molle, 105; repens, 105; Smithii, 105;
um, 105
Albizzia lophanta,
leurina, 19; elas stica, 21; Sas Peat
6; marchica, 16; olivacea, 20
Algae in the Chinese i hl Refer-
ences to the, 2
Algae, New British and American
speci Lobomonas: stud
morphogenesis of si a 123
1, 351; nycta-
nea, 351; ovata, cyte pumila, 351
ophila, 3
Altecnares. 332; Solani, 314, 323, 333-
33
Amaranthus er eee 243
Amauria dissect
Ammauriopais dis siieoccee, 359
Ambrosia, 53
American fossil hepatic, A new, 207
American species o a
study in m st seer eee of motile
algae, New British and, 123
o-t35
Amsinckia intermedia, 100; lycopsoides,
» 247
e development
of the PS a gery cei ete y ot
Martynia louisiana, ae
Andropogon saccharoides, 2
Anemone, 240; ng rosa, ae: spheno-
phylla, 239, 243.
Anislotus brachycarpus, 240, 24
oe ulus, 239, 245; trispermus, yh
Antennaria oe 181
Anthericum, 54
Antholyza
za, 55
Ll Rotieshinnes cigar 248
Aphanolejeunea, 4, 5, 7; sicaefolia, 5
AplopappusGooddingl, 44; past hi 44
79
el, 179; congesta, nied
lithophila, 179; subcongesta,
Aristida adscensionis, 242; seve tm
242; divergens, 242; Wrightii, 242
Aristolochia, 51; fimbriata, 51; Sipho,
pshencie: Ko read botanizing in
souther
Artemisia ita dai Brittonii, 250;
ularis, 102
Anton, JosEPH a New species
ot Uredi neae—XIV,
Asclepias scone ay eriocarpa
ge » 3553 ‘linaria, 247; pum-
98 not iosa, 98; syriaca, 97,
~ verticillata, 98
Ascobolus, —
ASHE, i ny am and shrubs
a the hic Mahe United States,
265
Aspergillus, 328, 376; niger, 375-
Asphodelus, 54
Aspidium simulatum, 63
sie
tragalus Nuttallianus, 245; parvi-
101
Aifoiat ni mney 238, a hastata,
97; patu ‘a, 97; rosea,
Audibertia capitata, 37
os reticulata, 228
Aven
hyedis Nailin; 246
Baccharis glutinosa, 250; sarothroides,
250
Bahia dealbata, 250; dissecta, 359
Baileya multiradiata, 238, 250
( 1 Ld? ra na L
399
400
Bartram, Epwin B., Midwinter bot-
anizing in so uthern Arizona, 237
Basidiouiycetee—V, Philippine, 223
Bebbia juncea, 2
etula, 172; lutea, 172; nigra, 172,293
Biatorelis. GaN —— 164, 78
echnum, 8; m, 3,
5;
Boletus aha, > Mees:
229
Bolivia, New = of trees of medical
interest from
Botanizing in gone ce Arizona, Mid-
05, oe cinerea, 305,
» 325, 327;
Siete filiorimis, Liew Parryi, 242;
Otnroc ii, 242
Brachiomonas, 75- , 80, 82, 84,
, 87, 89, 92, 125,131, a 135-137;
gracilis ths 76, 81, 89, 91; simplex,
—85, 89, 91; submarina, 75, 76, 78,
~: submarina
obtusa, 77, 79, 81, 89, 91
Brachiomonas, The peste of the
£i7>
Brachvoneiian. 207
Tuscus, 212, 213; Alleni, 211-
213
Brassica nigra, 244
British and American species of
Lo monas: a study in morpho-
genesis of motile Hs ner, 123
BRITTON, Ros
we
us, 104; eximius, ie
inermis, 94, 95, 100, 101; japonicus
rum, 100; vul garis exi-
176
Rucumers comes 195
Buechnera
gr es 21 inquinans, 15; nigrita,
ae “nigita, 16; pulla_ nigro-
olivacea
Bullaria, As Zorniae, 190
Cactaceae, Two new genera of, 251
California—1, Miscellaneous notes on
plants o uthern, 31;—
Calliandra eriophylla, 239, 244
Callisia, 271
Callistemon lanceolatum, 34
_Callitriche palustr ris, 246
Campan * sibirica, 100, 101
canpte
55
ei 154, 156
Caraguata, 12; Sintenesii
INDEX
Carex, 55, 103, 369; alata pulchra, 366;
aquatilis,176; atherodes, 1 h
tachya, 103; Bicknellii, 369; ‘brevios,
cumu-
foenea, ; Goodenovii, 177;
Heleonastes, 175, 177; Hookeriana,
103; hormathodes, 177, 361; hor-
4, 366; tenera, xa | = nera major,
362; tenera Richii, , 362; vesi-
Caria, 177
Care —-X1l, Notes on, 361
Carlowrightia arizonica, 249
Carpolithe ae II
Cassia Covesii, 245
Castanea ainifolia 266, 267; alnifolia
mila praecox, 267; pumila
Castilleja, 1 I9I,
Catinella, 15; elast tica, 16, 21; nigro-
oli , 18-20; gia 16
Catinella, The’ Lane
ste flexili 35,
Cecidomyia aos 163
Celtis palin 243
Centaurea picris,
peahies America, The genus Costus in,
2
Cephaleurps, 12
Cephalothecium roseum, 320
Cercidium i erexiies 245
ereus gigant 247
Chaetochloa macros. 242; viri-
dis, 2 2
Chamaerhodos erecta, 100
Chetenthes Covillei, 32; 33:
Lindheimeri, 239, 241; myriophy la
glei, 241; Wri ghtii, 241
C m, 95; album, 95- oie feru-
ast 96, 97 le eptophy lusts 96, 97;
6, 243; pratericola, 97
Carbon disulphide, corgi of
lants
Chilopsis pee 249
INDEX
marr 4 occidentalis, 355; umbel-
ata,
Chinese Serge References to the
algae in t
Chlamydomonas, 75, 78, 79, 82, 84, 85,
87-89, —12
89, 92; metastigma, 128;
culus, 123; reticulata, 127; aoe ete,
23; subcaudata, 85, 86
28
m Leucanthemum, 100
Chee lavicifolia, 249
Chrysom 19
Chrysothamnus lortubata: 101; graveo-
ens, IOI; nauseosus, IOI
Cirsium neomexicanum 238, 250
Citrus Lim ng
3)
on
BG
SS
2.5
S
FB.
JN
merica 5
eee nee lanceolata Pelceondi, 352;
umbellata 352 |
Clematis Dr rummondii, a
Clusia havetioides, 5, 6, Oo. ag
Cobaea scandens, 53, 55
CocKERELL, T. D. A., A new genus of
fossil Liliaceae, 211
Coenogonium, 7
Coleosanthus’ baccharideus, 249;
Coulteri, 249; Wrightii, 249
Collema, 7
Colletotrichum, 332; lagenarium, 323,
325, 329, ae ; Lindemuthianum, 333;
i
> +
°
s
Cololejeunea, 4, 6, 7; diaphana, 4-7
olorado, Two new plants from west-
Commelina, 270-
pee i Co vouchnhatin: Anew
__ genus of the, 269
1a
272-—
oe
w
275
' Commelinantia, A new genus of the
mmelinaceae, 2
Commelyna, 269, 270; anomala, 269;
rhodantha, 2 :
Commicarpus scandens, 243
Condalia spathulata fide
Copelandia, 232; papilionacea, 232
Compe ae m, 97; hyssopifolium, 97;
itidum, 97; villosum, 97
. aan, 166
Costus, 283, 284; Bakeri, 285, 2 288, 289;
b eatus, oe 285, 292; comosus,
288; 85
3
8; Danthonia pinetorum,
401
Malortieanus, 285, 28 9, 290; maxi-
2
92; villosissimus, 284,
Costus a cou America, The genus,
Covil: iea Stain 237, 245
Cracca ten ella,
Crassina
Crepidotus, 232; pogo
Crepis capillarie, | 103; podria Hs 103
ronartium, 191; coleosporioides, 191;
194; filamentosum, 191, 194; Hark-
nessii, 194; Quercuum, 193; ribicola,
re nae 319-321, 323, 329-331, 333,
ctiforme, 19h 194
Crossotolejunes
Gr yptantha calycosa, rags gracilis, 39,
40; s Hillmanii, 39; pareoere
- a : maritisia, ok cemosa,
8; ramosissima, 38, 39; recurvata, 39
Giecae 376; Pepo, 54, 382
uscuta, 107; yet 2,0 ROG;
arvensis, 98; calycina, 98; Choisiana,
107; Coryli, 98; corymbosa grandi-
flora, 109; cozumeliensis, 108;
dentatasqu a, 307; an-
ama ur
gana, 109; floribunda, 109;Gronovii,
98, Gronoviicurta, 98; macrocephala,
108; partita, 10 ntagona, 98;
plattensis, 98; tinctoria, 109; um-
ellata,
uscuta from Mexico, Three new
107
Eotindeodadion: scoparium, 320, 328,
330, : :
Cy ee a 242; diandrus, 103;
sculentus,
Cpettieta ieee. 323, 330, 333
Dacryomyces, 166; ie sat 167,
168; hyalinus, 1
Daedalea, 225; amanitoides, g25° a
a, 226; avid:
225; inconcinna, pers, indica,
intermedia, 2o7: lurida, 225; Palisoti,
eed alvanrierse 225, 227; repanda,
225; sanguinea, 224; striata, a
subconfragosa, 225; versatilis, 2
anaea,
175;
spicata
pinetorum, 175; thermalis, 175
402
Dasylirion Wheeleri, 239, 242
Dasyochloa _ pulchella, 242
Daucus p 24
Desmidiu
um, 4
Development of the flower and embryo-
Martynia louisiana, The,
141
Diabole, 194; cubensis, 194
Diapedium Torre i, 24
we Polygoni-alpini, 189
Dichorisandra, 271
Didymellina Iridis, 329,
Dioscorea, 55; qui eieuetis. 51, 55
Diplasiolejeunea emia , 4-7
Diplocarpon Rosae, 330
Dipterostemon cations 242
Discocactus, 2 he
ecg Bac. Lachnea with a
botryose e conidial seal 301
Eect of external and
internal factors on he : germination
ot fungous spores,
DospaLL, LouIss, Boece of the
beget stage of Puccinia Taraxaci,
235
Draba___corrugata,
— :
352; corrugata
352; cuneifolia, 239, 244;
xosa, 352
licaceagkeken Moldavica, 100; parvi-
iflorum, I
Drepanolejeunea campanulata, 5-7;
crucianella, 7
ryopetalon, 240, 247; runcinatum,
239, 244
Dryopteris spinulosa, 105; Thely-
omanesagay 130
cae LiaAs J. The genus Cati-
15
Dys nite a poro a 250
Earliella contatiala aah 24; cubensis, 224
Echin tus, r eguinii, 252;
bicolor, 251; conoideus, 252; hexae-
ophorus, 7: we othele, 251;
Wislizeni, 2
inocereus Fendleri ri, 2 247
Echinodorus cordif 35
Effect of external and yh tab’ factors
re sisi germination of fungous
» 313
Blaphoglossm, 3; latifolium, 4, 6, 8-
Eiatiae californica, 35
—— cladophora, 226; ves-
Encelia "farinosa 240,
Endothia parasitica, pia
INDEX
Ephedra antisyphilitica, 241; trifur-
cata, 241; viridis, 241
icampes ringens, 242
Epidendrum, 7
Epiphyllous plants of certain regions
in Jamaica,
Eragrostis megntincll ins. 242
Eremolithia Rixtordii, 351
wd hae piechioae 250; grandiflorus,
I
Encwodtnt apiculatum, 35; fascicu-
latum, 351; fasciculatum fiavoviride,
350; fasciculatum foliolosum, 351;
fasciculatum ifolium, 350,
su 50; nodosum Jaegeri,350;
ovalifolium, 179; pinetorum, 2
lycladon, 243; Wrightii, 24
ae
ium cicutarium, 245; texanum, 245
Erucastrum, Pollichii 1;
246; polycarpa, 246
Evolvulus Wiha, 2 247
Evonymus atropurpureus, IOI
Fagus pumila, 267
Farr, Ciirrorp H., Quadripartition
y ing in Sisyri inchium, 51
Favolus, 228; cucullatus, 227; curtipes,
227; orus, Rows! Itiplex,
228 , 228; spathulatus, 228
244
tuca Hallii, 104; viridula, 104
issidens,
Floscopa, 271
Fomes, 172; Bakeri, 172; rimosus, 172;
_ robustus tee
Se Ossi : ie modern plant,
. A abae Dhak 207°
“ossil La, iaceae, A new genus of, 211
ouquieria splendens, 24
fohcahta. 53
*ranseria rate, 249
pranera arryi,
?raxin us attenuata, 247
“reesia, 55
ritillacia, 56
coides erectus, 207
uct us, 135
Funalia versatilis, 226,
Fungous spores, Effect ‘of external and
pie nal factors on the germination
Galactia Wrightii, 245
INDEX
Galium, 358; angustifoliumpinetorum
357; californicum, 359; grande, 359}
Hallii, 358; multifiorum, 358; occi-
dentale, 358, ; gc Ha 359;
stellatum, 249, 3
» 24
ntiana humilis, 349; A got ex, 349
pep pees of fungous spores, Effect
rnal and internal factors on
t “yi
Glacier National Park, Unreported
plants Fides 175
Gleoca
Giosoporiat 228; candidu
us, 229; con-
pi nines ides, 228; " dichrous, 229; retic-
tus, 228
fo pats ig rufomaculans, 314, 320,
328, 330 meee
Gnaphalium alpinum, 181; Wrightii,
Goalaus pectorale, 127, 128
pefieong
FF, bare W., Philippine Basidio-
predilst Si 223; Unreported plants
from Glacier National Park,
angii, 262
Gymnogramme triangularis, 32
Gymnosperma co sum, 2
Gymnosporangium clavipes, 323, 330,
331, 334, 335; Juniperi-virginianae,
318, 321-323, 330, 331, 334
Hapalopilus subrubidus, eek
Haplop hy ton cimicidum
Harting annia — osa,
AZ ey ‘British and
peat
TS fe) onas: a
udy in morphogenesis of motile
algae, 123; The phylogeny of the
Dn
_ genu » 75
Hedeoma nana, 248; oblongifolia, 248
Hedychium, 3; coronarium, 3, 4
: scens, 4
elenium montanum,
Helianthus, 53; annuus, 382; giganteus,
IOI; serratus, IOI;
miliani, 101; ‘ditidus: IOI; tuberosus,
101
Hepatic, A new American fossil, 207
Herbert, 208; adunca
Hermannia pau uciflora, 230, 246
Herniaria cinerea, 35
Hexagonia, 227, 228; albida, 22
apiaria, 227; —" 229° ciliata, Son
403
cladophora, 226; Clemensiae, 27
cucullata, 227; glabra
Taxodii, 2 ve 226
ibiscus Coulteri, 246; denudatus, 246
Hoffmanseggia densiflora 245
Holacantha Emoryi,
HOLiick, ARTHUR; HowE, MARSHALL
Ac, & new American fossil
hepatic, 207
Howe, MARSHALL A., & HOotticx,
ARTHUR, ew American fossil
a aah 19; Synge a 16; macchica,
6, 20; olivacea, 16
pdncdiet von, 78, 299
ymenoclea monogyra, 249
Hymenogramme spathulata, 228
Hymenophyllum, 3
Inga vera, 195
Inonotus Clemensiae, 227
Iowa, Quercus lyrata in, 293
Tris, 55 :
Isocoma Hartwegi, 249
Ixia,
Jacquemontia Pringlei, 247
Jamaica, —- llous plants of certain
regions in,
Jamesoniella Rc MERE 208
Janusia gracilis, 2
i i
M., peng Pump A.
n plants of
nia—I, me -—II, 349
titperus scopulorum, 239, 241
Karschia, 15
Koeberlinia, spinosa, 245
Osa, 245
nitzkia maritima, 38; ramosissima,
Lachnea, 301, 305; undans, 301,
302, 304, 305; Pay a6t, 303, 304
chnea with botryose conidial
stage, Pe 301
Lactuca sativa, 53; virosa, 103
Laminaria
Lamium am lexica
ppula cenchrusoide s, 99 cilia 180;
carulitn foliosa, 99; floribu ae 99;
Lappula, 99; occide ntalis, 99; texana
homosperma, 99
404
Laschia, 229; minima, 229; spathulata,
228
nora, 7
Lejeunea flava, 4-7
Lemalis, 18; pulla, 18; pulla nigro-
olivascens, 18; rufo-olivacea, 16, 18
Lemna, 103; trisulca, 3
Lentinus, 229, 232; brachatus,
phyi 226, 231; ae eas!
panda, 225; striata,
ata densiflorum, os. 94; Draba,
94; hirsutum, 244; lasiocarpum, 239,
Lesquerella Fendleri, tts 244; pur-
purea, 239, 244
A new genus of fossil, 211
124, 128,
133, 135-137; Francei, 123-125, 127,
129, 130; pentogonia, 124, 125, 127,
9, 139; rostrata, 124, 127, 130, 139;
cetiace. ae
Lobomonas: tudy in morpho-
genesis of reo algae, New British
d American species of, 123
Lolium remotum, 104; rigidum, 104;
4
m macrocarpum, IOI
padium
Lophocolea, 5- Z
204; affinis, 198, 199,
Asch rnii, 204; Barkeri, 197;
bicolor, 204; bilineatus, 197; car-
nosulus, 204; chihuahuensis, 204;
Clarkei, 197; densiflorus, 200, 201
M
130-|M
INDEX
203; scohwittecntie Layneae, 203; tri
fidus, 204; umbellatus, 204; vallests,
204; vallicola si 2
Lupinus succulen and L. s,
Studies in the rae india Ae
197
ern comosa, 349
Lychnis alba, 95
Lycium, 357; Cooperi, ih Fremontii,
oe arviflorum, ; Spencerae,
356, 357: Torreyi, 248
Lycopus communis, I
ar ee KENNETH KENT, Notes on
C LL, 326%
Malacocarpus, 251
Malus elongata, 268; elongata Popens.
268; platycarpa par
Malva borealis, 95; crispa, con dk:
cillata,
Matcastruns 353
M ammillaria conoidea, 252; Grahami,
4d Se ee
Mangifera indica, I, 5, 10, 13
Marattia
Marchanti tia Pealei, 207
35
Martynia, 141, 150, 154; fragrans, 141,
142; lutea, I
The Divcionnent
of the fl rand embryogeny of, 141
ecardonia pedunculars,
Medicag
shines My hodocsnee 191;
roanensis,
odora scabra,
Merulius fugax, 168° oe striatus, 231;
subaurantiacus, I
Metzgeria furcata, 4-7
ee new species of Cus-
07
1 ;
Ehrenbergii, 200, 201; Hartwegi, | Microlejeunea, 4,
197; hirsutulus, 204; leucophyllus, | Micros stylis ‘monophillos, 349
205; Liebmanni, 199; micranthus, | Midwi botani in southern
204; micranthus microphyllus, 204; yhoo “4 237
Nanus, 199, 2 205; niveus, 197, | Mim asperata, 194; pigra, 194
204, 205; pachylobus, 204; persistens, | Mimulus Langsdor fii, 248
204; Pipersmithii, 204; polycarpus, Miscellaneous note plants of
99; ramosissimus,
200, 201; ‘eatichangers 204; sabulosus,
losus, 204; succulentus,
197-200; succulentus Brande esgeei,
s on
uthern California—I, 31; —II,
ee ium perenne, 5, 6
Modern plant fossil, A, 63
INDEX
Mollisia umbonata. 19
Mollugo verily 35
Fe pe retia,
oclea Gotischei, 6
e meahanie
ublenbebela dumosa 242; micro-
ma, 242; Port sh =
Muounz, Pur A., NSTON, IVAN
M., Miscellaneous ae on plant s of
South rn California—I, 31; —II,
oe
Mycoidea parasi
Mycological eh yah 1920, 163
Oporum serratum, 15.
Myosurus cupulatus, 36
Naias, 54; flexilis, 35
Nama hu humifusum, 356
261; coto, 260
ie “faery 1645 ‘resinae, 164
olloydia, 2
po 252
New prerican fossil hepatic, A, 207
merican sic of
New British a
b rpho-
a >=
A, 211
ommelinaceae,
269
Nee jee from western Colorado,
153
New haa and other notes on North
akota plants, 93
New species of trees of medical interest
from Bolivia, 25
species of Uredineae—XIV,
muicrocarpa,
242
North + eng oe - records
oe er notes o
Nost aap aes 299
Notes te 1920, Mycological, 163
Notes on Carex—XII, 361
Notes on — Dakota plants, New
records and other, 9
Notes on plants of Southera California,
239, 241; rryi, 32; sinua
sinuata integerrima, 31, 241; tenera,
Nuttallia, 183; marginata, 183
2
ea,
Nyssa sylvatica, 166
1; Beguinii, 252; conoi-
189
59; glauca, 248;
405
Occurrence of the pycnial stage of
Puccinia Taraxaci,
Oenothera, 354; cardiophylla, 3553
mahi i soleudene, 354; speci-
4
Ginna CARRIE, Stimulation of
plants by = disulphide, 375
Olneya tesota
O phioglossum’ Aes 211
Gober monophyllos, 349
Opuntia arborescens, 247; Bigelovii,
238, 247: chlorotica, 247; fulgida,
246; si peirsameges 247; Toumeyi, 247
Osmunda, 52; ustris aurea, 52;
, Two new plants
from western Colorado, 183
OVERHOLTs, ., Mycological notes
for 1920, 16
Panaeolus, 232; ‘campanulatus, 232°
papilionaceus, 232
Pandorina, 12
Panicum perlongum, 103
Pannaria rubiginosa, 7
Panus, 2 =
Papulos
Poriathcia Ohta 243
Parkinsonia ccitonoiie: 245
-6
a, 349
Parosela Greggii, 245; Polen 245
Parthenium incanum,
Patellaria applanata, 16 Ai hirneola,
16; olivacea, 16, 20; , 19; pulla
nigro-olivac 16, e: violacea, 16
Patinella applanata, 16, 19; hirneo
16, 19; olivacea, 16, 19; Violacea, 16,
I
Pectocarya scent 247
Pediastrum, 131, I
| Pellaea compacta, rey mucronata, 239,
241
Penicillium apr my 334, 375
Penstemon, 43, 189; albomarginatus,
44; antirrhinoides, 43; antirrhinoides ,
m us, 43; barbatus, 40;
barbatus Torreyi, 404 calcareus :
eA gi 41, 357; Clevelandi con-
3573 rig ma Bem? Stephensi
x; 357; ‘desertorum, 42; Eatoni, 40;
ruticosus, 180; Grinnellii, 42;
ziesii, 180; microphyllus, 43% Munzii ii,
40; Palmeri, 41, 42; Palm ri Grin-
nellii, 42; petiolatus, 42; Plgamerae,
: po he es 41; subulatus, 40;
Perezia Weightii, 249
Peridermium, 193; erebrum, 192, 193
406
pragma I9I, 194; aia egg
I
I-193; sta jenoigp sen 19I,
Perityle dissecta
pat ane detantting 323; pys-
» 335
Paasie: Louis J., ea Rae —
certain regions in Jama
sc
cea, 16, 18, 19;
vi 15-1 einer -atra, 16, 19, 20
Phacelix Franklin
Phaeopezia, 19; ved enal aI:
, 20; marchica, I
fusco-
S, 37
P iladelphus microphyllus, 244
Philibertella linearis,
Philippine 223
hlox, 99; andicola, 99; rnardina,
356; dolichantha, 336% Dow glasii, 99;
oodii, 99 nsburyi brevifolia,
356; supe 7
Photalenron gt eee el 243; cali-
fornicum distans leucocarpum, 349;
macrophyllum, 2
Phragmidium violaceum,
Phycopeltis, 3-7, 10; seieccjactl;
Phyllonoma ruscifolia, 212
e,
Physalis ixocarpa,
248
Phytophthora infestans, 323, 327, 330,
333
Picea, 191
Pilacre oe 165, 173
Pinnular a, 4-6
inus poy
193; oes
Virginia
193; ponderosa, 164,
193;. Strobus, 167;
164
iper, 3
Pittosporum tobira, 4
Plagiochila, 6
Plasmodiophora Brassicae, 322, 323
asmopara viticola, 323, 330, 333, 334
Platanus occidentali is, 293; Wrightii,
244
Platymonas, 80; subcordiformis, 80
ae i 229 ; labellatus, 229
Pluchea, 194; fasti tigiata, 194; Quitoc,
I
Plumbago scandens,
Poa Bigelovii, 242; og 176;
ae 6
S, 3; coriaceus, 5, 7
Polemoniam confertum, 179
Pol
Posbioharkies 80, 128
INDEX
Polygala macradenia, 240, 246
num, 189; pat neha
tachyum, 97
Polypodium, 2,66) 8¢ Este gperte 78
petals, 170,
I7E; t74; pete on a 224; dichrous,
229; Ellisianus, 172; fusco-badius,
incanus, 224; indecorus, 224;
lacerus, 226; leucoplacus, 223; mega-
porus, 228; minimus, 229; nigr
purpurascens, 229; paleaceus, 224;
pelleporus, 229; scabrosus, 224;
su ubrubidus, 228;tegularis, 224; thele-
Polystichum opty 9 8
pee 226; a: 7 Sig ; Persoonii,
; thel ephoroi es, Bp versatilis,
Pin 243; deltoides, 293; Wislizeni,
Por a, 223; leucoplaca, 223
Poesia juniciforme, 250
ee ERFIELD, JR. W. ences
to thealgaein the Chinese aaa: 297
Potentilla argentea
Preissites Wardii 207
Prosopsis glandulosa, 245
Prunus fasciculata, 35
edicactane 317; Tri-
123
Ptiloria pauciflora, 249
Puccinia, es Antirrhini, 316, 323, 330;
biocellata, 194; coronata, 321, 323,
332, se dispersa, 323; graminis,
321-323, 329, > inva! ta, 195;
Larici, 323; Malvacearum, 314, 322,
323, 329 ; offuscata, 190; Phlei-
pratensis, 323, 330; Plucheae, 194;
Polygoni-alpi 189; rubigo-vera,
323i A tenet ee Tare, 235; Zor-
190
Puccini Taraxaci, oop ceaa of the
pycnial stage
Pucciniastrum eecaniien. 190; arcti-
cum americanum, 190
Pyramimonas, 80
renula, 7
Pyrola picta, 355; umbellata, 355
| Quadripartition by furrowing in Sisy-
ir pat
nchium
INDEX 407
Quercus, 170; acuminata, 268; alba, | Senecio aureus, 181; crocatus, 181;
170; aquatica, elongata 268; ari- Lemmoni, 250; manitobensis, 102
zonica, 243; iinokeae 293; colora- ip poe hee Gladioli, 330; Lycoper-
densis, 268; Emoryi, 293, 243;
hy poleuca, 239, 2433 hig pigs Pane l * Quercus lyrata in Iowa,
ma ee 268, 293, ; migia|. 29
elongata, 268; nigra cadiuure Shrubs of the southeastern eee
268; oblongifolia, 239, 243; stellata, tates, eh - ee and, 265
268; virgini ana, 268 icyos angulatu
Quercus lyrata in Iowa, 293 Sideranthus pate 249; Gooddingi,
44
gaa ences to the algae in the Chinese Silene dichotoma, 95; wig 179;
Rha assics, 29 Fabaria, 95; noctiflora,
amnus. tomentella, 2 ir Silphium, 147
Rhizina nigro-olivacea, 19 mmondsia californica, 246
Si
are pte igricans, 314, 1386, 327, 335, Siphonoglossa longiflora, 249
84 Sisymbrium Loeselii
Rividestina I, 6, 191; catawbiense, Sisyrinchium, 55 ue 30 195; Ber-
190; punctatum, 190 font he Ae a ;
i 6 ’ 1, 6,
i mudia
epegie “ ee
_N., Britton, N. L., & Two! Sisyrinchiuin, _ uadripartition e ur
new genera of Cactaceae 251 SMITH, Cuatias PrPER., Studies in the
eae stupas Amodern p lant fossil, genus Lupinus—VII. L. succulentus
3 and eus, I
ROWLEE, W. W., The fe Costus in Solenia ta ek 1733 villosa,
Central America, 283
Rubus neglectus, 190; occidentalis, 190 Solidago humilis, 180; Purshii, 180
Ruellia tuberosa occidentalis, 249 Sonchus arvensis, 102; uliginosus, 102
mex Britannica, 105; hymenose- Sophia intermedia, 94; ochroleuca, 244
palus, 243; paucifolius, 189 Southern California, Miscellaneous
Russy, H. H., New species of trees of nies on plants of, —I, 31;—Il, 349
medical interest from Bolivia, 259 Sphaeralcea, 353; mbigua, a, 246, 353:
Ruscus, 212, 213; hypoglossum, 212; cise eae 1 A 246; F , 353
oe eervium, 213; latifolius, 213 xo
Rutosma purpurea, 246 Spirodela polyrhiza, 103
Salix orbicularis, 178; — 175, | Sporobolus asper, 10
178, 179; saximontana, I | Stau im, 4
Salvia lanceolata, 100; pea i a7; oe ophiogl
: : ee
Salvinia, 211; reticulata, 211, 212 | ea
Samb 52
Bambaci ee Sterigmatocystis, 384, 385; nigra, 376,
STEVENS, O. A., New records and other
see : “be , 36 pee on North Dakota plants, 93
Stic 1 3) 4
Sonpnep ner _agaiicoides, ~ Stimulation of plants by carbon
232
1 » 232;| disulphide, 3
geeton ie pi ce Nin Ee eonsttiie arizonicus, 244
| Str
Scirpus poeide heals 103; validus, 103 | Streptolirion, 271
Sel lutinum, ula,
Sclasotinin. yo ail 314, 330, 331,| Studies in the genus chai
333, 335; Geranii, 301, 302, 305; L. succulentus and L. n = 197
) : Study in morphogensis og sania were,
Ricini , 208
Sconclohier 351; nitrophiloides, 351; ped mgnas sh and og rican species
: 5 5
i mtg Suckleya
Se oo geen eareitg oe ;
ytonema, 4-7
Sedum “Grifhithaii,
Selagi i fod 241; rupicola, 241 | Taxilejeunea, 5-7
408
Leppert B. C., Commelinantia, a new
genus of the ener 269
techies dolosa
elocactus, 251; bation, 251;hexa
orus, 251; lophot thele, onl
a aa 352
Thre — of Cuscuta from
Mexi
Thysanocarpus, 240, 247; amplectens,
Tilletia | Ioeteies, 330
inantia, 269, 270, 272; anomala, 270,
273; fugax, 270; Pringlei 5
pe ig 211; Alleni, 212
rtula inermis, 239; ruralis, 239
Trace antella, 271
ES Dea adits. 54,
269-273; anomala,
260, 271-273; Pringlei, 271, 272
Tragia ramosa,
pera, 223; corrugata,
224; incana, 223° aie 224; palea-
a, 22. ; versatilis
rapella sinensis, his
Trees and shrubs of the southeastern
United States, Notes on
Trees of medical interest from Bolivia,
ew species of, 259
Trentepohlia, 3; arborum, 2; aurea, 2,
4, 5, 7
oe fasciculata, 242
richomanes, 3, 5
A emanomy
new genera of Cactace e, 251
Two new plants from western Colona y
183
Ulmus americana, 293
Notes se — and
shrubs of the southeastern, 265
Unreported eer om "Glacies Nat-
ional Park, 163
INDEX
Uredineae—XIV, New species of, 189
Ur — —— 194; coleosporioides,
194; cumula, 195; curvata,
nominata, 194; Plucheae, 194; Zor-
niae, 190
romyces, 189; caryophyllinus, 316,
323; fuscatus, 189; Trifolii, 323
romycladium, 194; cubense, 194
Usnea, 4
Ustilago Hordei, 330; Tritici, 330
Vaccaria Vaccaria, 95
— occidentalis, 180; sylvatica,
omingensis, 180
Vannuatiot californica, 244
bento inaequalis, 314, 317, 318, 320,
325- aif oe he 334, 335
150; Wrightii,
Verbena 248;
Veronica Cusickii, 180; casera 98
Vicia, 381, 387; Faba, 380 :
— — Angers
in
» SI
Viola Shelton? 353, 354
edeliella incarnata, 243
Woodsia scopulina, 31
Woodwardia ie tetiolia, 63
Xanthium, ge acerosum, IOI; cana-
dense, 10 ita licum, 101; pennsyl-
vanicum,
ucca baccata, 33, 34, 239, 242;
baccata. macrocarpa, 34; elata, *
242; glauca, 196; mac rp ij
mohavensis, 33, 34; Schottii, eee
ec T. G., Three new pia es of
Cuscuta from Mexico, 107
Zannichellia raped 35
ea nae 376
Ca rin,
271
Zythia, 164, 173; resinae, 163, 164, 173
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