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BROOKLYN BOTANIC GARDEN
MEMOIRS
VOLUME |
DEDICATION PAPERS
SCIENTIFIC PAPERS PRESENTED AT THE DEDICATION OF THE
LABORATORY BUILDING AND PLANT HOUSES
APRIL 19-21, 1917
ISSUED JULY 6, 1918
BROOKLYN@N. Y.; U.S. A.
PRESS OF
THE NEW ERA PRINTING COMPANY
LANCASTER, PA.
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PREFACE af
The papers contained in this volume were presented on April 20-
21, 1917, either in person or by title, at the Scientific Program, which
formed part of the Dedication Exercises on the occasion of the com-
pletion of the Laboratory Building and Plant Houses of the Brook-
lyn Botanic Garden.
ORLAND E. WHITE,
Epcar W. OLIVE,
ALFRED GUNDERSEN,
Publication Committee
iti
-
ae
CONTENTS
PAGE
femmnsON, GEO. F. The genus Endogone.:~..... 2 2. ee ek I
BLAKESLEE, A. F., and B. T. Avery, Jr. A vegetative reversion
MMM ON. De eg, ee 18
Britton, N. L. The flora of the American Virgin Islands..... 19
Burns, GEorGE P. Weather conditions and plant development. 119
oon, Met T.. Modern applications of botany ............. 123
DopcE, B.O. Studies in the genus Gymnosporangium—l. Notes
on the distribution of the mycelium, buffer cells, and the
Beweination Of the aecidiospore. ....:...2.6.20:-es- 684s: 128
East, E. M. Intercrosses between self-sterile plants.......... 141
Harrer, R. A. Binary fission and surface tension in the de-
Belopment ofthe colony in Volvox....2. ie =. s.5 4s as 154
Harris, J. ARTHUR. Further studies on the interrelationship of
morphological and physiological characters in seedlings of
RIE. eee os ge ee coolers maaan am 167
HARSHBERGER, JOHN W. American heaths and pine heaths.... 175
Ho.iick, ARTHUR. Some botanical problems that paleobotany
Ree Wem LOrSGlVe.. . 2... ae ee he ee 187
Howe, MarsHatyt A. Further notes on the structural dimorph-
ism of sexual and tetrasporic plants in the genus Galaxaura.. 191
Daemon, H.S. The. Uredinales of Oregon.~ 2.) 4........52. 198
JEFFREY, Epwarp C. Evolution by hybridization............ 298
-KunkKEL, L. O. A method of obtaining abundant ees in
cultures of Macrosporium solani E. & M. ete ae . -306
MACFARLANE, JOHN MUIRHEAD. Sonchvontsat in i lang structures 313
Metcatr, HAvEN. The problem of the imported plant disease
aqillustrated by the white pine blister rust...) 7 22 2. <-. 32
MurriLy, WitiiAM A. The rosy-spored A garics of North America 334
OLIVE, EDGAR W. The cytological structure of Botryorhiza Hip-
PML Sos ss Snes ayer 2 Ss Space ots SEE: tent ee 337
OsTERHOUT, W. J. V. The nucleus as a center of oxidation.... 342
REED, GEORGE M. Physiological specialization of parasitic
2 I SS eS, ae 348
RowWLeEE, W. W. Relation of marl ponds and peat bogs....... 410
SHEAR, C.L. Pathological problems in the distribution of perish-
BN lA EMP ORGCS ee ten. fei os OREN es ly vn ase eo 415
STEWART, F.C. Tubers within tubers of Solanum tuberosum... 423
Ve
vi CONTENTS
SHULL, GEORGE HARRISON. The duplication of a leaf-lobe factor
in the ‘shepherd’ s"purse. .... 0... 0: Ween.” ghee eee 42
SINNOTT, EDMUND W. Isolation and specific change.......... 444
SmitH, Erwin F. The relations of crown-gall to other over-
stowthsan plants...0. 05.5. 0..2. ESE eeee eee 2 oe 448
Stone, GEORGE E.. ‘Contact stimulation =e -o—— ..).- ee 454
Strout, A. B. Duplication and cohesion in the main axis in
Cichoreum Intybus. ... i. SPR SE ee be 480
TayLor, NorMAN. A quantitative study of Raunkiaer’s growth-
forms as illustrated by the 400 commonest species of Long
Island, Ne -Y.. . sso Sees Fee + 486
TRELEASE, WILLIAM. The ancient oaks of America...... ..a@2
TRUE, RopNney H., and Harvey, R. B. The absorption of cate
cium salts by sauaen stedlinigs, 20 see elas. os 502
WHITE, ORLAND E. Inheritance studies on castor beans....... 513
THE GENUS ENDOGONE
GEO. F. ATKINSON
Cornell University
The genus Endogone was founded by Link in 1809, and for more
than a century its life history and taxonomic relationship have re-
mained very obscure. Notwithstanding this obscurity in relationship
and development, the structure of the mature plants is so simple and
characteristic, that comparatively few species have been accredited
to the genus which do not belong here. The fruit bodies are pulvinate,
rounded to reniform or irregular. In size they vary from 2-4 mm.
up to 2-3cm. In life relation they are saprophytes. In habitat they
are hypogeous, epigeous or epixylous; and occur under or on the
ground, among or underneath fallen leaves, among the rhizoids of
mosses or roots of ferns, on dead wood, on sphagnum or other mosses.
In color they are whitish to yellow, brown or blackish. Approximately
seventeen species are known. The genus is cosmopolitan in distri-
bution. Species are known from Europe (including European Russia),
Ceylon, Tasmania, North America, Central America, South America
and the Fiji Islands.
General structure of the fruit body.—There is a peridium of slender,
interwoven hyphae formed by the terminal branchlets of the hyphae
which spring from the basal region, branch profusely and radiate to
the periphery. The interior constitutes the ‘“‘gleba,’’ the base or
central portion of the ‘‘gleba’’ is sometimes hollow or of less density
than the broad peripheral zone. The “gleba”’ is usually packed with
numerous, large thick walled ‘resting spores,’’ oval, elliptical, or
spherical in form, and yellowish, grayish or dark brown in color
according to the species. These ‘‘resting spores’? are 40-100 mw in
diameter, are packed among the hyphae, and terminate certain
branches. They have been termed spores, sporangia, ampullae, or
asci, the latter term apparently having the preference, since it is
employed by a majority of writers. Several large elliptical spores in
an ‘‘ascus’’ have been described in one species (Endogone pisiformis),
a single large spore! in an ascus in another species (E. reniformis),
and numerous minute spores in an ascus in several species. It is
iad
. 1The spores in this species are probably the ordinary “resting spores,’’ the
‘“‘asci’’ of authors, for Bresadola (1896, p. 297) says that neither the asci nor their
mode of dehiscence was seen.
2 1
2 BROOKLYN BOTANIC GARDEN MEMOIRS
doubtful, however, if true sporulation has ever been observed. In
dead resting spores the contents often segregate into a number of
large, rounded or elliptical bodies. In F. rentformis the single spore
in an “‘ascus” is probably the “resting spore’’ itself, while in the
species described as having numerous minute sporidia in an ascus
(often with an interrogation), the supposed sporidia are probably the
numerous fat bodies conspicuous in some species.
ORIGIN AND DEVELOPMENT OF THE ‘‘ RESTING SPORES’’ IN
ENDOGONE LACTIFLUA BERK.
Character of the mycelium.—The only important contribution to
development in the genus Endogone, thus far, was made by Bucholtz
in 1912, in his study of Endogone lactiflua. This species is subter-
ranean and occurs in various parts of Europe. ‘The fruit bodies range
in size from 4 mm. to 2 cm. The material studied by Bucholtz was
found in a plantation of Abies sibirica in Livland, Russia. The
mycelium is coenocytic, profusely branched, the hyphae following a
very sinuous course, but the general direction is radial and toward the
periphery. It is non-septate, true cross walls being formed only in
connection with the reproductive organs, though false cross walls are
occasionally found in the vegetative hyphae. The hyphae vary
greatly in diameter, in general becoming more slender toward the
periphery of the fruit body, but irregularities in the form of swellings
occur. Terminal branches on the interior are often clavate. Sack-
like enlargements occur from which numerous branches often diverge.
The walls of the hyphae are thick. The nuclei are minute, very
numerous, and lie in a parietal layer of cytoplasm.
The progametes and conjugation.—The progametes are clavate and
of unequal size. They lie nearly parallel and their walls fuse at the
lateral point of contact a short distance from the free ends. The
nuclei in the progametes, lying near the periphery become larger,
more distinct, and undergo one division. The nuclei now retreat from
the terminal portion of each progamete and all except one are excluded
from each gametangium by a cross wall. A few sterile nuclei some-
times remain in the gametangium but soon degenerate. The origin
of the selected gamete nucleus in each gametange is not known, but
it is probably derived from one of the peripheral nuclei in the pro-
gamete, or one of the daughter nuclei after mitosis. These gamete
nuclei are much larger than the progamete nuclei, thus maintaining
the nucleo-cytoplasmic relation. Each one occupies the center of its
gametange. The walls at the point of contact are now resorbed.
The nucleus from the smaller gametange (antherid) migrates into the
larger one (oogone).
ATKINSON: THE GENUS ENDOGONE 3
Formation of the ‘‘ resting spore’’ and simple zygocarp.—The “resting
spore” in Endogone lactiflua is not formed in the immediate zygote
resulting from the fusion of the two gametangia nor in the oogone,
but in an outgrowth from the latter. During and immediately after
fusion of the two gametangia their walls become thickened and firm,
so that they can not yield to the pressure from the young growing
zygote. As a result there arises a sack-like outgrowth from the end
of the oogone into which the cytoplasm from the two gametangia flows
accompanied by the two gamete nuclei, the antheridial nucleus fol-
lowing the oogonial nucleus. The sac-like outgrowth enlarges into
an oval or broadly elliptical resting zygote. A thick, stratified,
hyaline, cartilaginous wall is formed next to the primary zygote
membrane, which entirely encloses the cytoplasm and other contents,
thus separating them from the empty oogone. The two gamete
nuclei lie side by side in the center of the zygote but do not fuse until
after the resting period, except in a small variety from Germany in
which the gamete nuclei fuse at once according to Bucholtz. During
the growth and ripening of the zygote it becomes enveloped by slender
branches which coil in a more or less spiral manner around it forming
a thick cover of small cells, 2-3 cell layers deep, the walls of these
cells become greatly thickened and fuse next the zygote, grading out
to the thin walls of the surface. Each resting spore, or zygote, with
its individual cellular envelope forms a simple fruit or simple zygocarp
(zygosporocarp, as Bucholtz terms it). The fruit body of Endogone
lactiflua is filled with these simple zygocarps intermingled with the
mycelium, and is therefore a compound zygocarp. Germination of
the “resting spores’? has not been observed. Endogone lactiflua is
the only species of the genus in which such simple zygocarps are known,
1. €.,a fruit body with a single zygote enclosed in its individual envelope.
ENDOGONE SPHAGNOPHILA
In July, 1916, a day or two before the close of a fungus foray
organized by Mr. F. C. Stewart at his camp on Seventh Lake, in the
Adirondacks, Mr. W. H. Sawyer, Jr., a member of the party, brought
in some sphagnum on which were rounded, pulvinate, orange-yellow
bodies resembling the plasmodiocarps of certain slime molds. A pre-
liminary examination of the internal structure revealed the fact that
it was not a slime mold, but apparently a phycomycete with large
resting spores having a thick, hyaline, stratified, cartilaginous wall,
and orange-yellow content. A pair of stalks, or suspensors, attached
to one end of the resting spores in different stages of development,
indicated that they had their origin in an interesting type of conjuga-
tion. In the afternoon of the same day (July 31, 1916) Mr. Sawyer
4 BROOKLYN BOTANIC GARDEN MEMOIRS
and I crossed the lake and visited the same spot in order to collect
more material. The dry weather during the latter part of July had
lowered the water in the ravines so that this particular sphagnum
moor was water-free although the ground was very soft and wet.
The fruit bodies of the fungus were not very abundant, but here and
there a single one was found on a sphagnum plant, rarely two or more.
In nearly all cases the fruit body was attached on the upper side of
the central part of the terminal rosette, or one of its radiating branches.
Rarely was a fruit body found attached to one of the lower branches.
Altogether some 30 or 40 fruit bodies were collected. A number were
fixed in Flemming’s solution, some in Biondi’s solution and some in
chrom-acetic solution. Other material was carried to Ithaca on the
living sphagnum, where a few more fruit bodies were fixed. Some
were kept during the winter in moist situations out of doors, and
others in doors in a dried state. Finally, during the winter of 1916—
17, it was revealed to me in a semi-vision, that this fungus was a
member of the interesting genus Endogone.
Structure of the fruit bodies, or complex zygocarps.—The plants are
2-4 mm. in diameter, pulvinate, concave below and convex above, so
that a section through the center parallel with the morphological axis
is reniform. The larger plants are slightly convoluted or mildly
lobed, the upper portion showing two to three broad, low convolutions.
As the resting spores mature the plants are orange-yellow in color,
but the pigment resides entirely in the spore content, the mycelium
and spore walls being hyaline.
The peridium is thin, white, and composed entirely of a dense,
pliant weft of the terminal, slender branches of the radiating mycelium.
The terminal branchlets are 3-5 w in diameter at.the base and taper
out to a very slender point 1 uw or less in diameter. The walls are
much thickened, so that the lumen of these narrow branchlets is
nearly closed, quite so toward the apex. Many of these slender
branchlets are free above the surface and give to the peridium a
minutely tomentose, felt-like surface. Many of these branchlets arise
very close together, and then are more or less dichotomously branched
at a distance, a peculiarity often quite characteristic of the stouter
internal mycelium.
Internal mycelium and hold-fast.—The internal mycelium has a
general radial direction from the basal depression, diverging in all
directions toward the peridium. The main hyphae are 12-15 in
diameter. The branching is di- or trichotomous, or 4 to 5 or more
branches arise close together, their point of origin often suggesting a
ganglion-like enlargement of the parent hyphae from which the
branches radiate. The course of the hyphae is more or less sinuous.
ATKINSON: THE GENUS ENDOGONE 5
No cross walls have been observed, except in the progametes after copu-
lation. The nuclei are minute, very numerous and lie in the peri-
pheral granular cytoplasm. There is a nuclear membrane and a
large nucleolus (?karyosome). The fruit bodies are quite firmly
attached to the living sphagnum leaves, but the mycelium does not
appear to be parasitic, although short haustoria have been seen pene-
trating the cell. The hold-fast is a rather compact lattice-like layer
of mycelium forming a kind of ‘‘sole,’’ very closely applied to the
surface of the leaves, from which here and there the short haustoria
arise. The fungus is probably nourished by organic and mineral
solutions carried by the sphagnum from the water of the humus
substrate in the capillary stream so well provided for in the peat
mosses.
Conjugation of the progametes.—While there is a great resemblance
in the process of conjugation and in the formation of the resting zygote
of Endogone sphagnophila to the situation in E£. lactiflua, the details
of the process are quite different in the two species. The progamete
branches lie nearly or quite parallel. In a few cases where they have
been observed just prior to conjugation they do not appear to be
differentiated from ordinary stout vegetative branches, except that
the cytoplasm is more dense and abundant. They do not appear to
_ be enlarged or clavate. In fact many of the vegetative branches are
clavate and sometimes they are in pairs lying closely side by side,
but in no case have I been able to determine with certainty that such
branches are progametes. The progametes also appear to be un-
differentiated before conjugation. They conjugate by lateral contact
of their walls at the tip. Immediately after contact the progametes
begin to swell into a clavate or fusoid form, and the wall at the point
of contact is resorbed for some distance, thus forming a broad com-
municating area where the cytoplasm of the two merges. During the
enlargement one of the gametes frequently becomes larger than the
other. The cytoplasm is very dense and fills the distal portion of the
progametes, while in the proximal direction the cytoplasm is less
abundant and lies chiefly in a peripheral zone next the wall. A cross
wall is now laid down in each progamete a short distance behind the
broad communicating pore, separating the gametangia from the stalks
or suspensors.
Formation of the resting spore or resting zygote.—At the time of
conjugation and resorption of the contact wall the conjugating game-
tangia resembles the same stage of conjugation in Eremascus fertilis.
The zygote is not formed by the enlargement of the copulating game-
tangia as in the majority of the Mucorales, but the young zygote
begins to grow at once in an apical direction. Sometimes the origin
6 BROOKLYN BOTANIC GARDEN MEMOIRS
of the young zygote is symmetrical in relation to the two gametangia,
that is, the tip of each gametange shares equally in the growth. In
other cases the new growth arises more from one than the other,
usually from the larger one where they are unequal in size. More
rarely does the new growth arise entirely from the larger gametange,
but the communication is so broad that both gametangia remain in
direct communication with the contents of the new growth. This new
growth, or progressive zygote, enlarges to a broadly elliptical structure,
35-60 x 30-45 uw. It stands on the two supporting gametangia, and
the protoplasm of the gametangia and new zygote is continuous.
When the new zygote has reached its full size the protoplasm in the
gametangia withdraws and merges with that in the zygote. A new
wall is now laid down inside of the primary zygote membrane. At
first thin, it increases in thickness, forming a white, stratified, carti-
laginous layer around the protoplasm, thus cutting off communication
with the empty gametangia. The two stalks which support the mature
resting zygote are not simply the suspensors, but the empty game-
tangia plus the suspensors. The resting zygote is nearly filled with
very minute rounded or slightly irregular hyaline bodies, which appear
to be fat bodies, since they stain red with Sudan III. There is a
rounded clear space in the center, 7. e., in the middle of the long axis,
but in some zygotes it lies on one side next the wall. In fixed and
stained material the center of this vacuole (?) appears to be occupied
with a coarsely granular body or mass of minute bodies.
Cytology of conjugation and zygote formation.—The number of
nuclei in the gametangia ‘is variable, probably from five to ten or more
in each. No evidence of nuclear division in the progametes or game-
tangia has been observed, and no evidence has been seen of a selection
of gamete nuclei. Nor does it appear that there is any nuclear de-
generation in the progametes before the formation in the cross wall
which differentiates the gametangia. Following this stage nuclei in
the suspensors may degenerate. The cytoplasm in the gametangia
is so dense and stains so deeply that it is sometimes difficult to differ-
entiate the nuclei. When the stain is not too deep the nuclei are
clearly seen. They are considerably larger than the vegetative nuclei,
the increase being due to growth. There is a nuclear membrane, a
clear court in which are sometimes visible a few delicate threads, per-
haps portions of the linen or chromatin. There is a prominent central,
spherical, nucleolus or karyosome, which stains red with Flemming’s
triple stain, dark with iron haematoxylon. The nuclei are disposed
in the cytoplasm of the gametangia without order. They gradually
migrate into the new zygote, as it is formed, along with the cytoplasm.
In the young zygote the cytoplasm is at first dense and rather
ATKINSON: THE GENUS ENDOGONE 7
homogeneous as in the gametangia. But as the new zygote enlarges
the cytoplasm becomes coarsely reticulate. The strands are coarse
and with an irregular outline. They radiate irregularly from the
center to the periphery and anastomose by irregular branches, forming
a large meshed network the strands of which are coarser in the central
region, thinner toward the periphery. During the early stages of
development of the young zygote the nuclei appear to have a general
distribution, but have not been observed near the periphery. As the
zygote approaches its full size the nuclei occupy the more central
region, being distributed from the center to a zone half way, or a
little more, to the periphery. Sometimes the centralization is more
marked. At this stage there appears to be a differentiation of the
cytoplasm, or rather, the appearance of a clear homogeneous plasma
occupying the nuclear region and in which the nuclei die. When the
nuclei are strongly centralized, the homogeneous plasma appears to
form a single large central area. When they are more widely dis-
tributed, the homogeneous plasma is separated into several areas,
each area containing several nuclei. During all this period the coarsely
reticulate cytoplasm occupies the entire zygote.
A provisional suggestion as to the function of this homogeneous
plasma is that it serves as a medium for the freer movement of the
nuclei than can take place in the coarsely reticulated cytoplasm; or
the homogeneous plasma may actually serve to move the nuclei to a
certain extent, possibly bringing them into closer proximity in the
center and later carrying them into the peripheral zone. In this
central region the nuclei are generally in pairs and the two nuclei of a
pair appear to fuse. This fusion of paired nuclei in the zygote corre-
sponds with the fusion of paired nuclei in certain of the Mucorales as
described by Gruber (1901) in Sporodinia grandis, Dangeard (1906) in
Rhizopus nigricans, and Moreau (1911-1913) in Rhizopus nigricans
and species of Zygorhyncus. The fusion of the nuclei of a pair is
suggested by the fact that they are found close together, sometimes
the nuclear membranes in contact, again an elongated nucleus (?
fusion) with one membrane and this constricted between the two
nucleoli, and further two nucleoli surrounded by one membrane which
is not constricted. Of course these relations might be the result of
nuclear division, but no figures appear at this stage which suggest a
division of the nuclei.
In a later stage of development the coarse reticulum of the cyto-
plasm disappears. The cytoplasm becomes more homogeneous.
The nuclei appear to occupy a zone about half way between the
“center and periphery. Figures are present which suggest a division
of the nuclei at this stage, for the nuclei are often two to three times
§ BROOKLYN BOTANIC GARDEN MEMOIRS
longer than broad, presenting the appearance of rods, deeply stained,
lying in a peripheral zone of the cytoplasm.
The material on which this cytological study was made was not
as well fixed as it might have been. The peridium of the fruit bodies
is very dense and tough, not permitting the rapid penetration of the
fixing fluids. Then in the later stages of zygote formation the thick
cartilaginous wall of the resting zygote very likely offers great resist-
ance to the penetration of the fluids. An attempt will be made
to collect more material during the present season, when the fruit
bodies will be cut open before placing them in the fixing solutions,
and also it is hoped that younger stages of development may be
secured.
Up to the present time no ‘one has succeeded in germinating the
resting zygotes of any species of Endogone. Link (1809), Fischer
(1897, p. 121, 124) and Bucholtz (1912) have described sporulation
in the ‘‘resting spores’’ (azygotes) of Endogone pisiformis, a partheno-
genetic species. In this species the wall of the resting spore is only
slightly thickened. According to Fischer and Bucholtz the content
of the resting spore is gradually divided into angular areas which
round up and form a number of large elliptical spores inside the wall
of the resting spore (or ? sporangium). Their study was not made on
living material, but on specimens preserved for several years. There
was no intersporal substance or epiplasm.
I have made several attempts to germinate the resting zygotes of
Endogone sphagnophila, but thus far without success. The first
attempts were made in December, 1916, with material kept on sphag-
num under cover of a bell jar in the shade on the north side of a
building. The cultures were made by tearing out mats of mycelium
with the resting zygotes in a thin layer of water on glass slides which
were kept in moist chambers. The cultures were examined day by
day for a period of two weeks. These cultures were then allowed to
remain out of doors on a window ledge with a southern exposure until
the middle of January, 1917, when they were brought inside and again
examined daily for a period of a week. During the latter part of
March and early in April, 1917, fresh cultures were started from the
same source, 7. e., from fruit bodies kept on sphagnum out of doors,
where they were subject to freezing and thaw. Thus far (Apr. 17,
1917) there has been no evidence of germination, although the great
majority of the zygotes appear to be alive and in good condition.
A few of the zygotes, however, appear to be dead. In many of these ~
the content is divided into irregular bodies. Others are filled with
elliptical or globose bodies, in some instances with intersporal sub-
stance. ‘These bodies, some of them, at least appear to be spores,
ATKINSON: THE GENUS ENDOGONE |
but I am strongly inclined to believe that they are spores of some
parasite.
RELATIONSHIPS OF ENDOGONE
The coenocytic mycelium with no true cross walls, except those
which separate reproductive organs from the mycelium, with the
formation of resting zygotes soon after the conjugation of gametangia,
are phycomycete characters. The method of conjugation of equal or
slightly unequal gametangia indicates a closer relationship with the
Zygomycetes than with the Oomycetes, although in Endogone lactiflua
the content of the small gametange, or antherid, flows into the larger
one which is comparable to the oogone. In Zygorhyncus the two
gametangia are very unequal in size, but the zygote is formed within
and includes all of both gametangia, a strictly zygomycete feature.
In Conidiobolus the gametangia are of very unequal size, and are
worthy of being distinguished as antherid and oogone. The content
of the antherid passes into the oogone within which the zygote is
formed. In Basidiobolus* also there is a supplying gametange and a
receiving one, but other features of these genera, especially conidial
formation, show such a close relation to Empusa that they are generally
regarded as members of the zygomycetes with a leaning toward the
oomycetous type of fructification, but not having reached the char-
acteristic feature of egg differentiation in the oogone. Endogone
departs from the usual type of zygote formation present in the zygo-
mycetes. The zygote is an outgrowth from the conjugation point of
the gametangia (E. sphagnophila), or from the larger gametange
(LE. lactiflua, rarely in E. sphagnophila). A similar situation, however,
is present in Piptocephalis freseniana and in Empusa (Entomophthora)
fresenii, where the zygote is an outgrowth of the point of conjugation
much as in Endogone sphagnophila. In Empusa occidentalis, echino-
spora, sepulchralis, etc., the zygote is an outgrowth of one of the
gametangia, a situation similar to that in Endogone lactiflua, or some-
times it arises from the point of conjugation in these species of Empusa.
In the selection of a single sex nucleus in each gametange, EL.
lactiflua presents a situation similar to that in the Peronosporales,
though there is no differentiation of ooplasm and periplasm in the
oogone of Endogone lactiflua, as there is in the Peronosporales. In E.
sphagnophila there is no sex selection of nuclei in the gametangia so
far as we can determine, for all of the nuclei without manifesting any
differentiation pass with the cytoplasm into the new outgrowth where
the zygote is formed. The nuclear behavior in the zygote has not
yet been made entirely clear. The present evidence suggests that
2 Basidiobolus by some is placed in a distinct family.
10 BROOKLYN BOTANIC GARDEN MEMOIRS
there is first a nuclear fusion in pairs, perhaps some of the nuclei
degenerating. If this is confirmed the situation in EL. sphagnophila
agrees in this respect with that in the Mucorales. It appears also that
later, when the nuclei lie in a peripheral zone, they undergo at least
one division. At the present stage of the investigation the possibility
is not excluded that fusion of nuclear pairs does not occur. It is also
possible that after the division of the nuclei in the peripheral zone of
the zygote all but two may degenerate, the two selected ones later
uniting to form the fusion nucleus. This would bring E. sphagnophila
more nearly in line with the process in EL. lactiflua as described. by
Bucholtz, the selection of the sex nuclei being postponed to a late
period in E. sphagnophila. I do not think, however, that this is the
case, but am inclined rather to believe that there is multinuclear fusion
in pairs,® similar to that which takes place in the Mucorales as de-
scribed by Gruber (1901) in Sporodinia, by Dangeard (1906) in Sporo-
dinia, and by Moreau (1911-1913) in Rhizopus, Zygorhyncus and
Sporodinia. That in certain species of Endogone there is fusion of
but one pair of sex nuclei in the zygote, while in other species there
may be fusion of several pairs of sex nuclei, is not incomprehensible
in view of the nuclear process in fertilization in Cystopus (See Stevens
1899, 1901), where in C. blitt and portulacae there are fusions of many
pairs of sex nuclei in the egg, while in other species there is fusion of a
single pair of sex nuclei.
There is another feature in E. lactiflua which is paralleled in
certain of the Zygomycetes. This is the hyphal envelope which
encloses each zygote. Crude tendencies to such an envelope are
present in Phycomyces and Absidia of the Mucorales and in Empusa
rhizospora of the Entomophthorales (Thaxter, 1888), while in Mor-
tierella there is a well-developed envelope. In no other species of
Endogone, however, is such an envelope around each zygote known,
not even a rudimentary one. In this respect £. lactiflua represents a
more advanced stage of evolution, which is manifested also in the
origin of the resting zygote as a distinct outgrowth of the larger
gametange. This species may possibly represent the type of a distinct
genus, so widely does it depart in these two respects from all the
other known species.
There is another feature, however, in which Endogone departs
widely from any other known phycomycete. The mycelium and
numerous zygotes (“‘resting’’ spores or ‘‘sporangia’’ in the partheno-
genetic species) are united into a compact and distinct fruit body, or
’ According to Leger (1896) in Sporodinia grandis, and according to Miss McCor-
mick (1912) in Rhizopus nigricans, all nuclei but two degenerate in the zygote, but
this has not been confirmed.
ATKINSON: THE GENUS ENDOGONE 1]
zygocarp with a definite and well-developed peridium. This repre-
sents a distinct progression in development over all other phycomy-
cetes, a cephalization of zygotes into a complex fruit body.
The heterogamous character of the gametangia of Endogone lacti-
jflua and the selection of a single gamete nucleus in each are oomycete
features. But the lack of differentiation in the cytoplasm in the
oogone, or gametangia, is a zygomycete feature. For these reasons
Bucholtz* interprets Endogone as occupying an intermediate position
between the zygomycetes and Oomycetes, but constituting a distinct
group, the Endogoneae.® He probably regards this intermediate posi-
tion as simply taxonomic, not phylogenetic.
RELATION OF ENDOGONE TO THE ASCOMYCETES
Endogone has been shifted in all the three great divisions of the
fungi. It was first placed in the Basidiomycetes near Rhizopogon by
Link (1809) who was followed by Fries (Syst. Myc. 2, 295, 1822).
For a long time it has remained in the Ascomycetes, being placed in
the Tuberaceae by Vittadini (1831), by Tulasne (1857), by Saccardo
(Syll. Fung. 8, 905, 1889). Schroeter (1889) placed Endogone with
some uncertainty in the Order Protomycetes, the highest order of the
Phycomycetes. He was followed by Saccardo (Syll. Fung. 14, 829,
1899), and it is significant that the genus Protomyces has by many
students been placed in the Phycomycetes. In 1897 Schroeter, while
still retaining Endogone in the Protomycetaceae, transferred the group
to the Hemiascineae.
Until we know the morphological and cytological phenomena in
connection with the germination of the resting zygotes of Endogone
we cannot say with any degree of precision what relation it bears to
the Ascomycetes, nor how near that relation is. It appears quite
probable that Endogone does bear an interesting relation to the
Protoascomycetes. If the resting zygotes germinate fructificatively
with free cell sporulation, somewhat as occurs in Dipodascus or
Protomydes, its relation to the Protoascomycetes would be very clear.
The question would then arise whether with its coenocytic mycelium
it should be placed on a level with Dipodascus or just below it, repre-
senting the highest level of the Phycomycetes. Even if the germina-
tion phenomena should prove to be of the phycomycete type, Endo-
4 He describes two large nuclei in the zygote of E. ludwigii, a sexual species.
®> The genus has been considered for a long time by a number of students to
represent a distinct family. Fries (Summa Veg. Scand. 1849) proposed the
family Endogonaceae and in view of Bucholtz’s studies it is interesting to note that
Schroeter in 1889 placed it along with Protomyces as the highest member of the
Phycomycetes.
12 BROOKLYN BOTANIC GARDEN MEMOIRS
gone would still represent the nearest approach of the phycomycete
type to the ascomycete type. The method of conjugation of the
gametangia, and the growth of the zygote, in Endogone sphagnophila,
is surprisingly like that in Eremascus fertilis and in Dipodascus. Even
without the knowledge of germination of the resting zygotes in Endo-
gone, the genus seems to offer more of the characteristics of a prototype
of the Protoascomycetes (and perhaps also of the Uredinales) than
any other known phycomycete.’ Endogone presents additional strong
evidence of the phycomycete ancestry of the Ascomycetes.
All of the evidence considered, it appears to point more strongly
to the zygomycete alliance as the source of the primitive ascomycete
stock, rather than to the oomycete alliance. In the oomycetes the
sexual organs and the processes of fertilization have become very
highly specialized. The sexual organs are highly differentiated; one
or more distinct eggs are differentiated in the oogone, in many cases
the protoplasm being differentiated into ooplasm and_ periplasm;
while a special fertilization tube from the antherid penetrates the
oogone, or in a rare and specialized case a motile sperm enters the
oogone through a pore (Monoblepharis).
In the zygomycetes the sexual organs have retained a simple and
generalized condition. Copulation is by pore formation with merging
of the content of the gametangia. In most cases the gametangia are
equal and the zygotes mature in situ, within and comprising all of
both gametangia. Progression in the zygomycetes, however, is mani-
fested in five directions.
1. Ina tendency to differentiation in size of the gametes.
2. A tendency to differentiation of the gametangia in function,
the larger one becoming the receiving gametange, the oogone, but
without differentiation of content into egg and periplasm; the other
serving as the supplying gametange, antherid (Conidiobolus utriculosus,
Basidiobolus ranarum, Dispira americana Thaxter, 1895, Endogone
lactiflua, etc.).
3. The progressive tendency shown in the germination, or out-
growth, of the young zygote immediately after fusion from the copu-
lation point, or from one of the gametangia, the ripe zygote being formed
outside the gametangia, not within them according to the typical
process. The tendencies in this direction appear at different levels
in the zygomycete alliance. Examples are Piptocephalis freseniana,
° The following forms among the phycomycetes have been suggested as proto-
types of the ascomycetes. The Peronosporales by de Bary (1881) for the usual
ascomycete type, Piptocephalis for the Eremascus type; Myzocytium and Protascus
by Dangeard (1903-06, 1910), Cystopus by Lotsy (1907) and Monoblepharis by
Nienburg (1914).
ATKINSON: THE GENUS ENDOGONE 13
Empusa (Entomopthora) fresenii, Empusa rhizospora, sepulchralis, etc.,
Endogone sphagnophila and lactiflua.
4. Progression in the direction of selection of sex nuclei. In
Endogone lactiflua one sex nucleus is selected from among all the
potential ones in each gamete. This illustrates how the situation in
Dipodascus, Eremascus fertilis and in Endomyces has arisen. This
situation is presaged in some of the lower zygomycetes in the degener-
ation of some of the gamete nuclei in Rhizopus, Sporodinia and Zygo-
rhynchus, if we may accept the account by Moreau (1911, I913).
In Endogone sphagnophila there are many potential and many func-
tional nuclei in the gametangia and these are carried on into the
young germinating, or progressive zygote. This type illustrates a
situation which has been retained in those ascomycetes with multi-
nucleate gametangia as in Monascus, the Gymnoascaceae, A scodesmis,
Pyronema, etc.
5. The postponement or moving forward of the moment of nuclear
fusion from the gametangia to the new outgrowth, or progressive
zygote. The examples are the same as those given in paragraph 3.
In the zygomycete alliance Endogone lactiflua represents the most
progressive stage in these directions of any known species, unless
Dipodascus should be regarded as a phycomycete. The sex pair of
nuclei is organized by the migration of the antheridial nucleus into
the oogone. The pair then migrates into the new outgrowth where
the sex nuclei lie side by side in the resting zygote, or fuse, in one
variety. As Bucholtz (1912) points out there is wanting here only
the conjugate division of the nuclear pair to parallel the situation in
the ascogenous hyphae of the true ascomycetes.
From the situation reached by progression in these directions, by
members of the zygomycete alliance, there is but a small gap over
which to bridge in reaching the protoascomycetes. The principal
steps may be indicated as follows:
1. Free cell sporulation of the progressive zygote of Endogone.
This alone would place this zygote in the category of a generalized
ascus, and Endogone would become a fit member of the protoascomy-
cetes. Intersporal substance present in the sporangium of some
phycomycetes during sporulation may possibly presage typical free cell
sporulation. Other steps toward the true ascomycetes are indicated
in the following.
2. The omission of a period of rest by the progressive zygote and
the immediate free cell sporulation of the same. This step alone
would reach the level of Dipodascus, Eremascus, Endomyces magnusit,
etc.
3. Vegetative growth of the zygote and postponement of free cell
14 BROOKLYN BOTANIC GARDEN MEMOIRS
sporulation until the second or third cell of the new growth, would
carry the zygote to the level of that in Sphaerotheca, in the true
ascomycetes.
4. Vegetative growth of the zygote with splitting of the same by
branching, thus multiplying the terminal branches of the zygote in
which free cell sporulation takes place, carries the zygote to the level
of Pyronema, Monascus, etc., in the true ascomycetes.
5. Organization of a fruit body by formation of a peridium of the
interwoven terminal branches of the mycelium. This is already
realized in the complex zygocarp of Endogone, the peridium being on a
level with that in the Gymnoascaceae.
6. Organization of a peridium by enveloping hyphal branches of
the type in Monascus, the Erysiphaceae, Aspergillus, etc. This feature
is already realized in the simple zygocarp of Endogone lactiflua.
RELATION OF THE ZYGOTES AND AZYGOTES IN ENDOGONE
Several species of Endogone are parthenogenetic (E. macrocarpa,
microcarpa, pisiformis, etc.), yet the ‘‘resting spores” are similar to
the resting zygotes of E. sphagnophila, ludwigii, etc., in all other
respects so far as known at present, with the exception of the number
of nuclei in the resting stage. According to Bucholtz the zygotes of
the sexual species, E. lactiflua and E. ludwigii, are binucleate in the
resting stage, while the azygotes of E. pisiformis, macrocarpa, micro-
carpa, etc., are multinucleate. However, the zygotes of the sexual
species E. sphagnophila, are probably multinucleate in the resting
stage. It is very probable that at the time of germination the zygotes
of E. lactiflua and E. ludwigit become multinucleate by successive
divisions of the fusion nucleus. It is, therefore, very likely that the
phenomena of germination, whatever the type of germination is for
Endogone, is the same in the sexual and parthenogenetic species. It is
quite evident that the branch in which the azygote is formed is the
morphological equivalent of a gametange, just as the sporangia and
gametangia of Monoblepharis, Saprolegnia, etc., are morphological
equivalents. If there were sex differentiation among the nuclei of
the zygotes followed by fusion, then the azygotes would be strictly
homologous with the zygotes, in fact they would be zygotes. Bucholtz
(1912) regards the simple zygocarp of Endogone lactiflua as homologous
with the germ sporangium (carposporangium) of the Mucorales and
Peronosporales. In a certain degree this is true, but it is also homo-
logous with the zygote of the Mucorales and Peronosporales. In the
Mucorales the germ sporangium is external to the zygote while in the
Peronosporales it is internal. The germ sporangium of Endogone is
not known, it may be internal or external to the zygote. The zygote
”
ATKINSON: THE GENUS ENDOGONE IS
of Endogone presages the beginning of a new structure, not more so
than does the zygote of Piptocephalis and of certain species of Empusa,
etc., but it is still homologous with the zygote of the other phyco-
mycetes.
In the Mucorales, aside from the meiotic divisions of the fusion
nuclei, the process of sporulation in the germ sporangium (sporangium
formed on germination of the zygote) is the same as that in the ‘‘asex-
ual’’ sporangium. The germ sporangium and the asexual sporangium
are morphological equivalents. The germ sporangium is not a new
morphological structure, though the zygote and early stages of its
germination presage the origin of a new morphological structure.
Historical experience turns it quickly back into the well-worn trail.
The very simple primordium of the new structure does not mark out a
new path until the accumulation of new experiences, together with
environment, provide the threshold for progress to the new structure.
If nuclear fusion does not occur in the azygotes of Endogone then
the chromosome history, so far as we know, would run from generation
to generation without change. The two situations may be repre-
sented thus:
Sexual species of Endo- Ix—> 1x—> Ix Ix 5 mae Ix— Ix etc.
gone. Mycelium and gametangia zygote | spores, mycelium.
Parthenogenetic species { Ix— Ix Ix Ix >Ix>| Ix Ix etc.
of Endogone. l Mycelium and gametangium | azygote | spores, mycelium.
As sporulating organs the zygote and azygote (or germ sporangium
of the same) of Endogone are homologous structures. The true asexual
sporangium has been eliminated. Likewise, in the protoascomycetes,
where the threshold from the phycomycetes has been crossed, the
zygote (‘germ sporangium,’’ a generalized ascus) of Dipodascus, and
the azygote (‘‘germ sporangium,”’ ‘‘generalized ascus’’) of Ascoidea,
as sporulating organs are homologous structures. Ascoidea is prob-
ably parthenogenetic, the ‘‘generalized ascus’’ being a transformed
gametangium. Free cell sporulation occurs in both genera., This
interpretation of the relation of the free cell sporulating organs of
Dipodascus and Ascoidea is supported by the situation in Hremascus
fertilis and Endomyces magnusit where the ascus is in some cases of
sexual origin, in others of parthenogenetic origin, a single gametange
becoming the ascus. Endogone, with several sexual species having
sexually produced sporulating organs (zygotes), and other species
with parthenogenetically produced sporulating organs (azygotes), is
interesting in that it illustrates the homology of these structures, and
suggests how the parthenogenetic sporulating organs (generalized
asci) of Ascoidea, Protomyces, etc., may have arisen. It is interesting
16 BROOKLYN BOTANIC GARDEN MEMOIRS
to note that the forms with parthenogenetic asci, generalized or not,
were not endowed with potentialities of progress, nor with the evolu-
tion of any important lines. They have made practically little progress
and are few in number. On the other hand, those forms with sexually
produced asci, even though the sexuality be of a very greatly reduced
type, were endowed with great potentialities as evidenced by the large
group of Euascomycetes with high specialization, and great divergence
of character in several different series.
TECHNICAL DESCRIPTION
Endogone sphagnophila’ n. sp. Plants (zygocarps), 2-4 mm. in
diameter, pulvinate, reniform, plain or subcerebriform with two to
three low lobes or convolutions, orange yellow when mature, Peridium
white, submembranous, tough, of interwoven coenocytic, profusely
branched hyphae, minutely tomentose or downy from free, terminal,
very slender branchlets, 4-5 » at base, 1 wor lessat the tips. Mycelium
of the zygocarp I0—-I5 w in diameter, coenocytic, stout, non-septate,
branched in a dichotomous, or trichotomous manner, or several
branches springing from enlargements, radial, the terminal branches
interlacing to form the peridium. Progametes equal. Gametangia
separated from mycelium by a cross wall, equal or usually slightly
unequal, multinucleate. ‘Resting spores’’ (zygotes) formed as an
outgrowth from the conjugation point of the gametangia, or more rarely
from the larger one, one resting spore formed in the primary zygote
membrane from each pair of gametangia, elliptical to oval, rarely
irregular, with orange yellow content and a thick, white, stratified
cartilaginous wall, 35-60 x 30-45 yw, germination unknown.
On sphagnum in a ravine in region of Seventh Lake, Fulton Chain,
Adirondack Mts., New York, July 1916, Aug. 1917; and in Cranesville
moor, Western Maryland, Sept. 1917 (rarely on other mosses or on
dead twigs). Thaxter (Bot. Gaz. 24: 12, 1897) reports it on sphagnum
in Maine.
Latin diagnosis. Pulvinatis, reniformibus, subcerebriformibus, aureis, 2-4 mm.;
peridiis albidis submenbranceis, lentis, floccosis intertextis, tomentosulis; myceliis
glebae 12-15 w, radiatis, dichotomis vel trichotonis vel plurichotomis, ramulis
terminalibus peridium formantibus; sporis orientibus ab gametangiis copulantibus,
ellipsoideis vel ovalibus, maximis, 35-60 x 30-45 », plasmate aureo. Hab. on sphag-
num, Adirondack Mts., New York, and in Maine and Maryland.
LITERATURE CITED
Atkinson, Geo. F. Phylogeny and Relationships in the Ascomycetes. Ann. Mo.
Bot. Garden 2: 315-376. Figs. 1-9. I914.
7 Closely related to E. ludwigit Bucholtz, but this species is subterranean with a
prominent germ pore in the thick wall of the zygote and the nuclei are reduced to
two which fuse at maturity.
ATKINSON: THE GENUS ENDOGONE 17
de Bary, A. Untersuchungen iiber die Peronosporeen und Saprolegnieen und die
Grundlagen eines natiirlichen Systems der Pilze. In de Bary und Woronin,
Beitr. z. Morph. u. Physiol. d. Pilze 4: 1-145. Pls. 1-6. 1881.
Brefeld, O. Die Hemiasci und die Ascomyceten. Untersuch. Gesammtg. Myk. 9:
1-156. Pls. 1-3B. 1891.
— Ascomyceten II. Ibid. 10: 157-378. Pls. 4-13. 1891.
Bucholtz, F. Beitrage zur Kenntnis der Gattung Endogone Link. Beihefte Bot.
Centr. 29: 147-225. Pls. 3-10. 1912.
Dangeard, P.A. Recherches sur le développement du périthéce chez les Ascomycetes.
Premiére partie. Les ancetres des champignon superieurs. Le Botaniste 9:
157-303. Pls. 1-18. 1903-1906.
— La fecondation nucléaire chez les Mucorinées. C. R. Acad. Sci. Paris. 1906.
— Recherches sur le développement du périthéce chez les Ascomycetes. Le
Botaniste 10: 1-385. Pls. I-91. 1907.
Fischer, E. Tuberaceen und Hemiasceen, in Rab. Krypt. Fl. 2nd Auf. Die Pilze
5th Abt. I-151. 1897.
— Pilze in Handworterbuch der Naturw. 7: 880-929. Ig12.
Gruber, E. Ueber das Verhalten der Zellkernc in den Zygosporen von Sporodinia
grandis. Ber. deuts. bot. Ges. 19: 51-55. Pl..2. Igor.
Léger, M. Recherches sur la structure des Mucorinées. Paris, Poitiers. 1896.
Hinks) ©. B.S: Pl. nat. 3: 33. 18009.
Lotsy, J. P. Vortrage iiber botanische Stammesgeschichte 1: I-FV and 1-828.
Heel ASO O07:
McCormick, F. L. A. Development of the Zygospore of Rhizopus nigricans. Bot.
Gaz. 53: 67-68. IgI2.
Moreau, F. Deuxieme note sur les Mucorinées. Fusions de noyaux et dégen-
erescence nucléaire dans la zygospore. Soc. Myc. France Bull. 27: 334-341.
IQIl.
Moreau, F. Sur la reproduction sexuée-de Zygorhynchus moelleri Vuill. C. R.
Soc. Biol. 73: 452-455. I9II.
— Les phénomeéenes intimes de la reproduction sexuelle chez quelques Mucorinées
hétérogames. Soc. Bot. France Bull. 58: 618-623. IgII.
— Une nouvelle Mucorinée héterogame Zygorhynchus dangeardi sp. nov.
Soc. Bot. France Bull. 59: LXVII-LXX. 1912.
— Recherches sur la reproduction des Mucorinées et quelques autres thallo-
phytes. Théses presentées a la Fac. Sci. Univ. Paris 1-136. Pls. 1-14.
1913.
— Les karyogamies multiples de la zygospore de Rhizopus nigricans. Soc. Bot.
France Bull. 60: 121-123. 1913.
Schroeter, J. Cohn’s Krypt. Fl. Schlesien. 3!: Pilze. Protomycetes. 257-260.
1886.
— Hemiascineae in Engler & Prantl. Pflanzf. 11: 143-149. 1894.
Stevens, F. L. The compound oésphere of Albugo bliti. Bot. Gaz. 28: 149-176,
225-245. Pls. 11-15. 1899.
— Gametogenesis and fertilization in Albugo. Bot. Gaz. 32: 77-98, 157-169,
238-263.) Pls. 1-4: F.. 10901.
Thaxter, R. New or peculiar American Zygomycetes. I. Dispira. Bot. Gaz. 20:
BUS 5LOn ie GlsmsAc. .TOO5.
Tulasne, L. R. et Ch. Fungi Hypogaei. Pp. I-XIX, 1-222. Pls. 1-21. 1851.
Vittadini. Monographie Tuberacées. 1831.
A VEGETATIVE REVERSION IN PORTULACA
(Abstract)
A. F. BLAKESLEE AND B. T. AVERY, JR.
Station for Experimental Evolution, Cold Spring Harbor, N. Y.
A single dwarf individual was discovered in a bed of plants from
commercial seed of Portulaca grandiflora. When the dwarf is selfed
it throws all dwarfs. Some of the dwarf offspring produce reverting
branches which differ from the dwarf stock upon which they are
borne by having red, instead of green stems and by having longer
internodes. The flowers on both dwarf stock and reverting branches
are red. Selfed seed from the reverting branches produces both
dwarfs with short internodes and normal plants with long internodes
as well as occasional dwarfs that show reverting branches.
18
THE FLORA OF THE AMERICAN VIRGIN ISLANDS
N. L. BRITTON
New York Botanical Garden
The islands St. Thomas, St. Jan and St. Croix recently purchased
by the United States from the kingdom of Denmark, are situated to the
east and southeast of the island of Porto Rico. My interest in their
flora was first aroused by the proximity of the Virgin Islands archi-
pelago to Porto Rico, both the Porto Rican islands Culebra and
Vieques being parts of the archipelago. I therefore took occasion in
1913, accompanied by Dr. J. N. Rose and with the help of Mrs.
_ Britton, Miss Delia W. Marble, and Dr. J. A. Shafer, to explore St.
Thomas and St. Jan quite thoroughly, and Dr. Rose made collections
on St. Croix, while we were engaged in studying the cacti of the West
Indies! In 1901, I had made a brief visit to St. Croix, with Mr.
John F. Cowell.?
The islands are all hilly, there being very little level land on either
St. Thomas or St. Jan, but more on St. Croix. The rocks are mostly
of plutonic origin, but there is some limestone on St. Croix and locally
other stratified rocks occur. The highest elevation is about 500
meters (1,550 feet), on The Crown of St. Thomas.
There is but little natural forest remaining on any of the islands,
and what there is is confined to the hilltops in a few places. Re-
forestation is the crying need of the new possession, and it will be
highly discreditable to the United States if this subject is not im-
mediately taken in hand. Most of the higher parts of all three islands
are not available for any but forest products and the supply of wood
for fuel needs to be increased and the rainfall conserved by a forest
cover, for most of the rain now runs off immediately. This destruc-
tion of the forest has doubtless eliminated a good many species from
the original flora of the islands.
The principal literature of the botany of the islands is as follows 33
H. West. ‘Bidrag til Beskrivelse over Ste Croix, med en kort udsigt over St.
Thomas, St. Jean, Tortola, Spanishtown og Crabeneiland.’’ Kiébenhavn.
Pp. 363. 1793. [German edition pp. 274, Copenhagen 1794.]
West enumerates and partly describes 542 species, of which 111 were culti-
1 Jour. N. Y. Bot. Gard. 14: 99-109.
2\ four. N. Y. Bot. Gard: 2: 166.
8 See also citations in the chapters on Hepaticae, Fungi and Algae.
19
20
André
BROOKLYN BOTANIC GARDEN MEMOIRS
vated, mainly of St. Croix, a few from St. Thomas and St. Jan. Some of
the descriptions are by Vahl. A number of the plants listed have not been
observed on St. Croix by subsequent collectors and some of them are obscure.
The author was rector of a school at Christiansted. The book is very rare.
I am indebted to The New York Public Library for a photostat copy.
Pierre Ledru. ‘Voyage aux iles de Ténériffe, La Trinité, Saint-Thomas,
Sainte Croix et Porto Ricco, exécuté par ordre du gouvernement frangais,
depuis le 30 septembre 1796 jusqu’ au 7 juin 1798, sous la direction du capitaine
Baudin, pour faire des recherches et des collections relatives a |’histoire natur-
elle; contenant des observations sur le climat, le sol, la population, l’agricul-
ture, les productions de ces iles, le caractére, les moeurs et le commerce de
leurs habitants.’”’ Ouvrage accompagné de notes et d’additions par M.
Sonnini. Avec une trés belle carte gravée par J. B. Tardieu d’aprés Lopez.
Rais eechVOl OL.
Ledru was the botanist and Anselme Riedlé the gardener of an expedition
sent out by the Paris Museum of Natural History 1796-1798. Unfortunately,
many of the specimens attributed to St. Thomas were in all probability col-
lected on Porto Rico. Many living plants were brought back to the Jardin
des Plantes. The botanical parts of the report are general and not extensive.
L. de Schlechtendal. ‘‘Florula insulae Sti. Thomae Indiae occidentalis.”
Linnaea, 3: 251-276; 4: 78-93; 5: 177-200, 682-688; 6: 722-772. 1828-1831.
About 400 species are enumerated. The plants were collected by C. A.
Ehrenberg, a merchant, in the years 1827 and 1828. The records are anno-
tated and there are some descriptions.
Henry Krebs. ‘(Catalogue of plants found on the island of St. Thomas, W. I.”
1852. [In John P. Knox: A historical account of St. Thomas, W. I., with
its rise and progress in commerce, missions and churches, climate and its
adaptation to invalids, geological structure, natural history and botany.
New York. ]
Over 1,200 plants are enumerated alphabetically, including many in culti-
vation and some algae. Many of the records can not now be substantiated.
Krebs had previously published an account of the geographic distribution of
the Flora of St. Thomas.
J. P. Knox. ‘‘Catalogue des plantes qui naissent spontanément dans I’isle de Saint-
1s
Thomas.” 1857. [Memorie della r. Acad. di Torino, II, 16: Ixxvi-
Ixxxix. ]
This is essentially the same document as the preceding.
A. Eggers. “St. Croix’s Flora.’’ Vidensk. Meddel. Kjgbenhavn. Pp. 33-
afew ditel7loy
Baron Eggers was a Danish official on St. Croix from 1869 to 1874, and
made extensive botanical collections. He records 738 species, with anno-
tations.
H. F. A. Eggers. ‘‘Flora of St. Croix and the Virgin Islands, West Indies.’’ Bull.
H, FE.
US. Nat Mica er pies O70:
Baron Eggers was in command of Danish troops on St. Thomas during
most of the period between 1874 and 1887, and visited St. Jan. In this cat-
alogue he enumerates 881 indigenous or naturalized species, with annotations,
and also records many of the plants in cultivation.
A. Eggers. ‘Supplement til St. Croix’s og Jomfrugernes Flora.” Vidensk.
Meddel. Kjgbenhavn, pp. 11-21. 1889.
This work contains additional records to those previously published by
the author.
BRITTON: FLORA OF THE VIRGIN ISLANDS 21
Otto Kuntze. ‘‘Um die Erde.” Pp. 514. Leipzig. 1881.
Dr. Kuntze visited St. Thomas in 1874, at the beginning of his extensive
travels.
Otto Kuntze. ‘‘Revisio Generum Plantarum” 1: 2: pp. 1009. Leipzig. 1891.
The author records specimens collected by him on St. Thomas in 1874.
F. Borgesen and Ove Paulsen. ‘‘Om Vegetationen paa de Dansk-Vestindiske Mer.”
Botan. Tidsskr. Kjébenhavyn, 22: 1-114, f.I-43. 1898. [Reprint pp. 114.]
Mr. Borgesen visited the islands in 1892, and again in 1895-6, on his second
trip accompanied by Mr. Paulsen. They made extensive collections, and
listed six Spermatophytes as additions to the known flora. The document is
mostly ecologic, and especially detailed as to the composition of the coastal
vegetation. It was translated into French by Mlle. S. Eriksson and pub-
lished in 1900 (Rev. Gen. de Bot. 12: 99-107; 138-153; 224-245; 289-297;
344-354; 434-446; 489-510). [Reprint pp. 108.]
C. F. Millspaugh. ‘‘Plantae Utowanae.”’ Field Col. Mus. Bot. 2: I-110; 113-135.
pl. 25. 1900.
During the cruise of the yacht Utowana, December, 1898, to March, 1899,
Dr. Millspaugh, botanist of the expedition, visited St. Thomas on January
17 and 18, 1899, and collected about 200 species, which are enumerated.
C. F. Millspaugh. ‘Flora of the Island of St. Croix.’’ Field Col. Mus. Bot. 1:
441-546. Map. 1902.
Annotated list of 1,029 species, based especially upon the large collections
made in 1895, 1896 and 1897 by A. E. Ricksecker and Mrs. J. J. Ricksecker,
with records taken from Baron Eggers Flora. Mr. Ricksecker published a
list of the species collected by him, pp. 4, not dated [1896]. Dr. Millspaugh
has a chapter upon the botanical history of St. Croix.
F. Borgesen. ‘Notes on the Shore Vegetation of the Danish West Indian Islands.”
Bot. Tidsskr. 29: 201-259. f.1-140; pl. 3-6. 1909.
Mr. Borgesen made a third trip to the islands during the winter of 1905-
1906, especially for algological studies. The paper is ecological, and supple-
mentary to his earlier publications.
E. G. Britton. ‘‘Mosses of the Danish West Indies and Virgin Islands.”’ Bull.
Torr. Club 42: 1-8. 1915.
Mrs. Britton lists, with annotations, 28 species of Mosses, including 3 de-
scribed as new; four of the plants enumerated were found only on Tortola.
H. G. Brock, P. S. Smith, W. A. Tucker. ‘‘The Danish West Indies, their Resources
and Commercial Importance.’’ I917.
The United States Department of Commerce has recently published as
Special Agents Series 129 (pp. 68, figs. 1-8), a valuable document in which
the vegetable products of commercial value are discussed.
There are a very large number of records of plants from the islands in
taxonomic monographs and lists of species by many authors.
As a literary curiosity record may be made of a manuscript list of the plants
of St. Thomas, undated, arranged upon the Linnaean system of classification,
preserved in the library of the New York Botanical Garden, presented some
years ago by the late Dr. T. F. Allen.
General comments upon the vegetation are to be found in several books
of travel.
The earlier collections of botanical specimens are practically all
to be found only in the herbaria of the Old World. Perhaps the oldest
a2 BROOKLYN BOTANIC GARDEN MEMOIRS
are those of Von Rohr and of Ryan, made about 1780, and preserved
for the most part in the herbarium of the Botanical Museum at Copen-
hagen, where the most complete and extensive collections from these
islands are to be found. .
Prior to 1800 collections were made by L. C. M. Richard, Isert,
West, Pflug, Ledru and Riedlé. During the nineteenth century the
principal collectors were Benzon, Bertero, Ravn, Hornbeck, Ehren-
berg, Breutel, Krebs, Oersted, Holton, Eggers, Krause, Warming,
Borgesen, Paulsen, A. E. Ricksecker, Mrs. J. J. Ricksecker, Otto
Kuntze and Millspaugh. Since 1900 collections have been made
by N. L. Britton, Mrs. Britton, J. F. Cowell, Miss Marble, J. A. Shafer
and J. N. Rose.
A collection made by Kirkman Finley in Trinidad was erroneously
labeled as from St. Thomas, and many errors have been made in citing
these specimens. A few plants collected by Kuntze in Porto Rico
have been erroneously recorded as from St. Thomas, and many col-
lected by Riedlé on Porto Rico have been similarly erroneously re-
corded. Conversely, some plants collected by Purdie on St. Thomas
have been cited as Jamaican.
For the purposes of the following list of plants I have examined
the literature and have studied the following series of specimens:
1. Duplicates of plants collected by Benzon, Hornbeck, Eggers
and Paulsen, received by the New York Botanical Garden in exchange
with the Copenhagen Botanical Museum.
2. The collection made by I. F. Holton on St. Thomas, preserved
in the herbarium of Columbia University.
3. Dr. Otto Kuntze’s St. Thomas plants, which came to the New
York Botanical Garden as a part of his herbarium, presented by Mr.
Andrew Carnegie.
4. The St. Croix collections made by Mr. Ricksecker and a portion
of that made by Mrs. Ricksecker in the herbarium of the New York
Botanical Garden and parts of the complete sets preserved in the
herbarium of The Field Museum of Natural History.
5. Part of the St. Thomas collection made by Dr. Millspaugh.
6. The St. Croix collection made by Mr. Cowell and myself in 1900.
7. The collections made by Dr. Rose, assisted by Mr. Fitch and
Mr. Russell on St. Croix in 1913.
8. The collection made on St. Thomas by Mrs. Britton and Miss
Marble in 1913.
g. The collection made by Dr. Shafer and myself on St. Thomas,
St. Jan and small adjacent islands in 1913.
Mrs. Britton has contributed the catalogue of the mosses, Dr.
Evans that of the hepatics, and Professor Riddle that of the lichens.
BRITTON: FLORA OF THE VIRGIN ISLANDS 23
Our knowledge of the fungi of the islands is but fragmentary and it
is therefore deemed wise not to attempt an enumeration of them at
this time; a mycological survey would doubtless reveal the presence
of several hundred species. Dr. Howe has contributed a note on the
algological collections and researches of Mr. Borgesen.
St. Thomas and St. Jan are two of the Virgin Islands, discovered
by Columbus in 1493, and were so called to commemorate the young
women who are fabled as having accompanied St. Ursula.
The Virgin Island group is usually regarded as composed of the
following islands, proceeding from the west eastward, (1) Culebra, or
Snake Island (Porto Rican); (2) St. Thomas, or San Thomé, and
(3) St. John or San Jan; (4) Tortola, (5) Virgin Gorda, and (6) Ana-
gada (British). Throughout this archipelago there are many islets
and keys, and the marine views from the hills are among the most
charming in America. If to the above mentioned larger islands we
add (7) Jost Van Dyck, the next largest, a British island near Tortola,
we have seven major Virgin Islands, eight if we include Vieques.
Tortola (British) is separated from St. Jan by little over a‘mile
of water. The purchase from the Danish government thus brings
our frontier close to that of the British Empire at another point.
Vieques, or Crab Island (Porto Rican), lies south of the axis of
the archipelago, and is perhaps not properly a member of the Virgin
Island group, although it is sometimes so considered.
These islands were originally inhabited by Arawak and Carib
Indians. St. Thomas was colonized by the Dutch in 1657, passed to
the British about 1667, and to the Danes in 1671, who have since held
it, except for short occupations by the British. St. Jan was colonized
by the Danes in 1684, and their occupancy has since been continuous.
St. Croix, or Santa Cruz, was also discovered by Columbus in 1493 or
early in 1494, colonized by both Dutch and English in 1625, passed
soon to the Spanish, and next to the French in 1651. The Danish
ownership dates from 1733. It is isolated in the sea, and not properly
of the Virgin Island group; in clear weather, it can be seen from the:
hills of Porto Rico and from those of St. Thomas and St. Jan.
All three islands are oblong in shape, with the longer axes nearly
east and west, the coast lines irregular. The hills of St. Thomas
rise to about 1,500 feet; those of St. Jan are somewhat lower (about
1,260 feet), while the highest point on St. Croix (Mt. Eagle) is 1,164
feet. St. Croix is about 21 miles long, 6 miles wide, and has an area
of about 84 square miles, being thus about one seventh larger than
Staten Island, New York (723 square miles). St. Thomas is 13 miles
long, 4 miles wide, with an area, including its islets, of some 32 square
miles; St. Jan is 9 miles long, about 5 miles wide, with an area, in-
24 BROOKLYN BOTANIC GARDEN MEMOIRS
cluding its islets, of about 21 square miles. The total area of the
three islands, including their contiguous islets, is thus about 138
square miles, or not quite twice that of Staten Island. The areas
here used for St. Thomas and St. Jan are approximate, because the
total area of the contiguous islets is not definitely recorded.
The harbor of Charlotte Amalia, coveted by commercial and naval
interests, is the most striking coastal feature of the islands, indenting
the southern coast of St. Thomas. It is something less than a mile
in diameter, a little longer than wide, and is nearly enclosed by the
hills, its mouth being’ approximately goo feet wide. It is as safe an
anchorage as any tropical harbor can be, and affords anchorage for as
many vessels as would be at all likely to need it at any one time, in
water which is up to 37 feet deep. It is not as spacious as Guantanamo
Bay on the southeast coast of Cuba, but as a naval base, with the
hills fortified, would immediately command the Virgin Passage.
Magen’s Bay on the north side of St. Thomas, where a long penin-
sula juts out into the sea, and Coral Bay at the east and Cruz Bay
at the west end of St. Jan, are also valuable harbors, and there are
several other small harbors or coves. The so-called harbors at
Christiansted and Frederiksted, St. Croix, are open roadsteads.
These islands, like Culebra, Tortola, and Virgin Gorda, are partly
plutonic in origin, being partly composed of rocks which have solidified
froma molten state. There is no present evidence of volcanic activity,
as there is in the Leeward and the Windward Islands farther south,
and there are no volcanic peaks. Conglomerate and other stratified
rocks, supposed to be Cretaceous, also occur. They are evidently
ancient, and show evidences of an enormous amount of erosion since
their upheaval; they have not been geologically surveyed.
The soil, except that of some sand beaches and mangrove swamps
and salt marshes, has directly resulted from the decay and erosion
of the rocks; it is of good agricultural quality and locally deep, but
on the steep slopes and hillsides it is meager, having been much
washed away since the cutting away of the forests. There are not
many sand beaches on St. Thomas or St. Jan, but there is a consider-
able area of beach on St. Croix. In sheltered coves and reaches with
shallow water, the mangrove is forming land, as everywhere in similar
situations on tropical coasts.
Along large portions of the coast lines, the rocks come directly
to the sea, forming fine cliffs and headlands, often rising from deep
water, and much of the coastal scenery is highly picturesque.
I have included records of the plants commonly cultivated either
for their products or for ornament and interest, but have made no
attempt to include the rarer or unusual garden plants. If the records
BRITTON: FLORA OF THE VIRGIN ISLANDS 25
by Krebs and Knox are correct, there was a greater variety of plants
in gardens at the middle of the last century than at present.
In citing synonyms for the names of plants, I have given the
original in cases where the species was first named in a genus other
than the one in which it is now included, and I also have indicated
the names used by previous authors dealing with plants of the islands,
in so far as I have been able to refer them, but no attempt has been
made to give complete synonymy.
I gratefully acknowledge aid from Mr. A. S. Hitchcock in the
determination of some grasses and from Miss Margaret Slosson and
Mr. W. R. Maxon for information regarding some ferns.
SPERMATOPHYTA
TY PHACEAE
TYPHA ANGUSTIFOLIA L. [T7. domingensis Pers.; T. angustifolia
domingensis Griseb.] Along rivulets and lagoons, St. Thomas; St.
fan; St. Croix.
PANDANACEAE
PANDANUS UTILIS Bory. [P. odoratissimus of Eggers.] Planted
for ornament.
ZANNICHELLIACEAE
RUPPIA MARITIMA L. [R. rostellata of Eggers.] Shallow, brackish
water, St. Thomas; Buck Island; St. Jan; St. Croix.
CYMODOCEACEAE
CYMODOCEA MANATORUM Aschers. Shallow, salt water, St. Croix;
St. Thomas; St. Jan.
HALODULE Wricuti Aschers. Shallow, salt water, St. Thomas;
St, Croix.
ALISMACEAE
ECHINODORUS CORDIFOLIUS (L.) Griseb. [Alisma cordifolia L.;
A. rostratum Nutt.; Echinodorus rostratus Engelm.] Wet grounds,
sc. thomas; ‘St. Croix.
ELODEACEAE
HALOPHILA BAILLONIS Aschers. In salt water, St. Thomas.
HALOPHILA ASCHERSONII Ostenfeld. In salt water, St. Croix.
HY DROCHARITACEAE
THALASSIA TESTUDINUM Konig. In salt water, St. Thomas; St.
fan St. Croix:
26 BROOKLYN BOTANIC GARDEN MEMOIRS
POACEAE
SACCHARUM OFFICINARUM L. Subspontaneous after cultivation,
St. Croix, where it is extensively cultivated for sugar; grown in small
patches on St. Thomas and St. Jan.
ANDROPOGON GLOMERATUS (Walt.) B.S. P. Doubtfully recorded
from St. Thomas by Hackel.
ANDROPOGON BICORNIS L. [Anatherum bicorne Beauv.|] On the
high hills of St.. Thomas and St. Jan.
ANDROPOGON LEUCOSTACHYUS H.B.K. St. Thomas.
ANDROPOGON JUNCIFOLIUS Desv. St. Croix.
ANDROPOGON CERIFERUS Hack. St. Thomas.
ANDROPOGON PANORMITANUS Parl. [A. saccharoides of Eggers;
A. Wrightit of Millspaugh.] St. Croix.
ANDROPOGON SCHOENANTHUS L. Cultivated for perfume.
HETEROPOGON contToRTUS (L.) Beauv. Krumbay, St. Thomas
(according to Eggers).
Hocus SorcHuM L. [H. saccharatus L.; Sorghum vulgare Pers.;
Andropogon Sorghum Brot.| Subspontaneous after cultivation, St.
Groix: St. Thomas.
ANTHEPHORA HERMAPHRODITA (L.) Kuntze. [Zripsacum herma-
phroditum L.; Anthephora elegans Schreb.; A. villosa Spreng.] Waste
and cultivated grounds, St. Thomas; St. Croix.
NAZIA ALIENA (Spreng.) Scribn. ([Lappago aliena Spreng.; Nazia
racemosa aliena Scribn. & Smith; confused by authors with Nazia
racemosa (L.) Kuntze = Tragus racemosus (L.) Haller.] Sandy fields,
thickets and waste grounds, St. Thomas; St. Jan (according to
Eggers); St. Croix.
VALOTA INSULARIS (L.) Chase. [Andropogon insularis L.; Pant-
cum leucophaeum H.B.K.; P. insulare Meyer; Tricholaena insularis
Griseb.; Syntherisma insularis Millsp.] Dry soil, St. Thomas; St.
Jan; St. Groix:
VALoTA EccersiI (Hack.) Hitche. & Chase. [Panicum Eggersu
Hack.] St. Thomas. Endemic.
SYNTHERISMA DIGITATA (Sw.) Hitche. [Muilium digitatum Sw.;
Digitaria setigera Roth; D. horizontalis Willd.; Syntherisma setigera
Nash; P. sanguinale vulgare of Kuntze in part.] Fields, hills and
cultivated grounds, St. Thomas; St. Croix.
SYNTHERISMA SANGUINALIS (L.) Dulac. [Panicum sanguinale L.;
Digitaria marginata Link.] Fields, hills and cultivated grounds, St.
Thomas; St. Jan; St. Croix.
SYNTHERISMA IsCHAEMUM (Schreb.) Nash. [Panicum Ischaemum
Schreb.] St. Croix (according to Hitchcock & Chase).
BRITTON: FLORA OF THE VIRGIN ISLANDS Li
The grass recorded by Eggers as Digitaria filiformis from Cowell
Hill, St. Thomas, has not been further identified.
ERLOCHLOA PUNCTATA (L.) Desv. [Milium punctatum L.; Helopus
punctatus Nees.] Moist grounds, St. Croix; St. Thomas.
ANASTROPHUS COMPRESSUS (Sw.) Schlecht. [Milium compressum
Sw.; Paspalum platycaulon Poir; P.compressum Rasp.| Wet grounds,
mie bnomas; St. Jan; St. Croix.
PASPALUM GLABRUM Poir. [P. Hellert Nash; Panicum plantagineum
of Millspaugh; ? P. Richardt Steud.| Wet grounds, St. Thomas; St.
wean; St. Croix:
PASPALUM PLICATULUM Michx. [P. undulatum Poir.; P. caespi-
tosum of Eggers, at least in part.] Hillside, Buck Island, St. Thomas;
St. Croix (according to Eggers).
PASPALUM PANICULATUM L. [P. hemisphaericum Poir.] St.
Thomas (according to Schlechtendal).
PASPALUM FIMBRIATUM H.B.K. Waste grounds and roadsides,
St. Croix.
PASPALUM ORBICULATUM Poir. [P. pusillum Vent.] St. Thomas
(Fluegge; according to Grisebach).
PASPALUM CONJUGATUM Berg. Grassy places, St. Thomas; St.
Jan; St.. Croix:
PASPALUM NOTATUM Fluegge. St. Thomas is the type locality of
the species, but the plant has not been found there by recent collectors.
PASPALUM VIRGATUM L. St. Jan; St. Croix (according to West).
PASPALUM SECANS Hitche. & Chase. Sandy soil, St. Croix.
PASPALUM DISTICHUM L. Wet grounds, St. Thomas; St. Croix.
PASPALUM VAGINATUM Sw. [P. distichum vaginatum Sw.| Wet
grounds, St. Croix.
PASPALUM SPATHACEUM Desv., recorded as from St. Thomas by
Schlechtendal, is a species not understood by modern botanists.
PASPALUM MOLLE Poir., described as from St. Thomas: is a species
not understood by modern botanists.
PANICUM GEMINATUM Forsk. [P. paspaloides of Eggers and of
Millspaugh; P. brizoides Lam., not L.; Paspalum appressum Lam.]
Wet grounds, St. Thomas; St. Jan; St. ene.
PANICUM BARBINODE Trin. [P. molle of Eggers. ] Moist grounds,
St. Croix.
PANICUM REPTANS L. [P. grossarium L.; P. prostratum Lam.;
P. prostratum pilosum Eggers; P. caespitosum Sw.| Hillside thickets,
et. Ehomas; St. Jan; St. Croix.
PANICUM FASCICULATUM Sw. [P. fuscum Sw.; P. fasciculatum
fuscum Griseb.; P. fasciculatum flavescens of Kuntze.] Banks, hill-
sides and thickets, St. Thomas; St. Jan; St. Croix.
28 BROOKLYN BOTANIC GARDEN MEMOIRS
PANICUM MILIACEUM L. Waste grounds, St. Croix.
PANICUM ADSPERSUM Trin. Hillside thicket, Bethania, St. Jan.
PANICUM CAYENNENSE Lam. St. Thomas (recorded with doubt
by Schlechtendal).
PANICUM DIFFUSUM Sw. Rocky hillsides, St. Thomas; recorded
from all three islands by Eggers and from St. Croix by Grisebach.
PANICUM MAXIMUM Jacq. [P. jumentorum Pers.; P. polygamum
Sw:| “Dry soil, :St- Thomas: St. Jan: St Crom:
PANICUM LAXUM Sw. Hillsides, St. Thomas.
PANICUM TRICHOIDES Sw. Barracks, St. Thomas (recorded by
Eggers as P. brevifolium L.).
PANICUM GLUTINOSUM Sw. St. Croix (according to West).
LASIACIS DIVARICATA (L.) Hitche. [Panicum divaricatum L.;
P. divaricatum glabrum Kuntze.| Thickets and hillsides, St. Thomas;
Ste lat ote Crom:
Lastacis HArristI Nash. St. Jan.
LASIACIS SORGHOIDEA (Desvaux) Hitchc. & Chase. [Panicum
sorghoideum Desvaux; P. latifolium of Millspaugh.] Thickets, St.
Thomas St..Croix.
LASIACIS LIGULATA Hitchc. & Chase. [Panicum diaricatum
puberulum Griseb.| Shaded bank, St. Peter, St. Thomas.
ECHINOCHLOA COLONUM (L.) Link. [Panicum colonum L.] Grassy
places, waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
OPLISMENUS HIRTELLUS (L.) Beauv. [Panicum hirtellum L.; P.
setarium Lam.; O. setarius R. & S.; Orthopogon setarius Spreng.]
Woodlands, St. Thomas; St. Jan; St. Croix.
CHAETOCHLOA GENICULATA (Lam.) Millsp. & Chase. [Panicum
geniculatum Lam.; P. imberbe Poir.; Setaria glauca imberbis Griseb.;
S. glauca of Eggers; Chaetochloa glauca of Millspaugh.] Woodlands,
waste and cultivated grounds, St. Thomas; St. Croix; St. Jan.
CHAETOCHLOA SETOSA (Sw.) Scribn. [Panicum setosum Sw.;
Setaria setosa Beauv.; Panicum caudatum Lam.; Setaria setosa caudata
Griseb.; Setaria macrostachya of Schlechtendal.] Hillsides, St. Thom-
as; St. Groix: ‘St; Jan: -
CHAETOCHLOA RARIFLORA (Mikan) Hitchc. & Chase. [Setaria rart-
flora Mikan.] Hillsides, St. Thomas; St. Croix.
CENCHRUS ECHINATUS L. [C. viridis of Millspaugh; C. echinatus
brevisetus Scribn.; C. echinatus tribuloides of Kuntze.] Fields and
hillsides, St. Thomas; St. Jan; St. Croix.
CENCHRUS CAROLINIANUS Walt. St. Thomas (according to Hitch-
cock & Chase).
CENCHRUS VIRIDIS Spreng. Dry soil, St. Thomas.
STENOTAPHRUM SECUNDATUM (Walt.) Kuntze. [Ischaemum secun-
BRITTON: FLORA OF THE VIRGIN ISLANDS 29
datum Walt.; S. glabrum Trin.; S. americanum Schrank.] Moist
grounds, St. Thomas; St. Jan; St. Croix.
OLYRA LATIFOLIA L. Woodlands, St. Thomas; Cinnamon Bay,
St. Jan (according to Eggers.)
PHARUS GLABER H.B.K. Woodlands, St. Thomas; St. Jan; St.
Croix.
ORYZA SATIVA L. St. Thomas (according to Pilger).
ARISTIDA ADSCENSIONIS L. [Aristida bromoides H.B.K.; A. stricta
Griseb., not Michx.; A. americana Pilger, not L.] Thickets and hill-
miessot. Lhomas; St. Jan; St. Croix.
ARISTIDA COGNATA Trin. & Rupr. [A. Swartziana Steud.] Hill-
sides, St. Thomas; St. Croix.
SPOROBOLUS viRGINICUS (L.) Kunth. [Agrostis virginica L.]
Saline soil, St. Thomas; St. Jan; St. Croix.
SPOROBOLUS BERTEROANUS (Trin.) Hitche. & Chase. [Vilfa
Berteroana Trin.; Sporobolus angustus Buckley.| Wet grounds, St.
ane -ot. Croix.
SPOROBOLUS ARGUTUS (Nees) Kunth. [S. domingensis of Miil-
spaugh; ? S. littoralis of Eggers.] Saline soil, St. Croix.
SPOROBOLUS INDICUS (L.) R. Br. [Agrostis indica L.; Vilfa
tenacissima Kunth.] Dry soil, St. Thomas; St. Jan; St. Croix.
SPOROBOLUS MURALIS (Raddi) Hitche. & Chase. [A grosticula
muralis Raddi; S. minutiflorus of Millspaugh.]| Waste grounds and
roadsides, St. Croix.
CApRIOLA DactyLon (L.) Kuntze. [Panicum Dactylon L.; Cyno-
don Dactylon Pers.| Dry soil, St. Thomas; St. Jan; St. Croix.
CHLORIS RADIATA (L.) Sw. [Agrostis radiata L.] Dry soil, St.
Thomas; St. Jan (according to Eggers); St. Croix.
CHLORIS PARAGUAIENSIS Steud. [C. barbata Sw.; C. ciliata of
Eggers.] Waste and cultivated grounds, St. Thomas; St. Jan; St.
Croix.
CHLORIS SAGRAEANA A. Rich. [C. eleusinoides Griseb.] St. Croix.
CHLORIS CILIATA Sw. Dry soil, St. Thomas; St. Croix.
BOUTELOUA AMERICANA (L.) Scribn. [Aristida americana L.;
Heterostega juncifolia Desv.; B. litigiosa Lag.| Hillsides and banks,
Seethomas; St. Jan; St. Croix.
GYMNOPOGON FOoLIOsUS (Willd.) Nees. [Chloris foliosa Willd.]
St. Thomas.
ELEUSINE INDICA (L.) Gaertn. [Cynosurus indicus L.| Waste
and cultivated grounds, St. Thomas; St. Jan; St. Croix.
_ DACTYLOCTENIUM AEGYPTIUM (L.) Richt. [Cynosurus aegyptius
L.] Waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
LEPTOCHLOA FILIFORMIS (Lam.) Beauv. [Festuca filiformis Lam. ;
30 BROOKLYN BOTANIC GARDEN MEMOIRS
L. mucronata (Michx.) Kunth; ZL. mucronata multiflora Eggers.] St.
Croix, along ditches (according to Eggers).
LEPTOCHLOA VIRGATA (L.) Beauv. [Cynosurus virgatus L.; ? L.
virgata gracilis Eggers; Chloris poaeformis H.B.K.] Moist or wet
grounds, St. Thomas; St. Jan; St. Croix.
DIPLACHNE FASCICULARIS (Lam.) Beauv. [Festuca fascicularis
Lam.; Leptochloa fascicularis A. Gray.] Ina ditch, St. John’s Estate,
Stroix:
PAPPOPHORUM ALOPECUROIDEUM Vahl. [P. laguroideum Schrad.|
Rocky hillsides, St. Thomas; Buck Island, St. Thomas (according to
Eggers).
ERAGROSTIS PILOSA (L.) Beauv. [Poa pilosa L.; E. poaoides of
Grisebach.] Dry soil, St. Thomas; St. Croix.
ERAGROSTIS TEPHROSANTHUS Schultes. Dry soil, St. Thomas; St.
Croix.
ERAGROSTIS CILIARIS (L.) Link. [Poa ciliaris L.; E. ciliaris laxa
Kuntze.] Dry soil, St. Thomas; St. Jan; St. Croix.
ERAGROSTIS AMABILIS (L.) Wight & Arn. [Eragrostis plumosa
Link.] Cultivated grounds, St. Jan.
ERAGROSTIS BARRELIERI Dav. [E£. minor of Millspaugh; £.
poaoides of Eggers.] Dry soil, St. Thomas (according to Eggers);
St: Croiss
ERAGROSTIS ELLIoTTIIS. Wats. Dry soil, St. Thomas.
UNIOLA VIRGATA (Poir.) Griseb. [Poa virgata Poir; U. racemt-
flora Trin.] Bolongo, St. Thomas; Little St. James Island, St. Jan.
ARTHROSTYLIDIUM CAPILLIFOLIUM Griseb. Flag Hill, St. Thomas;
Battery, St. Jan.
BAMBOS VULGARIS Schrad. Naturalized in wet grounds, St.
Thomas; St. Croix.
Corx LacryMA-Jospt L. Cultivated for ornament.
ZEA MAYS L. Cultivated for food.
CYPERACEAE
KYLLINGA BREVIFOLIA Rottb. [K. monocephala Thunb. of
Schlechtendal and of Eggers.] Moist, grassy places, St. Thomas;
St. Jan; St. JCrom:
KYLLINGA ODORATA Vahl. [K. triceps of Eggers; K. odorata minor
Boeckl.]| Moist, shaded banks, St. Thomas; St. Jan.
KYLLINGA PUMILA Michx. Moist grassy places, St. Thomas,
collected by Riedlé (according to Clarke).
KYLLINGA PUNGENS Link. Midland, St. Croix.
CyYPERUS ODORATUS L. _[C. polystachyus R. Br.; Pycraeus odoratus
Urban.] Crown, St. Thomas, at about 500 m. altitude (according to
BRITTON: FLORA OF THE VIRGIN ISLANDS 31
Eggers). Not found by us on St. Thomas, but collected on Tortola
at about the same elevation.
CYPERUS LAEVIGATUS L. [C. laevigatus albidus Eggers; C. mucro-
natus Rottb.; Juncellus laevigatus Clarke.| Wet grounds, St. Thomas;
St. Croix.
CYPERUS SURINAMENSIS Rottb. Wet or moist grounds, St.
Thomas.
CYPERUS OCHRACEUS Vahl. Moist grounds, St. Croix.
CYPERUS ELEGANS L. [C. viscosus Sw.] Wet saline grounds, St.
Mhomas; St. Jan; St. Croix.
CYPERUS SPHACELATUS Rottb. Pastures and hillsides, Signal Hill
and Crown, St. Thomas.
CypERUS ComPREsSUS L. Moist ground, Haven Sight, St. Thomas
(according to Eggers). Not found by us on St. Thomas, but collected
on Virgin Gorda, Vieques and Culebra.
CYPERUS DISTANS L. f. [Cyperus Eggersit of Millspaugh.] Pas-
tures and ditches, Signal Hill and St. Peter, St. Thomas; Mt. Eagle,
a A TOlx.
CYPERUS ESCULENTUS L. [C. esculentus macrostachyus Boeckl.]
St. Thomas (according to Clarke).
CYPERUS ARTICULATUS L. Wet grounds, St. Thomas; St. Croix.
CYPERUS ROoTUNDUS L. [C. Hydra Michx.] Waste and cultivated
grounds, St. Thomas; St. Jan; St. Croix.
CYPERUS CAYENNENSIS (Lam.) Britton. [Ayllinga cayennensts
Lam.; Mariscus flavus Vahl; Cyperus flavus Nees; Cyperus flavo-
mariscus Griseb.; Mariscus cayennensis Urban.] Grassy places, St.
Thomas; St. Croix.
CYPERUS GRANULARIS (Desf.) Britton. [Mariscus gracilis Vahl;
Kyllinga filiformis capillaris Griseb.; C. capillaris of Millspaugh.]
Sandy soil, near the coast, St. Croix.
CYPERUS TENUIS Sw. St. Croix (according to Clarke).
CypPERUS LIGULARIS L. [Mariscus rufus H.B.K.; M. lgularis
Urban.] Moist, especially saline soil, St. Thomas; St. Jan; St.
Croix.
CYPERUS CONFERTUS Sw. [Mariscus confertus Sw.] Hillsides and
thickets, St. Thomas; St. Croix (according to Grisebach).
CYPERUS PURPURASCENS Vahl. Coastal rocks, Water Island, St.
Thomas; St. Croix.
CYPERUS BRUNNEUS Sw. [C. brizaeus Vahl; C. Ottonis Boeckl.;
C. discolor Boeckl.; Mariscus brunneus Clarke.] Coastal sands, St.
Thomas; St. Jan; St. Croix.
CYPERUS FERAX L. C. Rich. [C. pennatus of Eggers; C. flexuosus
Vahl; C. odoratus of Eggers; Torulinium ferax Urban; C. Michauxt-
anus of Millspaugh.] Wet grounds, St. Thomas; St. Croix.
a2 BROOKLYN BOTANIC GARDEN MEMOIRS
Cyperus VAHLII (Nees) Steud. Moist soil on hills, St. Thomas;
St. Jan.
CYPERUS FILIFORMIS Sw. [TZorulinium filiforme Clarke; C. uni-
folius Boeckl.] Moist soil, St. Thomas; St. Croix.
CYPERUS FERRUGINEUS Poir. [Pycraeus ferrugineus Clarke] re-
corded from St. Thomas by Clarke on the evidence of a specimen in
the herbarium of the British Museum, is probably an error in locality.
CYPERUS sTRIGOSUS L. Recorded by Schlechtendal as found in a
garded on St. Thomas, is probably an error in name.
ELEOCHARIS INTERSTINCTA (Vahl) R. & S. [Scirpus interstinctus
Vahl.] Marshes, St. Thomas; St. Croix.
ELEOCHARIS MUTATA (Vahl) R. & S. [Scirpus mutatus Vahl; E.
cellulosa of Millspaugh.] Wet grounds, St. Croix; St. Jan (according
to Eggers).
ELEOCHARIS FLACCIDA (Spreng.) Urban. [Scirpus flaccidus Spreng.
E. ochreata Nees.| Wet grounds, St. Thomas.
ELEOCHARIS CAPITATA (L.) R. Br. [Scirpus capitatus L.] Wet
grounds, St. Thomas; St. Jan; St. Croix.
ELEOCHARIS RETROFLEXA (Poir.) Urban. [Scirpus retroflexus
Poir.; Eleocharis Chaetaria R. & S.] Moist grounds, St. Thomas.
ELEOCHARIS MINIMA Kunth. Krumbay, St. Thomas (according
to Clarke).
ELEOCHARIS NODULOSA (Roth) Schultes. [Scirpus mnodulosus
Roth.] Adventure, St. Croix (according to Eggers).
SCIRPUS SUBDISTCHUS Boeckl., described as from St. Thomas, has
not been identified by subsequent botanists.
SCIRPUS ARTICULATUS (Kunth) Griseb. is recorded as from St.
Croix by Kunth, presumably erroneously, it being an Old World
species.
FIMBRISTYLIS DIPHYLLA (Retz.) Vahl. [Scirpus diphyllus Retz.;
?.S. dichotomus of Schlechtendal; Scirpus brizoides Muhl.; Fim-
bristylis polymorpha Boeckl.] Grassy places, St. Thomas; St. Jan;
Dt row.
FIMBRISTYLIS FERRUGINEA (L.) Vahl. [Scirpus ferrugineus L.]
Moist, saline soil, St. Thomas; St. Jan; St. Croix.
FIMBRISTYLIS SPADICEA (L.) Vahl. [Scirpus spadiceus L.] Moist
soil near the coast, St. Thomas; St. Croix.
ABILDGAARDIA MONOSTACHYA (L.) Vahl. [Cyperus monostachyus
L.; Fimbristylis monostachya Hassk.]| Moist, shaded bank, Rosen-
berg, St. Jan.
DICHROMENA CILIATA Vahl. [Rynchospora pura Griseb.] Pas-
tures and hillsides, Signal Hill and Crown, St. Thomas; Bordeaux,
Se pan.
BRITTON: FLORA OF THE VIRGIN ISLANDS mS
DICHROMENA RADICANS Schl. & Cham. Shaded banks, St. Thomas.
RyncHosporA BERTERI (Spreng.) Clarke. [Hypolytrum Berteriw
Spreng.; Rynchospora pusilla (Sw.) Griseb., not R. pusilla Chapm.]
Pastures, Signal Hill, St. Thomas (according to Eggers).
RYNCHOSPORA PODOSPERMA C. Wright. St. Thomas; a specimen
in the Arnott Herbarium (according to Clarke).
SCLERIA DISTANS Poir. St. Thomas (according to Clarke).
SCLERIA LITHOSPERMA (L.) Sw. [Scirpus lithospermus L.; Scleria
filiformis Sw.] Rocky thickets, St. Thomas; St. Croix.
SCLERIA PTEROTA Presl. [Scleria pratensis. Nees; S. communis
of Millspaugh.] Moist woodlands, St. Thomas; St. Jan; St. Croix.
SCLERIA SCINDENS Nees. Forests, Signal Hill, St. Thomas.
ARECACEAE
CocCOTHRINAX ARGENTEA (Lodd.) Sarg. [C. sancti-thomae Bec-
cari; C. Eggersiana Beccari; C. Eggersiana sanctae-crucis Beccari;
Thrinax argentea Lodd.; ? T. parviflora of Eggers.] Hillsides, Water
Island and Flag Hill, St. Thomas; St. Jan; St. Croix.
ACROCOMIA ACULEATA (Jacq.) Lodd. [Cocos aculeata Jacq.]
Hillside, St. Peter, St. Thomas.
ROysTONEA REGIA (H.B.K.) O. F. Cook. [? Areca oleracea of
West; Oreodoxa regia H.B.K.; ?0O. oleracea of Kuntze.] Wooded
ravine, Tutu, St. Thomas; St. Croix. Planted for ornament.
Cocos NUCIFERA L. Spontaneous after planting, especially in
€aastal sands, St. Thomas; St. Jan; St. Croix.
SABAL Planted, Charlotte Amalia, St. Thomas.
BORASSUS FLABELLIFER L. Recorded by West as found on St.
Croix. An East Indian palm.
ARACEAE
ANTHURIUM ACAULE (Jacq.) Schott. [Pothos acaulis Jacq.;
Anthurium Huegelit of Eggers.]| On rocks and trees in shaded situa-
tions, St. Thomas; St. Jan; St. Croix (according to Eggers).
ANTHURIUM GRANDIFOLIUM (Jacq.) Kunth. [Pothos grandtfolia
Jacq.; A. macrophyllum of Eggers.| On rocks in woodlands, St.
Thomas; St. Jan.
ANTHURIUM CORDATUM (Willd.) D. Don. [Pothos cordata Willd.;
? P. macrophyllum of West.] On rocks in forests, St. Jan; St. Croix.
ANTHURIUM SELLOUM C. Koch. On trees and rocks in forests, St.
van.
‘PHILODENDRON Krepssit Schott. [P. hederaceum of Eggers.] On
trees in forests, Crown, St. Thomas.
4
34 BROOKLYN BOTANIC GARDEN MEMOIRS
PHILODENDRON OXYCARDIUM Schott. On trees in forests, St.
Thomas.
PHILODENDRON GIGANTEUM Schott. On rocks in dense forests,
Signal Hill and Crown, St. Thomas (according to Eggers).
DIEFFENBACHIA SEGUINE (Jacq.) Schott. [Arum Seguine Jacq.]
Caret Bay, St. Thomas (according to Eggers).
CALADIUM BICOLOR (Ait.) Vent. [Arum bicolor Ait.; ? C. smarag-
dinum of Eggers.] St. Thomas (according to Urban). Cultivated on
St Groin.
XANTHOSOMA ATROVIRENS C. Koch. Cultivated and naturalized,
St. Thomas; St. Croix (according to Eggers).
XANTHOSOMA SAGITTIFOLIUM (L.) Schott. [Arum sagittifolium L.;
Arum maculatum of Millspaugh.] Naturalized after cultivation, St.
Thomas; St. Croix. Cultivated for its roots.
XANTHOSOMA ? HASTATUM Eggers, recorded by Eggers as spon-
taneous after cultivation on all three islands, has not been identified.
Arum hastatum Vahl, cited by Eggers as a synonym, is, an unpublished
name, printed in West’s Flora of St. Croix.
PISTIA STRATIOTES L. [P. occidentalis Blume.] Naturalized in
gardens, St. Thomas (according to Eggers).
LEMNACEAE
LEMNA PERPUSILLA Torr. [L. minor Eggers; L. paucicostata
Hegelm.] In still fresh water, St. Croix; St. Jan (according to
Eggers).
BROMELIACEAE
BROMELIA PINGUIN L. Hillsides.and thickets; used for hedges,
St. Thomas; StxCroix: St) Jan:
WITTMACKIA LINGULATA (L.) Mez. [Bromelia lingulata L.;
Chevalliera lingulata Griseb.] On trees and rocks on hills, St. Thomas;
St. Jan.
PITCAIRNIA LATIFOLIA Sol. St. Croix (according to Mez).
PITCAIRNIA ANGUSTIFOLIA (Sw.) Redouté. [Hepetis angustifolia
Sw.| ‘On rocks; St; Thomas; St Jan s-St. Crom
CATOPSIS NUTANS (Sw.) Griseb. On trees in forests, high hills of
St. Thomas and St. Jan.
TILLANDSIA UTRICULATA L. On trees and. rocks, St. Thomas;
St. Crom
TILLANDSIA FASCICULATA L. On trees in woodlands, St. Thomas;
Si ian,
TILLANDSIA RECURVATA L. On trees, St. Thomas; St. Jan; St.
Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS ® fo)
Eggers records, in his supplementary list, another, undetermined
Tillandsia from Adrian, St. Jan.
DENDROPOGON USNEOIDES (L.) Raf. [Tuillandsia usneoides L.]
On trees and shrubs, St. Thomas; St. Jan; St. Croix.
ANANAS ANANAS (L.) Cook & Collins. [Bromelia Ananas L.;
Ananas sativus Lindl.] Cultivated for its fruit.
COMMELINACEAE
COMMELINA LONGICAULIS Jacq. [C. cayennensis L. C. Rich.; C.
communis of West; C. nudiflora Clarke, not L.] Moist shaded situa-
Hons, ot. Thomas; St. Jan; St. Croix.
COMMELINA ELEGANS H.B.K. [C. virginica of Millspaugh and of
Kuntze.] Moist grounds, St. Thomas; St. Croix.
CALLISIA REPENS L. Shaded situations, St. Thomas; St. Jan;
et. Croix.
CALLISIA MONANDRA (Sw.) Schult. [Tvadescantia monandra Sw.;
Callisia umbellulata Lam.] Among shaded rocks, Signal Hill, St.
Thomas (according to Eggers).
RHOEO DISCOLOR (L’Her.) Hance. [Zvadescantia discolor L’Her.]|
Waste rocky places, St. Thomas; St. Jan; St. Croix.
ZEBRINA PENDULA Schnitzl. Lawns and cultivated grounds, St.
Thomas; St. Croix. Naturalized.
PONTEDERIACEAE
PIAROPUS CRASSIPES (Mart.) Raf. [Pontederia crassipes Mart.;
Eichhornia crassipes Solms; E. azurea of Millspaugh.] In water, St.
Croix.
LILIACEAE
ALOE VERA L. [Aloe vulgaris Lam.; A. perfoliata of West.] On
limestone and in fields, St. Thomas; St. Jan; St. Croix. Naturalized.
Cordyline guineensis (Jacq.) Britton. [Aletris guineensis Jacq.;
Sanseviera guineensis Willd.| Hillsides, St. Thomas; St. Croix.
Naturalized.
Yucca ALorroyia L. [Y. Draconis L.] Planted for ornament.
Yucca GLoRIOSA L., is recorded by Eggers as naturalized in gardens
and near dwellings on St. Thomas and St. Croix. Planted for orna-
ment.
ALLIUM PORRUM L. Cultivated for food.
ALLIuM CEPA L. Cultivated for food.
ALLIUM FISTULOsSUM L. Cultivated for food.
ALLIUM SATIVUM L. Cultivated for food.
36 BROOKLYN BOTANIC GARDEN MEMOIRS
CONVALLARIACEAE
ASPARAGUS OFFICINALIS L. Planted for food.
SMILACEAE
SMILAX ILICIFOLIA Kunth. [S. havenensis of Eggers.] Hillside
thickets, St. Jan-(?)):) St. Gro:
SMILAX CORIACEA Spreng. [S. subarmata O. E. Schulz; S. popul-
nea of Eggers.] Hillside thickets, St. Thomas; St. Croix.
SMILAX ROTUNDIFOLIA L., cited by O. E. Schulz as from St. Croix,
is an error in record or determination.
SMILAX DOMINGENSIS Willd., cited by A. de Candolle from St.
Thomas, is an error in locality.
AMARYLLIDACEAE
ATAMASCO TUBISPATHA (L’Her.) Maza. [Amaryllis tubispatha
L’Her.; Zephyranthes tubtspatha Herb.] In fields and near dwellings,
St. Thomas; St. Croix; St. Jan (according to Eggers).
ATAMASCO ROSEA (Lindl.) Greene. [Zephyranthes rosea Lindl.;
? Amaryllis Atamasco of West.] Cultivated for ornament.
CRINUM ERUBESCENS Ait. Along rivulets, St. Croix (according
to Eggers); cultivated for ornament.
CRINUM GIGANTEUM Andr. Cultivated for ornament.
CRINUM LONGIFOLIUM Herb. Cultivated for ornament, St. Croix,
and seemingly an escape (according to Millspaugh).
HYMENOCALLIS DECLINATA (Jacq.) Roem. [Hymenocallis expansa
Herb.; Pancratium caribaeum of Eggers; P. declinatum Jacq.; ? P.
patens of Schlechtendal; H. caribaea of Millspaugh.] Rocky coasts
and hillsides, St. Thomas; St. Croix; St. Jan.
HyMENOCALLIS CARIBAEA (L..) Herb. [Pancratium caribaeum L.]
Planted for ornament.
HIPPEASTRUM PUNICEUM (Lam.) Urban. [Amaryllis puniceus
Lam.; A. equestris Ait.; Hippeastrum equestre Herb.] Rocky shores
and hillsides, St. Thomas; St. Croix; St. Jan.
AGAVE SISALANA Perrine. Persistent after cultivation, St. Croix.
Cultivated for fiber.
AGAVE MISSIONUM Trelease. [Agave americana of Eggers in part;
A. sobolifera and A. Morrisii of Eggers.] Hillsides, St. Thomas; St.
Jan. Known otherwise on the other Virgin Islands and on Porto Rico.
AGAVE EGGERSIANA Trelease. [A. americana of West and of
Eggers, in part, and of Millspaugh.] St. Croix. Endemic, but not
definitely known in the wild state. Planted for ornament.
FURCRAEA TUBEROSA Ajit. f. [F. cubensis of Eggers and of Muills-
BRITTON: FLORA OF THE VIRGIN ISLANDS a7
paugh; F. hexapetala of Urban, in part.] Thickets, St. Thomas;
St. Croix.
HypoxIS DECUMBENS L. Grassy banks, St. Jan.
AMARYLLIS BELLADONNA L. Planted for ornament.
POLIANTHES TUBEROSA L. Planted for ornament.
DIOSCOREACEAE
DIOSCOREA PILOSIUSCULA Bert. Forests, high hills of St. Thomas;
eit. jan.
DioscoREA ALATA L. Persistent after cultivation, St. Thomas;
et Croix; St. Jan.
DIOSCOREA SATIVA L. [D. altissima of Eggers, at least in part.]
Persistent after cultivation, all islands (according to Eggers). Culti-
vated for its roots.
RAJANIA CORDATA L. [R. pletroneura Griseb.; R. hastata of
Eggers.] Forests, hills of St. Thomas.
IRIDACEAE
Galatea bulbosa (Mill.) Britton. [Sisyrinchium bulbosum Mill.;
S. palmifolium Cav.; Cipura plicata Griseb.; Eleutherine plicata
Herb.] Valleys, St. Croix. Grown in flower gardens.
MUSACEAE
’ Musa PARADISIACA L. Cultivated for its fruit.
MUSA SAPIENTUM L. Cultivated for its fruit.
ZINGIBERACEAE
ALPINIA OCCIDENTALIS Sw. [Amomum sylvestre of West; Reneal-
mia occidentalis Sweet; R. sylvestris of Eggers.] Forests and shaded
situations, Golden Rock, St. Croix; Signal Hill, St. Thomas.
ZINGIBER ZINGIBER (L.) Karst. [Amomum Zingiber L.; Zingiber
officinale Rosc.] Spontaneous after cultivation. St. Thomas; St.
Croix.
CURCUMA LONGA L. Cultivated for tumeric.
LANGUAS SPECIOSA (Wendl.) Small. [Zeruwmbet speciosum Wendl.;
Alpinia nutans Rosc.| Planted for ornament.
CANNACEAE
CANNA INDICA L. Moist waste places, St. Thomas; St. Croix (ac-
cording to Eggers). The plant may have been mistaken for C.
coccinea Ait.
38 BROOKLYN BOTANIC GARDEN, MEMOIRS
CaNnNA LAMBERTI Lindl. Naturalized in gardens, all islands (ac-
cording to Eggers); escaped in places, St. Croix (according to Mills-
paugh).
CANNA EDULIS Ker. Cultivated for its tubers.
CANNA LUTEA Mill. Cultivated and escaped in gardens at Bassin,
St. Croix (according to Millspaugh).
MARANTACEAE
MARANTA ARUNDINACEA L. [Maranta indica Tuss. of Miuills-
paugh.] Escaped or spontaneous after cultivation, St. Thomas;
Sit. Croix.
ORCHIDACEAE
HABENARIA MONORRHIZA (Sw.) Rchb. f. [Orchis monorrhiza Sw.;
Habenaria maculosa of Eggers.|] Hillsides, St. Thomas; St. Croix
(according to Cogniaux.) .
HABENARIA ALATA Hook. Signal Hill, St. Thomas.
VANILLA EcceErsit Rolfe. [V. aphylla Eggers, not Blume.]
Thickets, St. Thomas.
Beadlea elata (Sw.) Small. [Satyriwm elatum Sw.; Spiranthes
elata L. C. Rich.]’ In leaf mould and on wet shaded banks on high
hillssSt. Thomas, St. jane St. ‘Crom.
IBIDIUM TORTILE (Sw.) House. [Satyriwm tortile Sw.; Spiranthes
tortilis L. C. Rich.] Grassy hillsides, St. Thomas.
STENORRHYNCHUS LANCEOLATUS (Aubl.) Griseb. In clayey soil
among rocks, Signal Hill, St. Thomas (according to Eggers.)
CRANICHIS MUSCOSA Sw. Woods between Crown and Signal Hill,
St. Thomas. ;
PRESCOTTIA OLIGANTHA (Sw.) Lindl. [Cvranichis oligantha Sw.;
Prescottia myosurus Rchb. f.] Grassy fields and banks, hills of St.
Thomas; shaded bank, Bordeaux, St. Jan.
PRESCOTTIA STACHYODES (Sw.) Lindl. [Cranichis stychyodes Sw.]
Wooded hills, Bordeaux, St. Jan.
PONTHIEVA GLANDULOSA (Sims) R. Br. [Neottia glandulosa Sims.]
Wet shaded banks, St. Thomas; St. Jan.
LipaRIs ELATA Lindl. Among rocks on high hills, St. Thomas.
LipAris EGGerstt Rchb. f. Bonne Résolution, St. Thomas.
Perhaps not distinct from the preceding species.
POLYSTACHYA MINUTA (Aubl.) Britton. [Epidendrum minutum
Aubl.; Polystachya luteola Hook.; Cranichis luteola Sw.] On rocks,
walls and trees, Signal Hill and St. Peter, St. Thomas.
EPIDENDRUM PAPILIONACEUM Vahl. [E. bifidum Sw.; E. papih-
onaceum grandiflorum Cogn.| On small trees and shrubs, St. Thomas;
St. Jan; St.cCroix,
BRITTON: FLORA OF THE VIRGIN ISLANDS 39
EPpIDENDRUM CILIARE L. On shaded rocks and trees, St. Thomas;
Beans St. Croix.
EPIDENDRUM COCHLEATUM L. On trees, Mt. Eagle and Jacob’s
Peak, St. Croix (according to Eggers).
EPIDENDRUM PATENS Sw. On rocks, Signal Hill, St. Thomas (ac-
co ding to Eggers).
EPIDENDRUM CARINATUM Vahl, of St. Croix, is a species unknown
to modern botanists.
TETRAMICRA ELEGANS (Hamilt.) Cogn. [Cyrtopodium elegans’
Hamilt.; Epidendrum subaequale Eggers.]| Rocky hillsides, St.
Mhomas; St. Jan; St. Croix.
BRASSAVOLA CUCULLATA (L.) R. Br. [Epidendrum cucullatum L.]
On rocks, St. Thomas.
IONOPSIS UTRICULARIOIDES (Sw.) Lindl. [Epidendrum utriculari-
oides Sw.] St. Thomas (according to Cogniaux).
Onciprum LErBo.Lp!I Rchb.f. Flag Hill, St. Thomas (according to
Cogniaux).
ONCIDIUM VARIEGATUM Sw. On shrubs and trees, rarely on rocks,
St. Thomas; St. Croix.
ONCIDIUM INTERMEDIUM Bertero. [O. Lemonianum Lindl.] For-
ests and thickets, rare, Picaria Peninsula and Fortuna, St. Thomas
(according to Eggers).
CASUARINACEAE
CASUARINA EQUISETIFOLIA L. Planted; occasionally spontaneous
on St. Thomas.
PIPERACEAE
PirpER AMALAGO L. [P. medium Jacq.; P. Sieberi C. DC.] Wood-
lands and forests, St. Thomas; St. Jan; St. Croix.
PIPER DILATATUM L. C. Rich. [Piper Bredermyeri of Eggers and
of Millspaugh.] Shaded valleys, St. Croix.
PIPER BLATTARUM Spreng. Forests, Crown and Signal Hill, St.
Thomas (according to Eggers); known otherwise only from Porto
Rico.
PIPER RETICULATUM L. St. Croix (according to West).
PIPER AURITUM Kunth, is recorded by C. de Candolle, with doubt,
as collected on St. Thomas by Friedericksthal; the record is probably
erroneous.
PIPER TENUIFLORUM Vahl, St. Croix (according to West). A
species ‘not understood by modern botanists.
PIPER INCURVUM Sieb., is recorded from St. Croix; the record is
questioned by C. de Candolle.
40 BROOKLYN BOTANIC GARDEN MEMOIRS
PIPER RETROFRACTUM Vahl. Cultivated on St. Thomas.
POTOMORPHE PELTATA (L.) Mig. [Piper peltatum L.; P. umbel-
latum L.] Forests, shaded banks and along rivulets, St. Thomas;
StCroix:
PEPEROMIA GLABELLA (Sw.) A. Dietr. [Piper glabellum Sw.]|
On trees and rocks in forests, St. Thomas; St. Jan.
PEPEROMIA ALATA C. DC. [P. cubana of de Candolle, in part.]
On trees, St. Croix.
PEPEROMIA PELLUCIDA (L.) H.B.K. [Piper pellucidum L.] On
walls and in wet shade, St. Thomas; St. Jan; St. Croix; in forests,
St. Croix (according to Eggers).
PEPEROMIA SCANDENS R. & P. is recorded by C. de Candolle as
found by Friederichsthal on St. Thomas (Prodr. 16': 434, 1869);
but in his description of West Indian Piperaceae (Urban Symb. Ant.
3: 229. 1902), St. Thomas is not cited. The earlier record is, pre-,
sumably, erroneous.
PEPEROMIA GUADALUPENSIS C. DC. [Piper acuminatum of West;
P. acuminata of Eggers, in part.] St. Croix, according to de Candolle,
collected by West; on rocks in forests, all islands (according to Eggers).
PEPEROMIA HAMILTONIANA Miquel. [P. Hamiltoniana emarginula
C. DC.; P. acuminata of Millspaugh.] Shaded rocks, St. Croix.
PEPEROMIA MAGNOLIAEFOLIA (Jacq.) A. Dietr. [Piper magnoliae-
folium Jacq.; ? Piper obtusifolium of West; Peperomia obtusifolia and
P. obtusifolia clusiaefoia of Eggers.| In woodlands, St. Thomas;
St. Croix:
PEPEROMIA HUMILIs (Vahl) A. Dietr. [Piper humile Vahl; Peper-
omia Langsdorffii Miq.; P. polystachya of Millspaugh.] Shaded rocks,
St. Thomas::St. Jans St: Croix.
PEPEROMIA MYRTIFOLIA (Vahl) A. Dietr. [Piper myrtifolium
Vahl.] St. Croix, collected only by Pflug. Endemic.
PEPEROMIA POLYSTACHYA (Ait.) Mig. [Piper polystachyon Ait.]
St. Croix (according to Hooker); among rocks in forests, all islands
(according to Eggers). Perhaps not distinct from P. humilis.
PEPEROMIA RUPERTIANA C. DC.(?) Wet, shaded bank, Rosen-
berg, St. Jan. Determined from a barren specimen, and identification
therefore doubtful.
SALICACKEAE
SALIX CHILENSIS Molina. [S. Humboldtiana Willd.] In water,
near Grove Place, St. Croix.
ULMACEAE
CELTIS TRINERVIA Lam. Forests and thickets, St. Thomas;
Sie lan:
BRITTON: FLORA OF THE VIRGIN ISLANDS 4]
MomisIA IGUANAEA (Jacq.) Rose & Standley. [Rhamnus iguanaea
Jacq.; Celtis aculeata Sw.; Celtis aculeata serrata Eggers.] Thickets,
ef Phomas; St. Jan; St. Croix.
TREMA MICRANTHUM (L.) Blume. [Rhamnus muicranthus L.;
Celtis micrantha Sw.; Sponia micrantha Dcne.| Woodlands, St.
Mhomas; St. Jan; St. Croix.
MORACEAE
CHLOROPHORA TINCTORIA (L.) Gaud. [Morus tinctoria L.; Mac-
lura tinctoria D. Don.] Woodlands, St. Thomas; St. Jan (according
to Eggers); St. Croix.
ARTOCARPUS INCISA L. f. Hillsides and valleys, naturalized and
planted, St. Thomas; St. Jan; St. Croix.
Ficus URBANIANA Warburg. [Ficus crassinervia of Eggers in part,
and of Millspaugh.] Woods, St. Croix. Sometimes planted.
FICUS CRASSINERVIA Desf. [Ficus trigonata of Eggers.] Forests,
St. Thomas; St. Croix.
Ficus LAEVIGATA Vahl. [Ficus lentiginosa Vahl; Ficus populnea
Willd.; F. thomae Miq.; F. sancti-crucis Miq.; F. pedunculata Vahl.]
Forests, woodlands and hillsides, St. Thomas; St. Jan; St. Croix.
Ficus Carica L. Planted for its fruit.
Ficus ELASTICA Roxb. Planted for shade and ornament.
CECROPIA PELTATA L. Forests and hillsides, St. Thomas; St.
Jan; St: Croix.
URTICACEAE
URERA ELATA (Sw.) Griseb. [Urtica elata Sw.] Spring Garden,
St. Croix, collected by West; Eggers records West’s specimen as
preserved in the Copenhagen herbarium.
URERA BACCIFERA (L.) Gaud. [Urtica baccifera L.] is cited by
Eggers as recorded from St. Thomas by Weddell in de Candolle’s
Prodromus 16!: 93, but an examination of pages 93 and 94 of that work
does not verify the citation, and the plant is otherwise unknown from
these islands.
UrRTICA ELONGATA Vahl (St. Croix, West) is a species unknown to
modern botanists.
FLEURYA AESTUANS (L.) Gaud. [Urtica aestuans L.] On rocks,
walls and in forests, St. Thomas; St. Jan; St. Croix.
PILEA MICROPHYLLA (L.) Liebm. [Parietaria microphylla L.;
Adicea microphylla Kuntze; P. microphylla trianthemoides and succu-
lenta of Eggers; Adicea microphylla trianthemoides and succulenta of
Millspaugh.] Rocky situations, St. Thomas; St. Jan; St. Croix.
PILEA TENERRIMA Miquel. Shaded banks, St. Jan.
42 BROOKLYN BOTANIC GARDEN MEMOIRS
PrLEA RicHarpiI Urban. St. Thomas, collected by L. C. Richard,
the specimen preserved in the Copenhagen herbarium (according to
Urban). Endemic.
PILEA INAEQUALIS (Juss.) Wedd. [Urtica inaequalis Juss.; Adicea
inaequalis Kuntze.] On rocks in forests, Signal Hill and Crown,
St. Thomas.
PILEA SANCTAE-CRUCIS Liebm. [Adicea sanctae-crucis Kuntze;
Pilea semidentata of Eggers; Pilea grandis of Eggers.] Forests, St.
Thomas: of jams) St. Croix,
PILEA NUMMULARIAEFOLIA (Sw.) Wedd. [Urtica nummulariae-
folia Sw.; Adicea nummulariaefolia Kuntze.] Shaded situations, St.
Thomas: ot. Croix,
PILEA GRANDIFOLIA (L.) Blume. [Pilea grandis Wedd.] Re-
corded by de Candolle (Prodr. 16!: 143) as from Jamaica and St.
Thomas, is confined to Jamaica, where there is a parish of St. Thomas.
ROUSSELIA HUMILIS (Sw.) Urban. [Urtica humilis Sw.; U. lappu-
lacea Sw.; Rousselia lappulacea Gaud.] Shaded situations, St.
Thomas.
OLACACEAE
SCHOEPFIA SCHREBERI Gmelin. [Codoniwm arborescens Vahl; S.
arborescens R. & S.]_ Woodlands, St. Thomas; St. Croix.
LORANTHACEAE
DENDROPEMON CARIBAEUS Krug & Urban. [Loranthus emar-
ginatus of Eggers; Phthirusa caribaea Engler.| On trees, St. Thomas;
Ste lanesSte Crore
PHORADENDRON CHRYSOCARPUM Krug & Urban. [Phoradendron
flavens of Eggers; P. martinicense of Millspaugh.] On trees, St.
Thomas7*St. ‘Crom
PHORADENDRON TRINERVIUM (Lam.) Griseb. is recorded by Trelease
as represented in the Ventenat Herbarium by a specimen from St.
Thomas; it is otherwise unknown from the islands.
PHORADENDRON RACEMOSUM (Aubl.) Krug & Urban. [P. penni-
nervium O. Kuntze] is recorded by O. Kuntze as from St. Thomas,
apparently erroneously; the specimen was probably from Porto Rico.
ARISTOLOCHIACEAE
ARISTOLOCHIA ODORATISSIMA L. Hillside thickets, St. Jan.
ARISTOLOCHIA TRILOBATA L. Thickets, St. Thomas; St. Jan; St.
Croix (according to West).
ARISTOLOCHIA ANGUICIDA L. Thickets, St. Croix.
ARISTOLOCHIA RINGENS Vahl. Cultivated on St. Croix (according
to West).
BRITTON: FLORA OF THE VIRGIN ISLANDS 43
POLYGONACEAE
CoccoLosis Kruciu Lindau. Rocky Hills, Little St. James
Island, St. Jan.
COccCOLOBIS PYRIFOLIA Desf. [C.Kunthiana Meissn.; C. pyrifolia
Jacquini of Eggers.]|_ St. Thomas (according to Lindau).
COCCOLOBIS OBTUSIFOLIA Jacq. [C. microstachys Willd.; C.
microstachya ovalifolia Meissn.; C. punctata microstachya of Eggers;
C. punctata parvifolia of Millspaugh.] Thickets, St. Thomas; St.
fans ot.. Croix.
CoccoLosis KiotrzscHIANA Meissn. St. Thomas and St. Croix
(according to Lindau). Endemic. Perhaps not distinct from the
preceding species.
COcCOLOBIS DIVERSIFOLIA Jacq. [C. barbadensis Jacq.; C. punc-
tata of Eggers; C. coronata. of Millspaugh.] Woods and thickets, St.
@homas; St. Jan; St. Croix.
COCCOLOBIS LAURIFOLIA Jacq. [C. leoganensis of Eggers.] Thick-
ets, St. Croix.
CoccoLoBIs RUGOSA Desf. St. Thomas (according to de Candolle,
a specimen being preserved in the Delessert Herbarium); not known
to be on St. Thomas at the present time but may have been there
before the forests were cut away; known otherwise only from Porto
Rico.
CoccoLtopis UviFERA (L.) Jacq. [Polygonum Uvifera L.; C.
leoganensis Jacq.; Uvifera leoganensis Kuntze.|] Coastal thickets and
locally on hills, St. Thomas; St. Jan; St. Croix.
CoccoLoBiIs VENOSA L. [C. excoriata L.; C. nivea Jacq.] Woods
and hillsides, St. Thomas; St. Croix.
ANTIGONUM CINERASCENS M. & G. [A. cordatum of Eggers and of
Millspaugh.] Roadsides, St. Thomas; cultivated for ornament, St.
Thomas and St. Croix.
FaGcopyruM FacopyrumM (L.) Karst. [Polygonum Fagopyrum L.|
Planted for food.
RuUMEX VESICARIUS L. Recorded by Eggers as cultivated.
MUHLENBECKIA PLATYCLADA (F. Muell.) Lindau. Planted for
interest.
CHENOPODIACEAE
CHENOPODIUM MURALE L. Walls and waste grounds, St. Thomas;
mt. Croix.
CHENOPODIUM AMBROSIOIDES L. [? C. cuneifolium Vahl.] Walls
and waste grounds, St. Thomas; St. Jan; St. Croix.
ATRIPLEX PENTANDRA (Jacq.) Standley. [Axyris pentandra Jacq. ;
Atriplex cristata H. & B.; Obione cristata Moq.] Coastal sands, St.
Thomas; St: Jan; -St. Croix.
44 BROOKLYN BOTANIC GARDEN MEMOIRS
SALICORNIA PERENNIS Mill. [S. ambigua Michx.] Salt marshes,
St.-Croix:
BETA VULGARIS L. Cultivated for food.
AMARANTHACEAE
CELOSIA NITIDA Vahl. [? C. paniculata of Schlechtendal.] Woods
and thickets, St. Thomas; St. Jan; St. Croix.
CELOSIA ARGENTEA L. [C. margaritacea L.] Waste and culti-
vated grounds, St. Thomas; St. Croix.
CELOSIA CRISTATA L. Planted for ornament.
CHAMISSOA ALTISSIMA (Jacq.) H.B.K. [Achyranthes altissima
Jacq.; Kokera paniculata Kuntze.] Forests and thickets, St. Thomas;
St Croix.
AMARANTHUS DuBIUS Mart. [A. tristis Willd., not L.; A. pantcu-
latus of Eggers and of Millspaugh.] Waste grounds, St. Thomas;
St. Janz ot. Croix:
AMARANTHUS SPINOSUS L. Waste and cultivated grounds, St.
Thomas; St. Croix.
AMARANTHUS CRASSIPES Schl. [Scleropus amarantoides Schrad.]
Dry soil, waste and cultivated grounds, St. Thomas; St. Croix.
AMARANTHUS CAUDATUS L. St. Croix (according to West).
AMARANTHUS POLYGONOIDES L. [Amblyogyne polygonoides Raf.]
Sandy soil, roadsides and waste grounds, St. Thomas; St. Croix.
AMARANTHUS GRACILIS Desf. [Chenopodium caudatum Jacq.;
? Amaranthus oleraceus of West; Euxolus caudatus Moq.; E. oleraceus
of Eggers.] Waste and-cultivated grounds, St. Thomas; St. Jan;
St. Croix,
AMARANTHUS GANGETICUS L._ [A. incomptus Willd.; A. tricolor L.]
Planted for ornament.
CENTROSTACHYS INDICA (L.) Standley. [Achyranthes aspera indica
L.; Achyranthes aspera obtusifolia Griseb.; A. aspera simplex
Millsp.] Waste and cultivated grounds, St. Thomas; St. Croix.
CENTROSTACHYS ASPERA (L.) Standley. [Achyranthes aspera L.;
A. argentea Lam.] is recorded from the islands by Eggers, but I have
seen no specimens nor find any other record; the plant occurs, how-
ever, on Tortola.
ACHYRANTHES POLYGONOIDES (L.) Lam. [Gomphrena polygonoides
L.; Alternanthera polygonoides R. Br.; Alternanthera paronychioides
St. Hil.] Waste grounds, St. Thomas.
ACHYRANTHES REPENS L. [Alternanthera Achyrantha R. Br.;
A. paronychioides of Millspaugh.] Rocky waste places, St. Thomas;
St. Grom
ACHYRANTHES FICOIDEA (L.) Standley. [Gomphrena ficoidea L.;
BRITTON: FLORA OF THE VIRGIN ISLANDS 45
Illecebrum ficoideum L.; Alternanthera ficoidea R. Br.] Moist places
and on shores, St. Thomas.
ACHYRANTHES PORTORICENSIS (Kuntze) Standley. [Alternanthera
portoricensis Kuntze.] Rocky hills, Little St. James Island, St. Jan.
GOMPHRENA GLOBOSA L. Subspontaneous after cultivation, St.
Thomas; St. Croix.
IRESINE ANGUSTIFOLIA Euphr. _ [J. elatior L. C. Rich.] Thickets,
and banks, St. Thomas; St. Croix.
PHILOXERUS VERMICULATUS (L.) R. Br. [Illecebrum vermiculatum
L.; Lithophila vermiculata Uline; Iresine vermicularis Miq.] Saline
soil along the coasts, St. Thomas; St. Croix.
LITHOPHILA MUSCOIDES Sw. Rocks on the shore, Judith’s Fancy,
et. Croix.
NYCTAGINACEAE
MIRABILIS JALAPA L. [M. dichotoma L.! Waste grounds, St.
Thomas; St. Croix.
BOERHAAVEA ERECTA L. Waste and cultivated ground, St. Croix.
BOERHAAVEA COCCINEA Mill. [B. paniculata L. C. Rich.; B.
diffusa Sw.; B. decumbens Vahl; B. hirsuta Willd.; B. diffusa panicu-
lata Kuntze; B. repens of Millspaugh.] Dry soil, St. Thomas; St.
ox sSt. Jan.
COMMICARPUS SCANDENS (L.) Standley. [Boerhaavea scandens L.}|
Rocky hillsides, St. Thomas.
PISONIA ACULEATA L. Hillsides, woods and thickets, St. Thomas;
Si, Croix:
PISONIA SUBCORDATA Sw. [P. nigricans of West.] Thickets and
woods, St. Thomas; St. Croix.
TORRUBIA FRAGRANS (Dum.-Cours.) Standley. [Pisonia fragrans
Dum.-Cours.; Pisonia inermis of Eggers, of Kuntze and of Millspaugh;
? P. coccinea of West.] Forests and thickets, St. Thomas; St. Croix.
BOUGAINVILLEA SPECTABILIS Willd. Cultivated for ornament.
EGGERSIA BUXIFOLIA Hook. f. [Neea buxifolia Heimerl.] Dry
hillsides, St. Thomas; St. Jan.
BATIDACEAE
BATIS MARITIMA L. Shores of salt water lagoons, St. Thomas;
oh. Croix.
Pry TOLACGACE AE,
RivinA HumILIs L. [R. laevis L.; Tithonia humilis Kuntze.]
Dry, shaded situations, St. Thomas; St. Jan; St. Croix.
TRICHOSTIGMA OCTANDRUM (L.) H. Walt. [Rivina octandra L.;
Villamilla octandra Hook. f.; Rivina scandens Mill.| Woods and
thickets, St. Thomas; St. Jan; St. Croix.
46 BROOKLYN BOTANIC GARDEN MEMOIRS
PETIVERIA ALLIACEA L. Dry, shaded situations, St. Thomas;
St.lane St Croiz
MICROTEA DEBILIS Sw. Waste grounds, St. Thomas; St. Croix.
AIZOACEAE
MOLLUGO VERTICILLATA L. Dry soil, Buck Island, St. Thomas;
MOLLUGO NUDICAULIS L. Rocky soil and banks, St. Thomas.
St. /Crotx.
SESUVIUM PORTULACASTRUM L. [Halimus portulacastrum Kuntze. |
Saline soil, St. Thomas; St. Jan; St. Croix.
TRIANTHEMA PORTULACASTRUM L. [T7. monogynum L.] Waste
and rocky places. St. Thomas; St. Croix.
CyYPSELEA HUMIFUSA Turp. Around a small fresh-water lagoon,
Water Island, St. Thomas (according to Eggers).
PORTULACACEAE
TALINUM TRIANGULARE (Jacq.) Willd. [Portulaca triangularis
Jacq.] Rocky soil, St. Thomas; St. Croix.
TALINUM PANICULATUM (Jacq.) Gaertn. [Portulaca paniculata
Jacq.; P. patens Jacq.; Talinum patens Willd.] Rocky soil, St.
Thomas: St: Jan; St-Gror.
PORTULACA OLERACEA L. _ [P. oleracea macrantha and (?) micrantha
Eggers.] Sunny situations, St. Thomas; St. Jan; St. Croix.
PoRTULACA PILOSA L. Cultivated and waste grounds, St. Thomas;
St. Cron.
PORTULACA HALIMOIDES L. Sunny situations, St. Thomas; St.
Jan; St» Croix
PORTULACA QUADRIFIDA L. Waste and cultivated grounds, St.
DPhomassoe aan jobeCrom.
PORTULACA BRASILIENSIS West, of St. Croix, is not described.
The plant was probably one of the preceding species.
BASELLACEAE
BOUSSINGAULTIA LEPTOSTACHYS Mogq. [B. baselloides of Eggers.]
Naturalized in gardens and cultivated, St. Thomas; St. Croix.
BASELLA RUBRA L. Cultivated on St. Croix.
ALSINACEAE
DRYMARIA CORDATA (L.) Willd. [Holostewm cordatum L.; D.
cordata diandra Eggers.] Shaded moist places, St. Thomas; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 47
NYMPHAEACEAE
CASTALIA AMPLA (DC.) Salisb. [Nymphaea ampla DC.; N. ampla
parvifolia Eggers.] In rivulets and ponds, St. Croix.
MENISPERMACEAE
CISSAMPELOS PARIERA L. [C. muicrocarpa DC.] Woods and
thickets, St. Thomas; St. Croix; St. Jan.
HYPERBAENA LAURIFOLIA (Poir.) Urban. [Cissampelos laurifolius
Poir.; Cocculus laurtfolius of Eggers.| Woodlands, St. Thomas.
HYPERBAENA DOMINGENSIS (DC.) Benth. [Cocculus domingensis
DC.| Forest, near St. Peter, St. Thomas (according to Eggers). The
species is not accredited to St. Thomas by Urban (Symb. Ant. 1: 306).
Eggers’s record probably refers to H. laurtfolia.
ANNONACEAE
OXANDRA LAURIFOLIA (Sw.) A. Rich. [Uvaria laurifolia Sw.;
U. excelsa Vahl.] St. Croix (according to West).
GUATTERIA CARIBAEA Urban. [G. Ouregou Griseb., not Dunal.]
St. Thomas (according to Grisebach).
ANNONA MURICATA L. Woods and thickets, St. Thomas; St. Jan;
et. Croix.
ANNONA PALUSTRIS L. [A. glabra L.; A. laurifolia Dunal.]
Borders of marshes and coastal thickets, St. Thomas; St. Croix.
ANNONA SQUAMOSA L. [Annona cinerea Dunal.] Woods, hill-
sides and thickets, St. Thomas; St. Jan; St. Croix.
ANNONA RETICULATA L. Woods and hillsides, St. Thomas; St.
Waits ot. Croix.
ANNONA MONTANA Macf. Fredericksted, St. Croix.
LAURACEAE
HUFELANDIA PENDULA (Sw.) Nees. [Laurus pendula Sw.; H.
thomae Nees.| St. Thomas (DC. Prodr..15!: 65); recorded from St.
Thomas as collected by Riedlé (Mez, Jahrb. Bot. Gart. Berlin 5: 21).
ACRODICLIDIUM SALICIFOLIUM (Sw.) Griseb. Forests, St. Thomas;
Sian; St. Croix.
PERSEA PERSEA (L.) Cockerell. [Laurus Persea L.; Persea
americana Mill.; P. gratissima Gaertn. f.] | Spontaneous after plant-
in@, St. Thomas; St. Croix.
PHOEBE CUBENSIS Nees. [P. antillana cubensis Meissn.; P.
elongata of Eggers.] St. Croix (West, according to DC. Prodr. 15!:
31, and also recorded by Mez).
48 BROOKLYN BOTANIC GARDEN MEMOIRS
OcCOTEA LEUCOXYLON (Sw.) Mez. [Laurus leucoxylon Sw.; Ore-
odaphne leucoxylon Nees.| Forest, Signal Hill, St. Thomas.
OcOTEA FLORIBUNDA (Sw.) Mez. [Laurus floribunda Sw.] Wood-
ed hillside, Bordeaux, St. Jan.
NECTANDRA ANTILLANA Meissn. Forests, St. Thomas.
NECTANDRA MEMBRANACEA (Sw.) Griseb. [Laurus membranacea
Sw.] Dense forests, Signal Hill, St. Thomas; Will’s Bay, St. Croix
(according to Eggers).
NECTANDRA CORIACEA (Sw.) Griseb. [Laurus coriacea Sw.]
Forests, St. Thomas; St. Jan; St. ‘Croim:
LAURUS INDICA West, of St. Croix, is unknown to modern botanists.
LAURUS LONGIFOLIA Vahl, collected on St. Croix by West, is not
further determined.
CINNAMOMUM ZEYLANICUM Blume is recorded by Eggers as natur-
alized in a few places in shaded valleys on St. Croix.
CASSY THACEAE
CASSYTHA AMERICANA Nees. Coastal thickets, St. Thomas; St.
Croix. : :
PAPAVERACEAE
ARGEMONE MEXICANA L. Waste and cultivated grounds, St.
Thomas; St. Jan; St: Croix.
CAPPARIDACEAE
CLEOME SPINOSA Jacq. [C. pungens Willd.] Waste and culti-
vated grounds, St. Thomas; St. Jan; St. Croix.
CLEOME GYNANDRA L. [C. pentaphylla L.; Gynandropsis penta-
phylla DC.] Waste and cultivated grounds, St. Thomas; St. Jan;
Sit. Croix:
CLEOME viscosa L. [Polanisea icosandra of Millspaugh.] Waste
and cultivated grounds, St. Thomas; St. Croix.
CAPPARIS CYNOPHALLOPHORA L. [C. jamaicensis Jacq.; C. toru-
losa of West; C. jamaicensis marginata and siliquosa Eggers; ? C.
linearis of West.] Thickets and hillsides, St. Thomas; St. Jan; St.
Croix:
CAPPARIS INDICA (L.) Fawc. & Rend. ([Breynia indica L.; Cap-
paris Breynia Jacq.; C. amygdalina Lam.; C. Grisebachii of Mills-
paugh.] Thickets, woods and hillsides, St. Thomas; St. Jan; St.
Croix.
CApPARIS BADUCCA L. [Capparis frondosa Jacq.| Forests and
hillsides, St. Thomas; St. Croix; St. Jan (according to Eggers).
CAPPARIS FLEXUOSA L. [Capparis cynophallophora of Eggers and
BRITTON: FLORA OF THE VIRGIN. ISLANDS 49
of Millspaugh; C. saligna of West; C. cynophallophora saligna Eggers. |
Woods, thickets and hillsides, St. Thomas; St. Jan; St. Croix.
CAPPARIS COCCOLOBIFOLIA Mart. [C. cynophallophora latifolia
Griseb.] Thickets and hillsides, St. Thomas; St. Jan; St. Croix.
CAPPARIS PORTORICENSIS Urban. Hillside thicket between Be-
thania and Rosenberg, St. Jan.
CAPPARIS VERRUCOSA Jacq. St. Thomas (according to Grisebach) ;
Virgin Islands (according to Eggers).
MorIsONIA AMERICANA L. Wooded hillsides, St. Thomas; St.
Jan; St. Croix (according to Eggers). .
BRASSICACEAE
LEPIDIUM VIRGINICUM L. [L. apetalum of Millspaugh.] Waste
and cultivated grounds, St. Thomas; St. Jan; St. Croix.
LEPIDIUM SATIVUM L. Cultivated for condiment.
CAKILE LANCEOLATA (Willd.) O. E. Schulz. [C. aequalis L’Her.]
Coastal sands and rocks, St. Thomas; St. Jan; St. Croix.
SINAPIS ARVENSIS L. Naturalized near Anguilla, St. Croix (ac-
_ cording to Eggers).
BRASSICA INTEGRIFOLIA (West) O. E. Schulz. [Sinapis integrifolia
West; S. brassicata Griseb., not L.; S.juncea of Millspaugh.] Waste
and cultivated grounds, St. Thomas; St. Croix; St. Jan.
BRASSICA OLERACEA L. Cultivated for food.
BRASSICA CAMPESTRIS L. [B. Rapa L.| Cultivated for food.
SISYMBRIUM NASTURTIUM L. [Nasturtium officinale R. Br.| In
rivulets, St. Thomas; St. Croix.
RAPHANUS RAPHANISTRUM L. Recorded by West.
RAPHANUS SATIVUS L. Cultivated for food.
KONIGA MARITIMA (L.) R. Br. [Clypeola maritima L.| Culti-
vated for ornament.
MORINGACEAE
MorincGa MortneGa (L.) Millsp. [Guilandina Moringa L.; Hyper-
anthera Moringa Vahl; Moringa pterygosperma Gaertn.| Roadsides
and hillsides, St. Thomas; St. Jan; St. Croix.
CRASSULACEAE
BRYOPHYLLUM PINNATUM (Lam.) Kurz. [Cotyledon pinnata Lam. ;
B. calycinum Salisb.] Dry soil, St. Thomas; St. Jan; St. Croix.
ROSACEAE
A number of different kinds of roses are grown for ornament.
5
50 BROOKLYN BOTANIC GARDEN MEMOIRS
AMYGDALACEAE
CHRYSOBALANUS Icaco L. Woods, hillsides and thickets, St.
Thomas; St. Janest. Croix.
MIMOSACEAE .
INGA LAURINA (Sw.) Willd. [Mimosa laurina Sw.; M. laurifolia
of West.] Woodlands and forests, St. Thomas; St. Jan; St. Croix.
PITHECOLOBIUM UNGutIs-caTI (L.) Mart. [Mimosa unguts-cati L.;
Inga Unguts-cati Willd.; P. Unguis-cati forfex Griseb.| Thickets,
hillsides and pastures, St. Thomas; St. Jan; St. Croix.
ENTEROLOBIUM SAMAN (Jacq.) Prain. [Mimosa Saman Jacq.;
Calliandra Saman Griseb.; Pithecolobium Saman Benth.] Roadsides
and near settlements, St. Thomas; St. Croix.
AvBizziA LEBBECK (L.) Benth. [Mimosa Lebbeck L.; M. speciosa
Jacq.; Acacia Lebbeck Willd.| Fields and hillsides, St. Thomas; St.
Croix.
Anneslia portoricensis (Jacq.) Britton. [Mimosa _ portoricensis
Jacq.; Calliandra portoricensis Benth.] Forests, King’s Hill, St. Jan.
Anneslia haematostoma (Bert.) Britton. [Acacia haematomma
Bert.; Calliandra haemotomma Benth.] Flag Hill, St. Thomas.
Anneslia purpurea (L.) Britton. [Calliandra purpurea Benth.;
Mimosa purpurea L.; Inga purpurea Willd.| Cultivated on St. Croix
(according to West).
ACACIA NUDIFLORA Willd. Hiullsides and woods, St. Thomas; St.
Jan.
ACACIA RIPARIA H.B.K. [A. sarmentosa Griseb., not Desv.;
Mimosa paniculata of West; A. Westiana DC.] Hillsides and thickets,
St. Thomas; St. Jan; St. Croix (according to West).
Acacia CatEecHu Willd. [Mimosa catechu L. f.] Naturalized in
shaded valleys, St. Croix.
ACACIA MACRACANTHA H. & B. [Mimosa lutea Houst.; Acacia
lutea Hitche., not Leavenw.; A. macracantha glabrens Eggers.] Hill-
sides and thickets, St. Thomas; St. Jan; St. Croix.
Acacia ToRTUOSA (L.) Willd. [Mimosa tortuosa L.| Hillsides,
St, Dhomas St. Croix.
ACACIA ARABICA Willd., is planted for ornament, and is recorded
by Eggers as naturalized near dwellings on St. Thomas and St. Croix.
VACHELLIA FARNESIANA (L.) W. & A. [Mimosa Farnesiana L.;
Acacia Farnesiana Willd.| Hillsides and thickets, St. Thomas; St.
Jan; StVCrax:
LEUCAENA GLAUCA (L.) Benth. [Mimosa glauca L.; Acacia
frondosa Willd.] Fields and hillsides, St. Thomas; St. Jan; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 51
Mimosa pupicA L. Fields and hillsides, St. Thomas; St. Croix.
Mimosa CERATONIA L. [? M. sensitiva of West.] Hills and
thickets, St. Thomas; St. Jan; ? St. Croix (according to West).
Mimosa picrA L. [M. asperata L.] St. Thomas (according to
Grisebach), who indicates that he saw a specimen from that island,
but the plant has not been found there by recent collectors.
ACUAN VIRGATUM (L.) Medic. [Mimosa virgata L.; Desmanthus
virgatus Willd.; D. strictus Bertol.; D. virgatus strictus Griseb.; A.
virgatus albiflorus Kuntze.| Fields and hillsides, St. Thomas; St.
Croix.
ACUAN DEPRESSUM (H. & B.) Kuntze. [Desmanthus depressus H.
& B.| Hillsides, St. Thomas; St. Jan.
NEPTUNIA PUBESCENS Benth. Buck Island, St. Thomas (accord-
ing to Eggers).
ADENANTHERA PAVONINA L. Naturalized, St. Thomas; St. Jan
and St. Croix (according to Eggers); not seen by us on any of the
islands, except as a planted tree.
CAESALPINACEAE
HYMENAEA COURBARIL L. Woods and hillsides, St. Thomas; St.
wan: St. Croix.
TAMARINDUS INDICA L.,. In various situations, St. Thomas; St.
Jan; St. Croix. Planted for its fruit and for shade.
BAUHINIA TOMENTOSA L. Spontaneous after cultivation, waste
grounds, St. Thomas; St. Croix.
BAUHINIA MONANDRA Kurz. [B. Kappleri Sagot; B. Krugii
Urban.] Spontaneous after planting, St. Thomas; St. Croix.
BAUHINIA UNGULA Jacq., recorded by Grisebach as found on St.
Thomas, is probably an error in record or determination.
Cassia Fistuta L. Naturalized in shaded valleys, St. Croix
(according to Eggers). Planted for shade and for its fruit.
CASSIA GRANDIS L. Naturalized on St. Thomas and St. Croix.
CASSIA QUINQUANGULATA L. C. Rich. [C. bacillaris of Eggers, of
Kuntze and of Millspaugh.] Woods and thickets, St. Thomas.
CASSIA BICAPSULARIS L. Waste grounds and hillsides, St. Thomas;
ee jan; St. Croix.
CASSIA SIAMEA Lam. [C. florida Vahl.] Naturalized near towns,
St. Thomas (according to Eggers). Planted for shade and ornament.
CASSIA OCCIDENTALIS L. [C. planisiliqua L.] Waste and culti-
vated grounds, St. Thomas; St. Jan; St. Croix.
“CassiA Tora L. [C. obtusifolia L.] Waste and cultivated grounds,
St. Thomas; St. Croix.
52 BROOKLYN BOTANIC GARDEN MEMOIRS
CASSIA POLYPHYLLA Jacq. [C. biflora angustisiliqua of Eggers.]
Hillsides, St. Thomas; St. Croix.
CassIA OBOVATA Collad. Naturalized on St. Thomas (according
to Eggers).
CAssIA HIRSUTA L. Sugar Estate, St. Thomas (according to
Eggers).
CassiA ALATA L. Waste grounds, St. Thomas; St: “janie
Cro.
CAssIA AUGUSTIFOLIA Vahl. Planted on St. Croix (according to
West).
CASSIA TRIFLORA Vahl, collected on St. Croix by Rohr, is “‘a doubt-
ful species’ according to Eggers; it is not the same as Cassia triflora
Jacquin, a prior name.
CassIA GRANDIS L. [C. mollis Vahl.] Planted for shade.
CHAMAECRISTA GRAMMICA (Spreng.) Pollard. [Cassia grammuica
Spreng.] Rocky soil, Little St. James Island, St. Jan.
CHAMAECRISTA SWARTZII (Wickstr.) Britton. [Cassia Swarizi
Wickstr.; C. glandulosa of West; Cassia glandulosa stricta and ramosa
of Eggers; Chamaecrista glandulosa and C. glandulosa ramosa of Mills-
paugh; Chamaecrista complexa Pollard; Cassia Chamaecrista pubi-
caulis Kuntze.] Fields and hillsides, St. Thomas; St. Jan; St. Croix.
CHAMAECRISTA CHAMAECRISTA (L.) Britton. [Cassia Chamae-
crista L.; Cassia nictitans of Eggers and of Millspaugh; C. diffusa DC.]
Dry, grassy situations, St. Thomas; St. Jan; St. Croix.
PARKINSONIA ACULEATA L. Coastal thickets, St. Thomas; St.
Jango Croix.
HAEMATOXYLON CAMPECHIANUM L. [Sabinea florida of Mills-
paugh.] Coastal thickets and hillsides, St. Thomas; St. Jan; St.
Croix.
DELONIX REGIA (Bojer) Raf. [Poinciana regia Bojer.| Spon-
taneous after planting, St. Thomas; St. Croix.
GUILANDINA CrisTA (L.) Small. [Caesalpinia Crista L.; Gutlan-
dina Bonducella L.| Coastal sands, St. Thomas; St. Jan; St. Croix.
GUILANDINA DIVERGENS (Urban) Britton. [Caesalpina divergens
Urban; Guwilandina Bonduc of Schlechtendal and of Eggers.] Coastal
thickets, St. Thomas; St. Jan; St. Croix.
GUILANDINA MELANOSPERMA Eggers. [Caesalpinia melanosperma
Urban.] Coastal thickets, St. Croix.
CAESALPINIA CORIARIA Willd. [Poinciana coriaria Jacq.; Lebidibia
coriaria Schl.| Hillsides, St. Thomas. Planted on St. Croix.
CAESALPINIA GILLIES Wall. [Poinciana Gilliesti Hook.] Plant-
ed for ornament.
CAESALPINIA PUNCTATA Willd. Planted on St. Thomas.
BRITTON: FLORA OF THE VIRGIN ISLANDS 53
CAESALPINIA SAPPAN L. Recorded by Krebs as planted on St.
Thomas. .
CAESALPINIA ELATA Sw. Attributed by Eggers to St. Croix, pre-
sumably erroneously.
POINCIANA PULCHERRIMA L. [Caesalpinia pulcherrima Sw.| Spon-
taneous after cultivation, St. Thomas; St. Croix.
KRAMERIACEAE
KRAMERIA IxtnA L. [K. Ishami Millsp.]_ Dry rocky soil, Bovoni
and Water Island, St. Thomas.
FABACEAE
MYROSPERMUM FRUTESCENS Jacq. Naturalized near dwellings,
St. Croix (according to Eggers).
SOPHORA TOMENTOSA L. Coastal sands, St. Thomas; St. Croix.
CROTALARIA RETUSA L. Waste and cultivated grounds, St.
Thomas; St. Jan; St. Croix.
CROTALARIA JUNCEA L. Field at Bassin, St. Croix; recorded by
West as cultivated prior to 1793.
CROTALARIA VERRUCOSA L. Waste and cultivated grounds, St.
@uemas; St: Croix; St. Jan.
CROTALARIA INCANA L. Waste and cultivated grounds, St.
Thomas; St. Croix.
CROTALARIA LOTIFOLIA L. Thickets and hillsides, St. Thomas;
Sian, St. Croix.
CROTALARIA LABURNIFOLIA L. Cultivated on St. Croix (according
to West). ;
INDIGOFERA SUFFRUTICOSA L. [J. Anil L.] Thickets and hill-
sides, St. Thomas; St. Jan; St. Croix.
INDIGOFERA GUATEMALENSIS Mog. & Sessé. St. Thomas.
INDIGOFERA TINCTORIA L. Thickets, St. Thomas; St. Jan; St.
Croix (according to Eggers, who notes its former cultivation).
MEeEDICAGO SATIVA L. Planted on St. Croix.
PAROSELA DOMINGENSIS (DC.) Millsp. [Dalea domingensis DC.;
D. phymatodes of Eggers.]_ Dry soil, St. Jan, collected only by Eggers.
CRACCA CINEREA (L.) Morong. [Galega cinerea L.; G. littoralis L.;
Tephrosia cinerea Pers.; Cracca villosa cinerea Kuntze; Tephrosia
cinerea littoralis of Eggers.| Dry sandy soil, St. Thomas; St. Jan;
b.. Croix.
CRACCA PURPUREA L. [Galega purpurea L.] Cultivated on St.
Croix (according to West).
SABINEA FLORIDA (Vahl) DC. [Robinia florida Vahl.| Hillsides
and thickets, St. Thomas; St. Jan.
54 BROOKLYN BOTANIC GARDEN MEMOIRS
BENTHAMANTHA CARIBAEA (Jacq.) Kuntze. ([Galega caribaea
Jacq.; Cracca caribaea Benth; Brittonamra caribaea Kuntze.| Thick-
ets and hillsides, St. Thomas; St. Croix.
COURSETIA ARBOREA Griseb., recorded by Grisebach from St. Jean,
is erroneously quoted by Eggers as from St. Jan. There is a place
called St. Jean in French Guiana.
SESBAN SERICEA (Willd.) DC. [Coronilla sericea Willd.] Thick-
ets, Flag Hill, St. Thomas.
Sesban Sesban (L.) Britton. [Aeschynomene Sesban L.| Planted
on St. Crom.
AGATI GRANDIFLORA (L.) Desv. [Aeschynomene grandiflora L.;
Sesbania grandiflora Pers.| Roadsides and near dwellings, naturalized
St: Thomas; St. Jane ot. Croix.
PICTETIA ACULEATA (Vahl) Urban. [Robinia aculeata Vahl;
R. squamata Vahl; Aeschynomene aristata Jacq.; Pictetia squamata
DC.; P. aristata DC.] Woods, hillsides and thickets, St. Thomas;
St. Jans ot. Grom.
AESCHYNOMENE AMERICANA L. [Ae. americana depila Millsp.]
Grassy places, St. Thomas; St. Jan; St. Croix.
STYLOSANTHES HAMATA (L.) Taubert. [Hedysarum hamatum L.;
Stylosanthes procumbens Sw.] Dry soil, St. Thomas; St. Croix; St.
Jan. ;
STYLOSANTHES VISCOSA Sw., recorded by West from St. Croix.
Eggers thought perhaps a mistake for the preceding species, which is
probable.
ARACHIS HYPOGAEA L. Subspontaneous after cultivation, St.
.Thomas; St. Croix. Hardly persistent.
ZORNIA DIPHYLLA (L.) Pers. [Hedysarum diphyllum L.; Z. reticu-
lata Smith.] Pastures, high hills of St. Thomas; St. Croix (according
to'de Candolle).
CoDARIOCALYX GYRANS (L. f.) Hassk. [Hedysarum gyrans L. f.;
Desmodium gyrans DC.] Planted for interest.
MEIBOMIA TRIFLORA (L.) Kuntze. [Hedysarum triflorum L.;
Desmodium triflorum DC.; Meibomia triflora pilosa Kuntze.] Fields
and moist grassy places, St. Thomas; St. Jan; St. Croix.
MEIBOMIA SUPINA (Sw.) Britton. [Hedysarum supinum Sw.;
H. incanum Sw.; Desmodium supinum DC.; D. incanum DC.]
Fields, hillsides, woods and thickets, St. Thomas; St. Jan; St. Croix.
MEIBOMIA AXILLARIS (Sw.) Kuntze. [Hedysarum axillare Sw.;
Desmodium axillare DC.| Shaded banks and ravines, St. Croix.
MEIBOMIA MOLLIS (Vahl) Kuntze. [Hedysarum molle Vahl;
Desmodium molle DC.| Grassy places, St. Thomas; St. Croix.
MEIBOMIA SPIRALIS (Sw.) Kuntze. [Hedysarum spirale Sw.; Des-
BRITTON: FLORA OF THE VIRGIN ISLANDS 55
modium spirale DC.] Hillsides and banks, St:-Ehomas; (St. Jan:45¢-
Croix.
MEIBOMIA TORTUOSA (Sw.) Kuntze. [Hedysarum tortuosum Sw.;
Desmodium tortuosum DC.|° Banks, hillsides and thickets, St. Thomas;
St Croix.
MEIBOMIA SCORPIURUS (Sw.) Kuntze. [Hedysarum scorpiurus
Sw.; Desmodium scorpiurus Desv.] Grassy places, St. Thomas (ac-
cording to Grisebach); St. Croix (according to Eggers).
LOUREA VESPERTILIONIS (L.) Desv. [Hedysarum vespertilionis L.]
Naturalized in gardens, St. Thomas; St. Croix (according to Eggers).
Planted for ornament.
ALYSICARPUS NUMMULARIFOLIuS (L.) DC. [Hedysarum nummu-
larifolium L.; H. vaginale L.; Alysicarpus vaginalis DC.] Waste and
cultivated grounds, St. Thomas; St. Jan; St. Croix.
Ecastophyllum Ecastophyllum (L.) Britton. [Hedysarum Ecasto-
phyllum L.; Pterocarpus Ecastophyllum of West; Ecastophyllum
Brownei Pers.; Dalbergia Ecastophyllum Taubert.] Coastal thickets,
oie ehomas; St. Jan; St. Croix.
DREPANOCARPUS LUNATUS (L. f.) Meyer. [Pterocarpus. lunatus
L. f.| Coastal thickets, St. Thomas; St. Croix (according to West
and to Eggers).
ICHTHYOMETHIA PiscipuLa (L.) Hitche. [Erythrina Piscipula L.;
Piscidia Erythrina L.; P. Piscipula Sargent.] Thickets and wood-
lands, ot. Fhomas; St. Jan; St. Croix.
ANDIRA JAMAICENSIS (W. Wright) Urban. [Geoffraea jamaicensis
(inermis) W. Wright; G. tmermis Sw.; Andira inermis H.B.K.;
Vouacapoua americana of Millspaugh.] Woods and along rivulets,
Seeanomas; St. Jan; St. Croix.
Aprus Asrus (L.) W. F. Wight. [Glycine Abrus L.; Abrus
praecatorius L.] Thickets and hedges, St. Thomas; St. Jan; St.
Croix.
CLITORIA TERNATEA L. [Ternatea vulgaris H.B.K.] Thickets
and hedges, St. Thomas; St. Jan; St. Croix.
BRADBURYA VIRGINIANA (L.) Kuntze. ([Clitoria virginiana L.;
Centrosema virginianum Benth.; C. virginianum angustifolium Griseb. ]
Banks, fields and hillsides, St. Thomas; St. Jan; St. Croix.
BRADBURYA PLUMIERI (Turp.) Kuntze. [Clitoria Plumierit Turp.;
Centrosema Plumieri Benth.] Sugar Estate, St. Thomas.
TERAMNUS LABIALIS Spreng. [Z. uncinatus albiflorus Eggers.|
Thickets, St. Thomas; St. Jan; St. Croix.
ERYTHRINA CORALLODENDRON L. Hillsides, St. Thomas; St.
Croix. Planted for shade and ornament.
ERYTHRINA HORRIDA Eggers. Hillside, Flag Hill, St. Thomas.
Recorded from all three islands by Eggers.
56 BROOKLYN BOTANIC GARDEN MEMOIRS
MucuNA PRURIENS (L.) DC. [Dolichos pruriens L.] Shaded
valleys and rocky banks, St. Thomas; St. Croix.
GALACTIA DUBIA DC. [G. tenuztflora of Eggers, partly; G. regularis
of Millspaugh; G. dubia Ehrenbergit Urban; G. filiformis minor +
villosa f. albida Kuntze.] Hillsides and thickets, St. Thomas; St.
Jans) St-oGroix:
GALACTIA STRIATA (Jacq.) Urban. [Glycine striata Jacq.; G.
striata tomentosa Urban; G. filiformis of Eggers; G. tenuiflora of
Millspaugh.] Thickets, St. Thomas; St. Jan; St. Croix.
GALACTIA EGGERsII Urban. [G. tenuiflora of Eggers; G. pendula
of Knox.] Hillside, Flag Hill, St. Thomas; Bordeaux, St. Jan.
Endemic.
CANAVALI RUSIOSPERMA Urban. [C. parviflora of Eggers.] For-
est, Signal Hill, St. Thomas.
CANAVALI ENSIFORMIS (L.) DC. [Dolichos ensiformis L.; C.
gladiata ensiformis of Eggers.] Naturalized in provision grounds,
Signal Hill, St. Thomas (according to Eggers). Cultivated for its
seeds.
CANAVALI LINEATA (Thunb.) DC. [Dolichos lineatus Thunb.;
D. rotundifolius Vahl; Dolichos obtusifolius Lam.; Canavalia obtusi-
foha DC.| Coastal.sands, St. Thomas; St. Jan; St; Crom
CajAN CaJAN (L.) Millsp. [Cytesus Cajan L.; Cajanus flavus DC.;
Cajanus indicus Spreng.| Spontaneous after cultivation, St. Thomas;
St. jane SienCroim.
DOLICHOLUS RETICULATUS (Sw.) Millsp. [Glycine reticulata Sw.;
Rhynchosia reticulata DC.; R. reticulata latifolia Kuntze.]. Roadsides
and thickets, St. Thomas; St. Jan; St. Croix.
DOLICHOLUS PHASEOLOIDES (Sw.) Kuntze. [Glycine phaseoloides
Sw.; Rhynchosia phaseoloides DC.| Forest, Signal Hill, St. Thomas
(according to Eggers).
DOLICHOLUS MINIMUS (L.) Medic. [Dolichos minimus L.; Rhyn-
chosia minima DC.; R. punctata DC.; R. minima lutea Eggers; D.
minimus luteus Millsp.]| Banks, hillsides and thickets and in culti-
vated ground, St. Thomas; St. Jan; St. Croix.
PHASEOLUS LUNATUS L. Thickets, spontaneous after cultivation,
St. Thomas) St alaneist.. Grom
PHASEOLUS VULGARIS L. Spontaneous after cultivation, St.
Thomas; St. Jamsst- Croi,
PHASEOLUS LATHYROIDES L. [P. semierectus L.] Banks, fields
and hillsides, St. Thomas; St. Jan; St. Croix.
PHASEOLUS ALATUS L., recorded from St. Croix by West, is not
further determined.
VIGNA REPENS (L.) Kuntze. ([Dolichos repens L.; ? D. luteus of
BRITTON: FLORA OF THE VIRGIN ISLANDS . 57
West; Dolichos luteolus Jacq.; Vigna luteola Benth.; Bradburya
pubescens of Millspaugh, St. Thomas.] Moist thickets, St. Thomas;
pian ot. Croix.
VIGNA UNGUICULATA (L.) Walp. [Dolichos unguiculatus L.; D.
Catjang L.; Vigna Catjang Walp.] Edge of a cornfield near Doily
Hill, St. Croix; St. Thomas (according to Schlechtendal).
PACHYRRHIZUS EROSUS (L.) Urban. [Dolichos erosus L.; Pachyr-
rhizus angulatus L. C. Rich.] Hillside thickets, St. Thomas.
DouicHos LABLAB L. [Lablab vulgaris Savi; Dolichos benghalensis
Jacq.; Dolichos Lablab abliflorus (DC.) Millsp.] Thickets and spon-
taneous after cultivation, St. Thomas; St. Jan; St. Croix.
DOLICHOS SPHAEROSPERMUS (L.) DC. [Phaseolus sphaerospermus
L.| Cultivated for its seeds.
DOLICHOS SESQUIPEDALIS L. Cultivated for its seeds.
BROWNEA COCCINEA Jacq. Planted on St. Croix (according to West).
PisuM sATIvuM L. Cultivated for its seeds.
OXALIDACEAE
IONOXALIS INTERMEDIA (A. Rich.) Small. [Oxalis intermedia A.
Rich.; O. latifolia of Millspaugh.] Cultivated grounds, St. Croix.
IONOXALIS MARTIANA (Zucc.) Small. [Oxalis Martiana Zucc.]
Shaded banks, St. Thomas; cultivated grounds, St. Croix.
XANTHOXALIS CORNICULATA (L..) Small. [Oxalis corniculata L.;
O. corniculata microphylla of Eggers.] Dry soil, St. Thomas; St. Jan;
St. Croix.
GERANIACEAE
Pelargoniums are cultivated for ornament.
BALSAMINACEAE
IMPATIENS BALSAMINA L. [Balsamina hortensis Desp.] Grown in
flower gardens.
ERYTHROXYLACEAE
“ERYTHROXYLON BREVIPES DC. [E. ovatum of Eggers, of Mill-
spaugh, and of Kuntze; E. areolatum of West.] Hillsides and thickets,
Be, thomas; St. Croix.
ERYTHROXYLON AREOLATUM L. is doubtfully attributed to St.
Thomas by O. E. Schulz.
ZYGOPHYLLACEAE
’ GUAIACUM OFFICINALE L. Woods and thickets, St. Thomas;
formerly on St. Croix and St. Jan. Nearly exterminated. Planted
on St. Thomas.
58 BROOKLYN BOTANIC GARDEN MEMOIRS
TRIBULUS CISTOIDES L. [T. terrester cistoides Oliver.] Dry soil,
St. Crom:
KALLSTROEMIA MAXIMA (L.) T. & G. [Tribulus maximus L.]
Waste and cultivated grounds, St. Thomas; St. Croix; St. Jan.
RUTACEAE
ZANTHOXYLUM PUNCTATUM Vahl. [Fagara trifoliata Sw.; Tobinia
punctata Griseb.| Thickets and banks, St. Croix.
ZANTHOXYLUM THOMASIANUM Krug & Urban. [? Tobinia spinosa
of Eggers.] Forest, Flag Hill, St. Thomas; St. Jan. Endemic.
ZANTHOXYLUM SPINIFEX (Jacq.) DC. [Fagara spinifex Jacq.;
F. tragodes of West; Zanthoxylum microphyllum Desv.] Thickets, St.
Croix.
ZANTHOXYLUM MONOPHYLLUM (Lam.) P. Wilson. [Fagara mono-
phylla Lam.; Zanthoxylum simplicifolium Vahl; Z. Ochroxylum DC.|]
Hillsides, woods and thickets, St. Thomas; St. Jan; St. Croix.
ZANTHOXYLUM MARTINICENSE (Lam.) DC. [Fagara martinicensis
Lam.; Zanthoxylum Clava-Herculis of Eggers.] Woods and hillsides,
St. Thomas: St..jans? St./Croi-
ZANTHOXYLUM FLAVUM Vahl. [Fagara flava Krug & Urban.]
Bordeaux Hills, St. Jan, nearly extinct (according to Eggers). Not
found by us on St. Jan in 1913.
PILOCARPUS RACEMOSUS Vahl. Forest, King’s Hill, St. Jan.
AMYRIS ELEMIFERA L. [A. maritima Jacq.; 4. sylvatica of Eggers. ]
Woods and thickets, St. Thomas; St. Jan; St. Croix (according to
Eggers).
CHALCAS EXOTICA (L.) Millsp. [Murraya exotica L.] Spon-
taneous after cultivation, St. Thomas; St. Croix.
TRIPHASIA TRIFOLIA (Burm. f.) P. Wilson. [Limonia trifolia
Burm. f.; 7. ¢trifoliata (L.) DC.] Spontaneous after cultivation,
naturalized in thickets, St. Thomas; St. Jan; St. Croix.
Cirrus Mepica L. Recorded by Eggers as naturalized in gardens.
Cirrus LIMA Lunan. [C. medica Limonum of Eggers; C. Limetta
Wight.] Woodlands and thickets, naturalized, St. Thomas; St. Jan;
St. Croix.
Cirrus AURANTIUM L. Occasionally spontaneous after planting,
St. Thomas: St. Croix,
CITRUS \VULGARIS Risso. [C. Bigaradia Loisel.; C. Aurantium
bigaradia Griseb.] Occasionally spontaneous after planting, St.
Thomas? St. Crom:
CITRUS DECUMANA L. Planted for its fruit.
CITRUS BUXIFOLIA Poir. Planted for its fruit (according to
Eggers).
BRITTON: FLORA OF THE VIRGIN ISLANDS 59
CLAUSENA WAmpI Blanco. [Cookia punctata Sonn.| Planted for
shade.
SURIANACEAE
SURIANA MARITIMA L. Coastal sands, St. Thomas; St. Jan; St.
Croix.
SIMAROUBACEAE
QUASSIA AMARA L. Naturalized in and about gardens, St. Thomas;
St. Croix. Planted for shade.
CASTELARIA NICHOLSONI (Hook.) Small. [Castela Nicholsoni
Hook.; C. erecta of Eggers and of Millspaugh.] Thickets, St. Croix.
AESCHRION ANTILLANA (Eggers) Small. [Rhus antillana Eggers;
Quassia excelsa of West; Picrasma antillana Urban; Picraena excelsa
of Eggers and of Millspaugh.] Forests, St. Thomas; St. Jan; St.
Croix.
BURSERACEAE
ELAPHRIUM SIMARUBA (L.) Rose. [Pistacia Simaruba L.; Bursera
gummifera L.; Bursera Simaruba Sargent.] Woods and hills, St.
Thomas; St. Jan; St. Croix.
TETRAGASTRIS BALSAMIFERA (Sw.) Kuntze. [Hedwigia balsamtfera
Sw.; ? Icica altissima of West.] St. Croix (according to West).
MELIACEAE
SWIETENIA MAHAGONI Jacq. Hillsides and valleys, St. Thomas;
St. Croix. Often planted; perhaps not native.
Metia AZEDARACH L. Roadsides; occasional in woods, St.
moomas; St. Jan; St. Croix.
TRICHILIA HIRTA L. [T. spondioides Jacq.] Woods, thickets and
hillsides, St. Thomas; St. Jan; St. Croix.
_TRICHILIA WAWRANA ANTILLANA C. DC., described as from St.
Croix from a specimen in the Copenhagen herbarium, is otherwise
unknown.
GUAREA TRICHILIOIDES L., was recorded by West from St. Croix,
but the record was questioned by Eggers. It is abundant in Porto
Rico.
MALPIGHIACEAE
HIRAEA FAGINEA (Sw.) Ndz. [H. faginea glandulifera Ndz.],
recorded by Niedenzu from St. Thomas, is probably an error in
locality.
- BANISTERIA PURPUREA L. [Heteropteris purpurea H.B.K.; H.
parvifolia DC.] Thickets and hillsides, St. Thomas; St. Jan; St.
Croix.
60 BROOKLYN BOTANIC GARDEN MEMOIRS
BANISTERIA LAURIFOLIA L. [Heteropteris laurifolia A. Juss.; B.
laurifolia antillana Ndz., B. lancifolia of West.] St. Croix (according
to Niedenzu). .
STIGMAPHYLLON LINGULATUM (Poir.) Small. [Triopteris lingulata
Poir.; Banisteria periplocifolia Desf.; Stigmaphyllon periplocifolium
A. Juss.; S. Sagraeanum of Millspaugh.] Thickets and hillsides, St.
Thomas; St. Jan; St. Croix.
STIGMAPHYLLON CORDIFOLIUM Ndz. St. Thomas (according to
Niedenzu).
STIGMAPHYLLON CILIATUM (Lam.) A. Juss., recorded by Niedenzu
from St. Thomas, as collected by Finlay, was really from Trinidad.
STIGMAPHYLLON TOMENTOSUM (Desf.) Ndz. [Banisteria tomentosa
Desf.] Royiers, St. Jan (according to Niedenzu).
SPACHEA LITTORALIS A. Juss., recorded by A. Jussieu as collected
by Finlay on St. Thomas, was from Trinidad.
THRYALLIS GLAUCA (Cav.) Kuntze. [Galphimia glauca Cav.; G.
gracilis Bartl.}| Roadsides and about dwellings, naturalized, St.
Thomas; St. Croix.
TETRAPTERIS INAEQUALIS Cav. St. Croix (according to Niedenzu).
MALPIGHIA FUCATA Ker. [M. fucata elliptica Ndz.] St. Croix
(according to Eggers).
MALPIGHIA GLABRA L. [M. glabra antillana Urban & Ndz.]
Thickets, St. Thomas; St. Croix (according to Eggers and to Nieden-
Zu).
MALPIGHIA PUNICIFOLIA L. [M. punicifolia vulgaris and lancifolia
Ndz.; M. glabra of Millspaugh.] Hillsides and thickets, St. Thomas;
Sty Croix.
MALPIGHIA LINEARIS Jacq. [M. angustifolia L.; M. angustifolia
oblongata Ndz.; ? M. urens lanceolata Eggers.] Hillside thickets,
Water Island, St. Thomas; St. Jan.
MALPIGHIA BIFLORA Poir. [M. oaycocca Grisebachiana Ndz.]
St. Croix (according to Niedenzu).
MALPIGHIA PALLENS Small. [M. urens of Millspaugh? and of
Eggers.] Thickets along sandy beaches, St. Croix. Endemic.
MALPIGHIA INFESTISSIMA (A. Juss.) Rich. [M. urens of West;
M. urens infestissima A. Juss.; M. Cnide of Eggers.] Hillside thickets,
Water Island, St. Thomas; St. Jan. Also on Vieques and Culebra.
Endemic. St. Thomas is the type locality.
BUNCHOSIA GLANDULOSA (Cav.) DC. [Malpighia glandulosa Cav.;
M. Swartziana of Eggers.] Thickets, St. Thomas; St. Jan; St. Croix.
BYRSONIMA SPICATA (Cav.) DC. [B. coriacea of Millspaugh.]
Woods and thickets, St. Thomas; St. Jan; St. Croix.
BYRSONIMA CUNEATA (Turcz.) P. Wilson. [B. lucida DC.] St.
Thomas (according to de Candolle, and cited also by Niedenzu).
BRITTON: FLORA OF THE VIRGIN ISLANDS 61
BYRSONIMA MARTINICENSIS Krug & Urban. St. Croix (according
to Small).
POLYGAEACEAE
POLYGALA ANGUSTIFOLIA H.B.K. Thickets, southern side of St.
Thomas.
SECURIDACA BROWNEI Griseb. [S. scandens West.] Naturalized
around Christiansted, St. Croix, and on St. Thomas (according to
Eggers).
SECURIDACA ERECTA L. Dry soil, St. Croix; St. Thomas (accord-
ing to de Candolle).
EUPHORBIACEAE
SAVIA SESSILIFLORA (Sw.) Willd. [Croton sessiliflorum Sw.]
Thickets and hillsides, St. Thomas; St. Jan; St. Croix.
Asterandra grandifolia (L.) Britton. [Phyllanthus grandifolius L.]
St. Thomas (according to Urban).
PHYLLANTHUS NIRuURI L. Waste and cultivated grounds, St.
Thomas: St. Croix:
PHYLLANTHUS ACUMINATUS Vahl is accredited to St. Thomas by
Mueller (DC. Prodr. 15?: 381) who records a specimen in the Candol-
lean herbarium, but the shrub is not known to inhabit St. Thomas now.
Cicca pDisTicHA L. [Phyllanthus distichus Muell. Arg.] Spon-
taneous after planting, St. Thomas; St. Jan; St. Croix.
MARGARITARIA NOBILIS L. f. [Phyllanthus nobilis Muell. Arg.;
Cicca antillana A. Juss.; P. nobilis antillanus Muell. Arg.] Forests,
St. Thomas; St. Jan; St. Croix (according to Eggers).
SECURINEGA ACIDOTHAMNUS (Griseb.) Muell. Arg. [? Adelia
Acidoton of West; Flueggea Acidothamnus Griseb.] Thickets, St.
Thomas; Little St. James Island, St. Jan; eastern St. Croix (according
to Eggers).
DRYPETES GLAUCA Vahl. St. Croix (according to Eggers).
CROTON ASTROITES Dryand. [C. phlomoides Pers.| Thickets, St.
Whomas; St. Jan; St. Croix.
CROTON BETULINUS Vahl. Thickets, St: Thomas; St. Jan; St.
Croix.
CROTON FLAVENS L. [C. balsamifer Jacq.; C. flavens rigidus
Muell. Arg.; Oxydestes flavens Kuntze.] Thickets, St. Thomas; St.
fam; St.-Croix.
CROTON DISCOLOR Willd. Rocky thickets, St. Thomas; St. Croix.
CROTON LoBATUS L. [Oxydectes lobata Kuntze.] Waste and
cultivated grounds, St. Thomas; St. Croix; St. Jan.
CROTON HUMILIS L. Hillside thickets, St. Thomas.
CROTON GLANDULOSUS L. St. Croix (according to Urban).
62 BROOKLYN BOTANIC GARDEN MEMOIRS
CROTON OVALIFOLIUS Vahl. [Oxydectes ovalifolia Kuntze.] Hill-
sidés, St. Thomas; St. Jan; St-.Croi.
CROTON HASTATUS West, of St. Croix (hyponym) is not identified.
An arboreus Croton, not found in flower, occurred on Flag Hill,
St. Thomas, according to Eggers.
DITAXIS FASCICULATA Vahl. [Argyrothamnia fasciculata Muell.
Are;| Thickets, St. Thomas; St. Jan; St--Croix.
ARGYTHAMNIA CANDICANS Sw. Thickets and hillsides, St. Thomas;
St, Jan; st. Crom.
RICINELLA RICINELLA (L.) Britton. [Adelia Ricinella L.; R.
pedunculosa Muell. Arg.] Hillsides and thickets, St. Thomas; St.
Janis: St. Grom:
ACALYPHA PORTORICENSIS Muell. Arg. Rocky slopes, St. Croix.
ACALYPHA CHAMAEDRIFOLIA (Lam.) Muell. Arg. [Cvoton chamae-
drifolius Lam.; Acalypha reptans Sw.; A. corchorifolia Willd.; A.
chamaedrifolia genuina and brevipes of Eggers.] Rocky soil, St. Thom-
asc. St. Croix.
ACALYPHA POLYSTACHYA Jacq. St. Thomas (according to Eggers).
The record is probably an error in determination.
TRAGIA VOLUBILIS L. Thickets, banks and hillsides, St. Thomas;
St.gan; Sts Croix:
DALECHAMPIA SCANDENS L. Thickets, St. Thomas; St. Jan; St.
Crom
RICcINUS COMMUNIS L. Waste and cultivated grounds, St. Thomas;
St-Jang St. -Croi:
ALEURITES MOLUCCANA (L.) Willd. [Jatropha moluccana L.|
Roadsides and near dwellings, St. Thomas; St. Croix.
JatTropHa Curcas L. Hillsides and near dwellings, St. Thomas;
Sit Janke ot. Croix
JATROPHA GOSSYPIFOLIA L. [Adenoropium gossypifolium Pohl;
J. gossypifolia staphisagriaefolia and elegans of Eggers.| In dry soil,
fields and hillsides, St. Thomas; St. Jan; St. Croix.
JATROPHA MULTIFIDA L. Roadsides and planted in gardens, St.
Thomas; St... Croix.
JATROPHA PANDURAEFOLIA Andr. Planted for ornament.
Maninot Maninot (L.) Cockerell. [Jatropha Manthot L.| Spon-
taneous or persistent after cultivation, St. Thomas; St. Jan; St.
Croix.
Saptum LaurocerAsus Desf. [Excoecaria Laurocerasus Muell.
Arg.; ? E. Laurocerasus laurifcha of Eggers.] A high tree in forests,
Cinnamon Bay, St. Jan, not seen flowering (according to Eggers).
Otherwise known only from Porto Rico.
HIPPOMANE MANCINELLA L. Coastal woods, St. Thomas; St.
Croix:
BRITTON: FLORA OF THE VIRGIN ISLANDS 63
GYMNANTHES LUCIDA Sw. [Sebastiana lucida Muell. Arg.] Woods
ane tuickets, St.Thomas; St. Jan; St. Croix.
HuRA CREPITANS L. Woods, roadsides and near dwellings, St.
Miomas; St: Jan; St. Croix.
CHAMAESYCE VAHLII (Willd.) P. Wilson. [Euphorbia Vahlit
Willd.] Rocky hills, Little St. James Island, St. Jan.
CHAMAESYCE BUXIFOLIA (Lam.) Small. [Euphorbia buxifolia
Lam.; E£. glabrata Sw.] Coastal sands, St. Thomas; St. Jan; St.
Croix.
CHAMAESYCE ARTICULATA (Aubl.) Britton. [Euphorbia articulata
Aubl.; E. linearis Retz.; E. linearis heterophylla Kuntze.] Coastal
rocks, St. Thomas; St. Jan; St. Croix (according to Retzius and
reported by Eggers).
CHAMAESYCE HIRTA (L.) Millsp. [Euphorbia hirta L.; E. piluli-
fera L.; E. pilulifera procumbens Boiss.] Roadsides, banks and culti-
vated grounds, St. Thomas; St. Jan; St. Croix.
CHAMAESYCE HYPERICIFOLIA (L.) Millsp. [Euphorbia hypericifolia
L.; E. hypericifolia hyssopifolia of Eggers.] Fields, banks and culti-
vated grounds, St. Thomas; St. Jan; St. Croix.
CHAMAESYCE SERPENS (H.B.K.) Small. [Euphorbia serpens
H.B.K.] Dry soil, St. Thomas.
CHAMAESYCE PROSTRATA (Ait.) Small. [Euphorbia prostrata Ait.;
? E. Chamaesyce of West.] Waste and cultivated grounds, St. Thomas;
er lan; ot. Croix.
CHAMAESYCE BRASILIENSIS (Lam.) Small. [Euphorbia brasiliensis
Lam.] Grassy places near Charlotte Amalia, St. Thomas, determined
by Millspaugh.
EUPHORBIA THYMIFOLIA Burm. is recorded by Eggers from all the
islands, but has not been found on any of them by other collectors,
and his determination of the species is therefore doubted.
AKLEMA PETIOLARIS (Sims.) Millsp. [Euphorbia petiolaris Sims.;
? E. cotinifolia of West and of Schlechtendal.] Hillsides and thickets,
St. Thomas; St. Jan; doubtfully recorded from St. Croix.
POINSETTIA HETEROPHYLLA (L.) Kl. & Garcke. [Euphorbia hetero-
phylla L.; E. heterophylla linifolia Kuntze.| Dry rocky situations,
St. Thomas.
POINSETTIA CYATHOPHORA (Murr.) S. Brown. [Euphorbia cy-
athophora Murr.; E. heterophylla cyathophora Griseb.]| Waste and
cultivated grounds, St. Thomas; St. Jan; St. Croix.
POINSETTIA OERSTEDIANA KI. & Garcke. [Euphorbia geniculata
of Eggers; Euphorbia Oerstediana Boiss.] Grassy places, St. Thomas;
Si. Croix.
POINSETTIA PULCHERRIMA (Willd.) Graham. [Euphorbia pulcher-
rvima Willd.| Planted for ornament.
64 BROOKLYN BOTANIC GARDEN MEMOIRS
EUPHORBIA NERIIFOLIA L. Planted for ornament.
EUPHORBIA SPLENDENS Bojer. Planted for ornament.
EUPHORBIA ANTIQUORUM L. Cultivated (according to Eggers).
PEDILANTHUS TITHYMALOIDES (L.) Poit. [Euphorbia tithymaloides
‘L.] Persistent after cultivation, St. Thomas. Grown in flower
gardens.
PEDILANTHUS PADIFOLIUS (L.) Poit. [Euphorbia tithymaloides
padifolia L.| Thickets in dry stony ground, St. Croix.
PEDILANTHUS ANGUSTIFOLIUS Poit. Thickets and hillsides, St.
Thomas; St. Jan.
CODIAEUM VARIEGATUM Blume. Planted for ornament.
BUXACEAE
TRICERA VAHLII (Baill.) Britton. [Buxus Vahliu Baill.; Tricera
laevigata sanctae-crucis Eggers.] On limestone, Stony Ground, St.
Croix. Known otherwise only on Porto Rico.
MANGIFERA INDICA L. Spontaneous after planting, St. Thomas;
St. Jans St. Croix:
ANACARDIUM OCCIDENTALE L. Woods, hillsides and along roads,
St-sinomas: ot lam: ob. Croix.
SPONDIAS PURPUREA L. Spontaneous after planting, St. Thomas;
St. Jans St. Crom
SPONDIAS MomBIN L. [Spondias lutea L.] Woods, hills and road-
sides: ot. Lhomas; St Jan; St. Crom.
SPONDIAS DULCIS Forst. f. Cultivated on St. Croix.
Comoc.LapIA DoponakEa (L.) Urban. [lex Dodonaea L.; Como-
cladia tlicifolia Sw.| Rocky coastal thickets, St. Thomas; St. Jan;
St. Crow
CELASTRACEAE
MAYTENUS ELLIPTICA (Lam.) Krug & Urban. [Senacia elliptica
Lam.; Rhamnus laevigatus Vahl; Ceanothus laevigatus DC.] Woods
and thickets, St. Thomas; St. Jan; St. Croix.
MAYTENUS CYMOSA Krug & Urban. [M. elaeodendroides of
Eggers.] Thickets, St. Thomas; St. Croix. Known otherwise only
from Vieques. Endemic.
RHACOMA CROSSOPETALON L. [Myginda pallens Sw.; M. latifolia
Vahl, not Sw.] Thickets, St. Thomas; St. Jan; St. Croix.
MYGINDA LATIFOLIA (Sw.) Urban. [Myginda latifolia Sw.]
Thickets, St. Thomas.
SCHAEFFERKA FRUTESENS Jacq. [S. completa Sw.| Thickets, St.
Thomas; St. Jan; St. Croix.
ELAEODENDRON XYLOCARPUM (Vahl) Urban. [Cassine xylocarpa
BRITTON: FLORA OF THE VIRGIN ISLANDS 65
Vent.; Celastrus polygamus Vahl; Rhamnus polygamus West.| Coast-
alihickets, St. Thomas; St: Jan; St. Croix.
HIPPOCRATEACEAE
HIPPOCRATEA VOLUBILIS L. is doubtfully accredited to St. Thomas
by Urban (Symb. Ant. 4: 367).
SAPINDACEAE
SERJANIA POLYPHYLLA (L.) Schum. [Paullinia polyphylla L.;
S. lucida Schum.; Paullinia curassavica of West.] Woods and
pmekets, St. Thomas; St. Jan; St. Croix.
PAULLINIA PINNATA L. St. Thomas (according to Radlkofer).
Dr. Millspaugh records Paullinia frutescens glabrescens (L.) Radlk.
from Midland, St. Croix, as perhaps cultivated.
CARDIOSPERMUM HaLicAcABuM L. Banks and thickets, spon-
taneous after cultivation, St. Thomas; St. Croix.
CARDIOSPERMUM MICROCARPUM H.B.K. Thickets, St. Thomas;
sienpar- St. Croix.
CARDIOSPERMUM CORINDUM L. Hillsides, St. Croix.
CARDIOSPERMUM BIPINNATUM West, is not known to modern
botanists.
ALLOPHYLUS OCCIDENTALIS (Sw.) Radlk. [Schmeidelia occidentalis
Sw.] Forests, St. Croix.
SAPINDUS SAPONARIA L._ [.S. inaequalis DC.] Forests, St. Thom-
fee oesjan: St. Croix.
ME LIcocca BIyUGA L. Hillsides, woods and along roads, St.
Thomas; St. Jan; St. Croix. Planted and naturalized.
CUPANIA TRIQUETRA A. Rich. [C. fulva of Eggers.] Woods and
hills, St. Thomas; St. Jan.
BLIGHIA SAPIDA Koen. Planted for its fruit.
DODONAEACEAE
DopoONAEA viscosa L. Coastal thickets, St. Thomas; St. Croix.
RHAMNACEAE
KRUGIODENDRON FERREUM (Vahl) Urban. [Rhamnus ferreus
Vahl; Ceanothus ferreus DC.; Condalia ferrea Griseb.] Woods and
thickets, St. Thomas; St. Jan; St. Croix.
REYNOSIA UNCINATA Urban. [R. mucronata of Eggers.] Coastal
thickets near Tague Bay, St. Croix.
ReynostA GuAMA Urban. [R. latifolia of Eggers.] Hillside
thickets, St. Thomas; St. Jan. Endemic.
6
66 BROOKLYN BOTANIC GARDEN MEMOIRS
SARCOMPHALUS RETICULATUS (Vahl) Urban. [Paliurus reticulatus
Vahl; Zizyphus reticulatus Vahl.]| Thickets, Fair Plain, St. Croix.
COLUBRINA COLUBRINA (Jacq.) Millsp. [Rhamnus Colubrina
Jacq.; Colubrina ferruginosa Brongn.] Coastal thickets and hillsides,
St: Daomas: St. Jan -St. Crom.
COLUBRINA RECLINATA (L’Her.) Brongn. [Rhamnus_ reclinatus
L’Her.; R. ellipticus Sw.| Woods and thickets, St. Thomas; St. Jan;
St) Croix.
GOUANIA LUPULOIDES (L.) Urban. [Banisteria lupuloides L.;
Gouania domingensis L.| Woods and thickets, St. Thomas; St. Jan;
St. Cror,
ZIZYPHUS JUjUBA (L.) Lam. [Rhamnus Jujuba L.| Planted for
its fruit.
RHAMNUS GLABRATUS West. A species not understood by modern
botanists.
VITACEAE
VITIS TILIIFOLIA H.& B. [V.caribaea DC.] Forests, St. Thomas.
VITIS VINIFERA L. Planted for its fruit.
Cissus sicyorbEs L. Woods, walls and thickets, St. Thomas;
Stadia: St. Crom
CISSUS TRIFOLIATA L. [C. acida L.] Dry thickets, St. Thomas;
Stan: St. Crom:
Cissus cAusTIcA Tuss. [C. trifoliata of Eggers and of Millspaugh.]
On trees and rocks, St. Thomas; St. Croix.
Cissus OBOVATA Vahl. St. Croix. Known otherwise from St.
Martin and eastern Porto Rico.
TILIACEAE
CoRCHORUS ACUTANGULUS L. Waste and cultivated grounds, St.
Thomas; St. Croix.
CoORCHORUS SILIQUOSUS L. Thickets, fields, waste and cultivated
grounds, St. Thomas; St. Jan; St. Croix.
CorcHorus HIRTUS L. Gardens and roadsides, St. Thomas and
St. Croix (according to Eggers).
CoRCHORUS HIRSUTUS L. Coastal thickets and hillsides, St.
Thomas; St-vjan.) St.Croix.
TRIUMFETTA EXCISA Urban. Bassin yard, St. Croix. Known
otherwise only from Porto Rico.
TRIUMFETTA RHOMBOIDEA Jacq. Thickets, St. Croix.
TRIUMFETTA SEMITRILOBA Jacq. [T. althaeoides Lam.; T. sem-
triloba havanensis Millsp.| Woods, banks and thickets, St. Thomas;
St. Jan; St:/Grorm:
BRITTON: FLORA OF ‘THE VIRGIN ISLANDS 67
TRIUMFETTA LAPPULA L. Thickets, St. Thomas; St. Jan; St.
Croix.
SLOANEA DENTATA L. Planted on St. Croix (according to West).
MALVACEAE
ABUTILON UMBELLATUM (L.) Sweet. [Sida uwmbellata L.] Rocky
thickets and hillsides, St. Thomas; St. Jan; St. Croix.
ABUTILON HIRTUM (Lam.) Sweet. [Sida hirta Lam.; Abutilon
indicum hirtum Griseb.; A. graveolens of Millspaugh.] Waste and
cultivated grounds, St. Thomas; St. Croix.
ABUTILON INDICUM (L.) Sweet. [Sida indica L.; A. subpapy-
raceum Hochreutiner.| Sandy waste grounds, St. Thomas; St. Croix.
ABUTILON LIGNOSUM A. Rich. St. Thomas and St. Croix (accord-
ing to Eggers).
GAYOIDES CRISPUM (L.) Small. [Sida crispa L.; Abutilon crispum
Medic.] Sandy soil, St. Thomas.
WISSADULA AMPLISSIMA (L.) R. E. Fries. [Sida amplissima L.;
? Abutilon periplocifolium albicans of Eggers; Sida hernandioides
L’Her.; W. hernandioides Garcke.] Banks and thickets, St. Jan;
Se Croix.
WISSADULA PERIPLOCIFOLIA (L.) Griseb. [Sida periplocifolia L.;
Abutilon periplocifolium Don.| Fields and hillsides, St. Croix.
MALVASTRUM COROMANDELIANUM (L.) Garcke. [Malva coro-
mandeliana L.; M. americana L.; M. tricuspidata Ait.; Malvastrum
tricuspidatum A. Gray.] Waste and cultivated grounds, St. Thomas;
eas ot. Croix.
MALVASTRUM SPICATUM (L.) A. Gray. [Malva spicata L.] Hill-
sides, waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
SIDA CILIARIS L. Dry, grassy and rocky situations, St. Thomas;
Sivan; ot. Croix.
SIDA ERECTA Macf. Dry soil, St. Croix.
SIDA sPINOSA L. [S. angustifolia Lam.; (?) S. spinosa polycarpa
Eggers; S. retusa of Millspaugh.]| Banks, fields and cultivated
Grounds, ot. Thomas; St. Jan; St. Croix.
SIDA GLOMERATA Cay. Banks and thickets, St. Thomas; St. Jan.
SIDA CARPINIFOLIA L.f. [S. carpinifolia acuta Millsp.; S. carpini-
folia antillana Millsp.; (2) S. carpinifolia brevicuspidata Eggers.]
Banks, fields, woods and thickets, St. Thomas; St. Jan; St. Croix.
SIDA RHOMBIFOLIA L. [S. rhombifolia retusa of Eggers.| Banks,
fields, waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
SIDA PROCUMBENS Sw. [S. pilosa Cav.; S. supina Sw.] Road-
sides, St. Croix.
SIDA CORDIFOLIA L. [S. althaetfolia Sw.; S. cordifolia althaeifolia
6
68 BROOKLYN BOTANIC’ GARDEN MEMOIRS
of Millspaugh.] Banks, fields and thickets, St. Thomas; St. Jan;
St. (Crom,
SIDA HUMILIS Cav. [Sida supina of Muillspaugh, St. Thomas;
Sida supina glabra of Millspaugh and of Eggers.| Banks, fields, and
thickets, St. Thomas; St. Jan; St. Croix.
SIDA GLABRA Mill. [S. ulmifolia Cav.; S. arguia Sw.|] Banks,
fields and thickets, St. Thomas; St. Croix.
SIDA GLUTINOSA Commers. [.S. nervosa DC.; (?) S. nervosa viscosa
Eggers.| Hillsides and thickets, St. Thomas; St. Jan; St. Croix.
SIDA ACUMINATA DC. [S. acuminata macrophylla Schl. and
microphylla Schl.| Hillsides, St. Thomas; St. Croix. ;
SipA Eccrersi E. G. Baker. St. Thomas, apparently (Eggers,
Suppl. 14). Otherwise known only from Tortola and Culebra.
Endemic.
SIDA JAMAICENSIS L. [S. tristis Schl.] Fields and hillsides, St.
Thomas St. Jan: St. Croix:
BASTARDIA viscosa (L.) H.B.K. [Sida viscosa L.] Dry fields,
hills and thickets, St. Thomas; St. Jan; St. Croix.
MALACHRA CAPITATA L. [M. palmata Moench.] Dry soil, St.
Croix.
MALACHRA ALCEIFOLIA Jacq. [M. rotundifolia Schrank.| Thick-
ets, waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
MALCHRA FASCIATA Jacq. [M. radiata Griseb., not L.; (?) M.
urens of Eggers.] Waste grounds, St. Thomas.
URENA LOBATA L. [U. americana L. f.; U. reticulata Cav.; U.
lobata americana Guerke.] Fields, woods, hillsides and cultivated
grounds, ot. lhomas; pt. Jan; St. Croix:
URENA SINUATA L. St. Thomas (according to Guerke).
PAVONIA SPINIFEX (L.) Cav. [Hibiscus spinifex L.] Thickets,
waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
MALACHE SCABRA B. Vogel. [Pavonia spicata Cav.; Althaea
racemosa Sw.; P. racemosa Sw.| Mangrove swamps, St. Croix (ac-
cording to Eggers).
PARITI TILIACEUM (L.) A. Juss. [Hibiscus tiliaceus L.] Coastal
woods, St. Thomas; St. Jan; St. Croix (according to West).
HIBISCUS BRASILIENSIS L. [H. phoeniceus Jacq.| Hillsides and
thickets, St. Thomas; St. Croix.
HIBISCUS CLYPEATUS L. St. Croix (according to West).
Hipiscus vitirotius L. Thickets and waste grounds, St. Thomas;
Sti Croix:
Hipiscus SABDARIFFA L. Spontaneous after cultivation, St.
Thomas; St. Croix.
Hipiscus RosA-sINENsIS L. Spontaneous after cultivation, St.
Thomas. Planted for ornament.
BRITTON: FLORA OF THE VIRGIN ISLANDS 69
HIBISCUS MUTABILIS L. Planted for ornament.
CIENFUEGOSIA HETEROPHYLLA (Vent) Garcke. [Fugosia hetero-
phylla; Kosteletzkya pentasperma of Eggers.] Moist soil, St. Thomas.
ABELMOSCHUS BSCULENTUS (L.) Moench. [Hibiscus esculentus L.]
Spontaneous after cultivation, St. Thomas; St. Jan; St. Croix.
THESPESIA POPULNEA (L.) Soland. [Hibiscus populneus L.|
Coastal woods and thickets, St. Thomas; St. Jan; St. Croix. Com-
monly planted.
GOSSYPIUM BARBADENSE L. Thickets and hillsides, spontaneous
after cultivation, St. Thomas; St. Croix.
GOSSYPIUM VITIFOLIUM Lam. is recorded by Schlechtendal as
naturalized in St. Thomas; Eggers suggests it may formerly have
been cultivated there.
ALTHAEA ROSEA Cav. Planted for ornament.
BOMBACACEAE
CEIBA PENTANDRA (L.) Gaertn. [Bombax pentandrum L.; B.
heptaphyllum of West; Eriodendron anfractuosum DC.| Hills, forests
and roadsides, St. Thomas; St. Jan; St. Croix.
ADANSONIA DIGITATA L. Naturalized in wooded valleys, St. Croix
(according to Eggers). Planted for shade on St. Thomas and St.
Croix.
QUARARIBAEA TURBINATA (Sw.) Poir. [Myrodia turbinata Sw.|]
Woods, St. Jan; Spring Garden, St. Croix (according to West).
PACHIRA ALBA Walp. Planted, St. Thomas.
PACHIRA AQUATICA Aubl. [Carolinea princeps L.f.] Planted, St.
Croix.
STERCULIACEAE
MELOCHIA NODIFLORA Sw. [Riedleia nodiflora DC.| Hillsides,
banks and thickets, St. Thomas; St. Jan; St. Croix.
Moluchia pyramidata (L.) Britton. [Melochia pyramidata L.|
Grassy places, waste and cultivated grounds, St. Thomas (according
to West); St. Croix.
Moluchia tomentosa (L.) Britton. [Melochia tomentosa L.| Hill-
sides and thickets, St. Thomas; St. Jan; St. Croix.
WALTHERIA AMERICANA L. [W. indica L.] Fields, banks and
hillsides, St. Thomas; St. Jan; St. Croix.
AYENIA PUSILLA L. Thickets and hillsides, St. Thomas; St. Jan;
Set. Croix.
THEOBROMA Cacao L. Naturalized in shaded valleys, St. Croix
(according to Eggers). Planted for its seeds.
GuAzuMA GuAzuMA (L.) Cockerell. [Theobroma Guazuma L.;
70 BROOKLYN BOTANIC GARDEN MEMOIRS
Guazuma ulmifolia Lam.; G. tomentosa H.B.K.] Fields, woods and
roadsides, St. Thomas; St. Croix.
HELICTERES JAMAICENSIS Jacq. Thickets, St. Thomas; St. Jan;
Ste Croix.
DILLENIACEAE
DAVILLA RUGOSA Poir. is recorded by Grisebach from the island
St. Thomas, and also from St. Thomas-in-the-Vale, Jamaica. It oc-
curs in the Jamaica parish, but is not known on our island.
OCHNACEAE
OURATEA LITTORALIS Urban. [Gomphia nitida of Eggers.] Coast-
al thickets, St. Thomas. Known otherwise only from Porto Rico.
TERNSTROEMIACEAE
Taonabo peduncularis (DC.) Britton. [Ternstroemia peduncularis
DC.; T. elliptica of West and of Eggers.] Forests, Bordeaux Hill, St.
Jan; Maroon Hill, St. Croix.
CLUSIACEAE
MAMMEA AMERICANA L. Forests, hills and roadsides, St. Thomas;
St= Jan; St. Croix:. Much planted.
CALOPHYLLUM CALABA Jacq. Forests, roadsides and valleys, St.
‘Thomas: St, Croix.
CLUSIA ROSEA Jacq. [?C. alba of West.] Hillsides and forests,
St. Thomas; St. Jan; St. Croix (according to West and to Eggers).
TAMARICACEAE
TAMARIX INDICA Willd. Planted for ornament.
BIXACEAE
BixA ORELLANA L. Spontaneous after planting, St. Thomas; St.
Croix. Grown for the dye stuff annato.
CANELLACEAE
CANELLA WINTERANA (L.) Gaertn. [Laurus Winterana L.;
Canella alba Murr.] Woods and thickets, St. Thomas; St. Jan; St.
Croix (according to Eggers).
VIOLACEAE
Calceolaria linearifolia (Vahl) Britton. [Viola linearifolia Vahl;
Hybanthus linearifolius Urban; Ionidium strictum Vent.] Rocky
thickets, Water Island, St. Thomas; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 71
FLACOURTIACEAE
Procxia Crucis L. [Trilix crucis Griseb.] Forests, St. Thomas;
St. Jan (according to Eggers); St. Croix.
MyYROXYLON BUXIFOLIUM (A. Gray) Krug & Urban. [Xylosma
buxifolium A. Gray; Drypetes laevigata of Eggers.] Cinnamon Gut,
St. Jan; St. Croix (according to Urban).
MYROXYLON NITIDUM (Hell.) Kuntze [Xylosma nitidum A. Gray],
is recorded ‘by Eggers as naturalized on St. Thomas. It is endemic
in Jamaica.
SAMYDA SPINULOSA Vent. [S. glabrata Grisebach and of Eggers,
not Sw.] Thickets, Crown, St. Thomas (according to Eggers).
Otherwise known only from Porto Rico.
SAMYDA DODECANDRA Jacq. [S. serrulata L.| Thickets, St.
@homas; St. Jan; St. Croix.
CASEARIA GUIANENSIS (Aubl.) Urban. [Iroucana guianensis
Aubl.; Casearia ramiflora Vahl; C. hirta of Millspaugh; C. nitida of
Kuntze.] Woods and thickets, St. Thomas; St. Croix; St. Jan.
CASEARIA DECANDRA Jacq. [C. parvifolia Willd.; Samyda de-
candra Jacq.; C. parvifolia microcarpa Eggers.| Woods and thickets,
St. Thomas; St. Jan; St. Croix (according to Eggers).
CASEARIA ARBOREA (L. C. Rich) Urban. [Samyda arborea L. C.
Rich; C. stipularis Vent.| St. Thomas, collected by O. Kuntze, whose
specimen is so labelled by him; Urban (Symb. Ant. 7: 75) indicates,
however, that it may have come from Porto Rico.
CASEARIA SYLVESTRIS Sw. [Samyda parviflora L. not Loefl.]
Woods, hills and thickets, St. Thomas; St. Jan; St. Croix.
TURNERACEAE
TURNERA DIFFUSA Willd. [J7. microphylla Desv.; T. parviflora
of Eggers.] Coastal thickets, St. Thomas; St. Jan; St. Croix.
TURNERA ULMIFOLIA L. [T7. ulmifolia acuta Urban; T. angusti-
folia Mill.| Hillsides and waste grounds, St. Thomas; St. Jan; St.
Croix.
PIRIQUETA VISCOSA Griseb. Hillside thickets, St. Thomas.
PASSIFLORACEAE
PASSIFLORA PALLIDA L. [P. suberosa L.; P. hirsuta L.; P.
minima L.; P. parviflora Sw.; P. peltata Cav.] Hillsides, banks and
thickets, St. Thomas; St. Jan; St. Croix.
_PASSIFLORA FOETIDA L. Banks, waste and cultivated grounds,
St. bhomas; St: Croix:
PASSIFLORA MULTIFLORA L. Thickets, St. Thomas (according to
Masters); St. Jan.
ie BROOKLYN BOTANIC GARDEN MEMOIRS
PASSIFLORA RUBRA L. Woods and thickets, St. Thomas; St. Jan;
St. Croix (according to Eggers).
PASSIFLORA LAURIFOLIA L. Forests and thickets, St. Thomas;
St Jan; St. Croix. Perhaps not indigenous; much planted.
PASSIFLORA INCARNATA L., recorded from St. Croix by West, must
be an error in determination.
PASSIFLORA QUADRANGULARIS L. Planted for its fruit.
PASSIFLORA MALIFORMIS L. Planted for its fruit.
CARICACEAE
CaricA Papaya L. Spontaneous after cultivation, St. Thomas;
St. Crom. Much planted:
BEGONIACEAE
Several kinds of Begonias are grown as garden flowers.
BEGONIA HUMILIs Ait., attributed to St. Thomas by A. de Candolle
as collected by Finlay, was really from Trinidad.
CACTACEAE
HYLOCEREUS TRIGONUS (Haw.) Safford. [Cereus trigonus Haw.;
C. triangularis of West and of Eggers.] On trees and rocks in forests
and valleys, St. Thomas; St. Jan.
HYLOCEREUS UNDATUS (Haw.) Britton & Rose. [Cereus undatus
Haw.; Cereus triangularis of authors.] Persistent after cultivation,
Sti: Phomas- St, Crom
SELENICEREUS GRANDIFLORUS (L.) Britton & Rose. [Cereus
grandiflorus L.| Persistent after cultivation, St. Thomas; St. Croix.
SELENICEREUS PTERANTHUS (Link & Otto) Britton & Rose.
[Cereus nycticalis Link.| Recorded by Millspaugh as naturalized on
stone walls of a neglected garden at Bassin, St. Croix.
CEPHALOCEREUS ROYENI (L.) Britton & Rose. [Cactus Royeni L.;
C. peruvianus of West; Cereus floccosus Otto; Pilocereus Fouchianus
Weber; Cereus armatus Otto.] Dry rocky hillsides, St. Thomas;
Stfagi St. Croc.
CEPHALOCEREUS NOBILIS (Haw.) Britton & Rose. [Cereus nobilis
Haw.; Cereus strictus DC.| Persistent after cultivation, St. Thomas.
ACANTHOCEREUS PENTAGONUS (L.) Britton & Rose. [Cactus
pentagonus L.] Persistent after planting, St. Thomas; St. Croix.
Cactus IntTortus Mill. [C. Melocactus of West; Melocactus
communis of Eggers; M. atrosanguineus Link & Otto.] Coastal hills
and cliffs, St. Thomas; St. Jan; St. Croix.
CoRYPHANTHA NIVOSA (Link) Britton. [Mamiullaria nivosa Link.]
BRITTON: FLORA OF THE VIRGIN ISLANDS 73
Rocky slopes and cliffs, Buck Island and Flat Cays, St. Thomas; St.
Jan and Little St. James Island, St. Jan.
OPUNTIA RUBESCENS Salm-Dyck. [O. catacantha Link & Otto;
O. spinosissima and tuberculata of Eggers.] Coastal hills, St. Thomas;
St. Jan; St. Croix. The spineless or nearly spineless race is com-
monly planted for interest, and occurs wild on Little St. James Island,
St. Jan, and on Culebra. .
OPUNTIA REPENS Bello. [Cactus curassavicus of West; O. curas-
savica of Eggers and of Millspaugh.] Dry fields and hillsides, St.
Moomas; St. Jan; St. Croix.
Opuntia antillana Britton & Rose, spec. nov.
Plant depressed, ascending or nearly prostrate, often forming
clumps I m. broad, seldom more than 4 dm. high. Joints obovate or
oblong-obovate, 2 dm. long or less, green, glabrous, readily detached;
leaves conic-subulate, 2-3 mm. long; areoles large, 2-3 cm. apart,
brown-woolly; spines mostly 3-6 at each areole, subulate, rather stout,
terete, I-6 cm. long, yellow fading gray or nearly white; glochids
many, yellow; flowers about 7 cm. broad; petals obtuse, bright
yellew or fading reddish; fruit red-purple, about 4 cm. long.
Rocky and sandy soil, St. Thomas, St. Croix, also on Tortola, Porto
Rico, Hispaniola, and St. Kitts. Type specimen collected on St. Kitts
(Rose, Fitch & Russell 3230).
OpunTIA DILLENIIT (Ker.) Haw. [Cactus Dillenit Ker.; Cactus
Opuntia of West; O. Tuna of Eggers and of Millspaugh; O. horrida
Salm-Dyck.] Banks, fields and hills, St. Thomas; St. Jan; St. Croix.
A hybrid with O. rubescens was observed on Buck Island, St. Thomas.
OPUNTIA TRIACANTHA (Willd.) DC. [Cactus triacanthus Willd.]
Coastal rocks, Buck Island, St. Thomas.
NOPALEA COCHENILLIFERA (L.) Salm-Dyck. [Cactus cochenillifer
L.; Opuntia coccinellifera Mill.] Persistent or spontaneous after
cultivation; recorded by Eggers as occurring on limestone, St. Thomas;
=. Croix.
PERESKIA PERESKIA (L.) Karst. [Cactus Pereskia L.; P. aculeata
Mill.] Spontaneous after cultivation, St. Thomas; St. Croix.
PERESKIA GRANDIFOLIA Haw. [P. Bleo of Eggers and of Mills-
paugh.] Spontaneous after cultivation, St. Thomas; St. Croix.
CEREUS NORTHUMBERLANDIA Lambert. [C. Jepidotus Salm-
Dyck.] Planted, St. Croix.
CEREUS HEXAGONUS (L.) Mill. [C. peruvianus (L.) Mill.] Planted
(according to Eggers).
Other species of Cacti are occasionally cultivated for interest.
74 BROOKLYN BOTANIC GARDEN MEMOIRS
THYMELAEACEAE
DAPHNOPSIS CARIBAEA Griseb. [Nectandra antillana of Mills-
paugh.] Forests and hillsides, St. Thomas; St. Jan; St. Croix.
LYTHRACEAE
AMMANNIA COCCINEA Rottb. Moist ground, St. Thomas; St.
Jans St, Crotx.
AMMANNIA LATIFOLIA L. [A. sanguinolenta Sw.| Moist ground,
St. Thomas; St. Croix:
GINORIA Rourit (Vahl) Koehne. [Antherylium Rohrit Vahl.]
Coastal thickets, St. Thomas; st. Jan; St. Croix:
LAWSONIA INERMIS L. Spontaneous after cultivation, St. Thomas;
St.Crom
LAGERSTROEMIA INDICA L. Commonly planted for ornament.
PUNICACEAE
PuNICA GRANATUM L. [P.nanaL.] Spontaneous after planting,
St; Thomas: ot..Jan> St:-Groix. »-Growm for its innit
RHIZOPHORACEAE
RHIZOPHORA MANGLE L. Mangrove swamps, St. Thomas; St.
Jan; St. Croix. Not very common.
COMBRETACEAE
TERMINALIA CATAPPA L. [Buceras Catappa Hitchc.] Hillsides,
valleys, and commonly planted, St. Thomas; St. Jan; St. Croix.
CONOCARPUS ERECTA L. [C. erecta procumbens Jacq.] Coastal
rocks and mangrove swamps, St. Thomas; St. Jan; St. Croix.
BucipA Buceras L. [Buceras Buceras Millsp.; Myrobalanus
Buceras Kuntze.| Moist soil, mostly near the coasts, but occasional
on hillsides, St. Thomas; St. Jan; St. Croix.
LAGUNCULARIA RACEMOSA (L.) Gaertn. [Conocarpus racemosa L.]
Coastal swamps, St. Thomas; St. Jan; St. Croix.
QUISQUALIS INDICA L. is commonly cultivated as an ornamental
vine,
MYRTACEAE
Pstip1um GuajAva L. Thickets, hillsides, and commonly planted
for its fruit, St: Thomas; St. Jan; St. Croix.
PsIDIUM AMPLEXICAULE Pers. [P. cordatum Sims.] Hillsides, St.
Thomas; St. Jan; planted on St. Croix. Occurs also on Tortola.
Apparently endemic in the Virgin Islands, although recorded from
Nevis.
BRITTON: FLORA OF THE VIRGIN ISLANDS 75
PsIDIUM AROMATICUM Knox, recorded from St. Thomas, is not
identified.
AMOMIS CARYOPHYLLATA (Jacq.) Krug & Urban. [Myrtus caryo-
phyllata Jacq.; Myrtus acris Sw.; M. Pimenta Ortega; ? Pimenta
vulgaris of Eggers; Pimenta acris Kostel.; A. caryophyllata grisea
Krug & Urban.| Hills and woods, St. Jan; St. Croix (according to
Eggers).
MyrCIA PANICULATA (Jacq.) Krug & Urban. [Eugenia paniculata
Jacq.; E. acetosans Poir.; E. marginata Pers.; Myrtus coriacea Vahl;
Myrcia coriacea DC.; M. coriacea Imrayana Griseb.|] Forests, St.
fnomas; St. Jan; St. Croix.
MyRcIA SPLENDENS (Sw.) DC., doubtfully accredited to St.
Thomas by Urban, as collected by Riedlé, probably was from Porto
Rico, where it is abundant.
CALYPTRANTHES THOMASIANA Berg. Signal Hill, St. Thomas;
Bordeaux, St. Jan. Endemic.
CALYPTRANTHES PALLENS (Poir.) Griseb. [C. Chytraculia ovalis
Berg.; C. Chytraculia suzygium Berg.; Chytraculia pallens Muillsp.;
C. Chytraculia of West.] Forests, rare, St. Thomas; St. Croix.
EUGENIA LIGUSTRINA (Sw.) Willd. [Myrtus ligustrina Sw.; M.
cerasina Vahl.] Woods and thickets, St. Thomas; St. Jan; St. Croix.
EUGENIA LANCEA Poir. [E. ludibunda Bert.; E. virgultosa of
Eggers and of Millspaugh; £. glabrata of Eggers; Myrcia thomasiana
DC.] Woods and thickets, St. Thomas; St. Croix.
EUGENIA MONTICOLA (Sw.) DC. [Myrtus monticola Sw.; Eugenia
Poirett Berg, not DC.; FE. foetida West; FE. flavovirens Berg.]
Woods and thickets, St. Thomas; St. Jan; St. Croix. .
EUGENIA BUXIFOLIA (Sw.) Willd. [Myritus buxifola Sw.; E.
foetida Poir.] Thickets, St. Thomas; St. Croix.
EUGENIA AXILLARIS (Sw.) Willd. [Myrtus axillaris Sw.| Thick-
Cts, ot. Croix.
EUGENIA RHOMBEA (Berg) Krug & Urban. [E. foetida rhombea
Berg.; E. Poiretit of Millspaugh; ? E. pallens of Eggers.]| Coastal .
thickets, St. Thomas (according to Berg); St. Croix.
EUGENIA PROCERA (Sw.) Poir. [Myrtus procera Sw.; M. cerasina
Vahl of Eggers.] Woods and thickets, St. Thomas; St. Jan; St.
Croix.
EUGENIA PSEUDOPSIDIUM Jacq. [E. portoricensis DC.; E. thomasi-
ana Berg.| Forests and wooded valleys, St. Thomas; St. Jan; St.
Croix.
.EUGENIA CORDATA (Sw.) DC. [Myrtus cordata Sw.; M. ramiflorus
Vahl; E. sessiliflora DC., not Vahl; E. lateriflora of Eggers.] Woods
and thickets, St. Thomas; St. Jan; St. Croix.
76 BROOKLYN BOTANIC GARDEN MEMOIRS
EUGENIA SESSILIFLORA Vahl. Hillsides, St. Thomas (according
to Eggers); St. Croix. Endemic.
EUGENIA FLORIBUNDA West. Woods, hillsides and thickets, St.
Thomas: St. jan; St: Grom.
EUGENIA UNIFLORA L. Spontaneous after planting, St. Thomas;
St. Croix. Grown for its fruit.
EUGENIA MICRANTHA Vahl, not DC., recorded from St. Croix by
West, is not further determined (hyponym).
EUGENIA EMARGINATA Vahl, not DC., recorded from St. Croix by
Vahl, is not further determined (hyponym).
EUGENIA PEDUNCULATA Raeusch. of St. Croix, is unknown to
modern botanists.
ANAMOMIS FRAGRANS (Sw.) Griseb. [Myrtus fragrans Sw.;
Eugenia punctata Vahl; Anamomis punctata Griseb.] St. Croix;
forests, St. Jan (according to Eggers).
JAmBos JAmBos (L.) Millsp. [Eugenta Jambos L.; Jambos vul-
garis DC.] Woods and valleys, naturalized, St. Thomas; St. Jan;
St Croix ‘
JAMBOS MALACCENSIS (L.) DC. [Eugenia malaccensis L.], planted
for its fruit, was naturalized in shaded valleys, St. Croix (according to
Eggers).
Myrtus CoMMuNIS L. Planted for ornament.
LECY THIDACEAE
COUROUPITA GUIANENSIS Aubl. Planted for ornament and in-
terest.
MELASTOMACEAE
TETRAZYGIA ANGUSTIFOLIA (Sw.) DC. [Melastoma angustifolia
Sw.; Miconia angustifolia Griseb.] Hillside thickets, St. Thomas;
St. Jan; St. Croix (according to Cogniaux).
TETRAZYGIA ELAEAGNOIDES (Sw.) DC. [Melastoma elaeagnoides
Sw.] Forests and hillside thickets, St. Thomas; St. Jan; St. Croix.
MICONIA MACROPHYLLA (D. Don) Triana. [Chitonia macrophylla
D. Don; Diplochita serrulata DC.] Wooded valleys, St. Thomas;
St. Croix (according to Eggers and to Cogniaux).
MICONIA IMPETIOLARIS (Sw.) D. Don. [Melastoma impetiolarts
Sw.] Forests, St. Croix (West); St. Thomas (according to Grise-
bach).
MICONIA LAEVIGATA (L.) DC. [Melastoma laevigata L.; M.
prasina of Millspaugh, St. Thomas.] Woods and hillsides, St. Thomas;
St. Jans; St2i\Grom:
MICONIA PRASINA (Sw.) DC., recorded by Naudin as collected on
BRITTON: FLORA OF THE VIRGIN ISLANDS 77
St. Thomas by Riedlé, was probably from Porto Rico. It occurs on
Tortola.
MICONIA STENOSTACHYA (Schr.) DC. [Miconia argyrophylla
Benth., not DC.], recorded by Naudin and others as collected by
Finlay on St. Thomas, was really from Trinidad.
MICONIA THOMASIANA DC. was not from St. Thomas, but from
Porto Rico.
MICONIA ACINODENDRUM (L.) Triana. [Melastoma acinodendrum
L.; Tshudya berbiceana Griseb.], recorded by Naudin and others as
collected on St. Thomas by Finlay, was really from Trinidad; St.
Croix (according to West).
MECRANIUM AMYGDALINUM (Desr.) C. Wright [Cremanium
amygdalinum Griseb.], attributed to St. Thomas by Naudin, was from
Hispaniola, collected by Poiteau.
CLIDEMIA SPICATA DC., is recorded by Cogniaux as collected on
St. Thomas by Finlay, but the specimen was from Trinidad. Eggers
erroneously records it from all three islands.
CLIDEMIA HIRTA (L.) D. Don, attributed to St. Thomas by Naudin
as collected by Riedlé, was probably from Porto Rico.
CLIDEMIA RUBRA Mart, accredited to St. Thomas by Naudin and
others as collected by Finlay, was really from Trinidad.
MOuRIRIA DOMINGENSIS (Tuss.) Spach. [Petaloma domingensis
Tuss.; P. Mouriri of West.] Borders of a stream, Spring Garden,
St. .Acroix.
NEPSERA AQUATICA (Aubl.) Naud., recorded by Cogniaux as col-
lected by Riedlé on St. Thomas, was probably from Porto Rico.
ONAGRACEAE
JUSSIAEA SUFFRUTICOSA L. [J. angustifolia Lam.; J. octovalvis
Sw.; J. suffruticosa ligustrifolia of Eggers.] Wet grounds, St. Thomas;
eewian; ot. Croix.
JUSSIAEA ERECTA L. St. Croix (according to West).
ARALIACEAE
DENDROPANAX ARBOREA (L.) Dene. & Pl. [Sciadophyllum capi-
tatum of Eggers; Aralia arborea L.; Gilibertia arborea E. March.]
Forests, hills of St. Thomas.
DIDYMOPANAX MICANS (Willd.) Krug & Urban. [Aralia micans
Willd.; Panax speciosum of Eggers.] Forest, King’s Hill and Bor-
deaux, St. Jan.
APIACEAE
ERYNGIUM FOETIDUM L. Moist grounds, Caret Bay, St. Thomas
(according to Eggers).
78 BROOKLYN BOTANIC GARDEN MEMOIRS
CELERI GRAVEOLENS (L.) Britton. [Apium graveolens L.; Peuce-
danum graveolens Benth.] Persistent after cultivation, St. Croix.
Grown for celery.
ANETHUM GRAVEOLENS L. Spontaneous after cultivation, St. Thomas;
St. Croix:
PIMPINELLA ANIsUM L. Spontaneous after cultivation, St. Croix.
Grown for anise.
Aptum PETROSELINUM L. [Petroselinum sativum Hoffm.] Cultivated
for food.
Daucus Carota L. Cultivated for food.
FoENICULUM FoENICULUM (L.) Karst. [F. vulgare Gaertn.] Cul-.
tivated for drug purposes.
CEREFOLIUM CEREFOLIUM (L.) Britton. [Anthriscus Cerefolium L.]
Cultivated for flavoring.
ERICACEAE
XOLISMA RUBIGINOSA (Pers.) Small. [Andromeda rubiginosa Pers.;
Lyonia jamaicensis of Eggers.] Bolongo, St. Thomas. Known other-
wise only from Hispaniola.
THEOPHRASTACEAE
JACQUINIA BERTERII Spreng. [J. Berterit retusa Urban.] Thick-
ets, St. Thomas; St. Jan; St. Croix (according to Mez).
JacguintA Barsasco (Loefl.) Mez. [Chrysophyllum Barbasco
Loefl.; J. armillaris Jacq.; J. armillaris arborea of Eggers.] Coastal
thickets, St. Thomas; St. Jan; St. Croix.
MYRSINACEAE
ICACOREA GUADALUPENSIS (Duch.) Britton. [Ardisia guadalu-
pensis Duch.; A. coriacea of West and of Eggers.] Forests and hills,
St. Thomas; St: Jan: St. Crorx.
STYLOGYNE LATERIFLORA (Sw.) Mez. [Ardisia lateriflora Sw.;
A. caribaea Miquel.] St. Thomas (according to Eggers). ;
PLUMBAGINACEAE
PLUMBAGO SCANDENS L. __[P. scandens densiflora Kuntze.] Banks,
woods and thickets, St. Thomas; St. Jan; St. Croix.
PLUMBAGO CAPENSIS Thunb. Grown in gardens.
PLUMBAGO ZEYLANICA L. St. Croix (according to West).
SAPOTACEAE
Sapota Acuras Mill. [Achras Sapota L.] Forests, and com-
monly planted, St. Thomas; St. Jan; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 79
LuUCUMA MULTIFLORA A. DC. [Achras multiflora Vahl, according
to Eggers.] Forests, St. Thomas; St. Croix.
SIDEROXYLON FOETIDISSIMUM Jacq. [S. mastichodendron Jacq.]
Forests, St. Thomas; St. Jan; St. Croix.
DIPHOLIS SALICIFOLIA (L.) A. DC. [Achras salicifolia L.; Bumelia
salicifolia Sw.| Forests and hillsides, St. Thomas; St. Jan; St. Croix.
BUMELIA OBOVATA (Lam.) A. DC. [Sideroxylon obovatum Lam.;
B. cuneata Sw.| Coastal hillsides, shores and borders of marshes,
ret nomas; St. Jan; St. Croix.
CHRYSOPHYLLUM CAINITO L. St. Thomas; St. Croix, occasionally
planted, perhaps spontaneous.
CHRYSOPHYLLUM EGGERsII Pierre. [C. microphyllum of Eggers;
? C. oliviforme monopyrenum of Eggers.] Woods and hillsides, St.
Thomas; St. Jan; St. Croix. Endemic.
CHRYSOPHYLLUM PAUCIFLORUM Lam. [C. glabrum of Eggers and
of Millspaugh; C. pauciflorum nervosum Pierre.| Forests and hill-
sides, St. Thomas; St. Jan; St. Croix. Otherwise known only from
Porto Rico. ’
CHRYSOPHYLLUM BICOLOR Pierre. St. Thomas (according to
Pierre and Urban); known otherwise only from Porto Rico.
Mimusops NITIDA (Sessé & Mog.) Urban. [Sapota Sideroxylon
of Eggers.] Forests, St. Jan. Determined from foliage only; its
flowers and fruit have not been collected by botanists, and the identi-
fication is, therefore, uncertain.
Mimusops ELENGI L. Planted, St. Thomas.
SYMPLOCACEAE
SYMPLOCOS MARTINICENSIS Jacq. Forest, Signal Hill, St. Thomas.
OLEACEAE
FORESTIERA SEGREGATA (Jacq.) Krug & Urban. [Myrica segregata
Jacq.; Forestiera porulosa Poir; F. porulosa Jacquinii Eggers.]
Thickets, St. Croix.
FORESTIERA EGGERSIANA Krug & Urban. Thickets, St. Thomas;
St. Jan. Known otherwise only on Culebra, Vieques and Virgin
Gorda, thus endemic in the Virgin Islands.
FORESTIERA RHAMNIFOLIA Griseb. [Drypetes laevigata of Mills-
paugh.] Bluffs of Salt River, St. Croix.
MAYEPAEA CARIBAEA (Jacq.) Kuntze. [Chionanthus caribaea
Jacq.; C. compacta Sw.; Linociera compacta R. Br.] Forests, St.
Thomas; St. Croix.
JASMINUM SamBac (L.) Soland. [Nyctanthes Sambac L.; J.
quinqueflorum Heyne.] Spontaneous after planting, St. Croix.
80 BROOKLYN BOTANIC GARDEN MEMOIRS
JASMINUM PUBESCENS (Retz) Willd. [Nyctanthes pubescens Retz.]
Spontaneous after planting, St. Thomas; St. Croix.
JASMINUM GRANDIFLORUM L._ [J. officinale of Millspaugh.] Spon-
taneous after planting, St. Thomas; St. Croix.
JASMINUM OFFICINALE L. Planted for ornament.
JASMINUM HUMILE L. [J. revolutum Sims.] Planted for orna-
ment.
OLEA EUROPAEA L. Planted on St. Thomas.
LOGANIACEAE
SPIGELIA ANTHELMIA L. Moist or exsiccated situations, St.
Thomas: St. "Crom:
APOCYNACEAE
ALLAMANDA CATHARTICA L. Spontaneous after cultivation, St.
Thomas.
PLUMIERA ALBA L. Coastal rocks and hills, St. Thomas; St. Jan;
St. Croix
PLUMIERA RUBRA L._ [P. obtusifolia of Millspaugh.] Planted for
ornament.
PLUMIERA OBTUSA L. St. Croix (according to West); naturalized
in gardens (according to Eggers).
CATHARANTHUS ROSEUS (L.) D. Don. [Vinca rosea L.; Lochnera
rosea Rchb.] Waste grounds, spontaneous after cultivation, and
much planted for ornament, St. Thomas; St. Jan; St. Croix.
TABERNAEMONTANA CAPENSIS L. Planted for ornament.
A species of Tabernaemontana was found in thickets at French-
man’s Bay, St. Thomas, according to Eggers, who, doubtfully, records
it as T. citrifolia.
RAUWOLFIA TETRAPHYLLA L. [R. nitida Jacq.] Woods, hillsides
and thickets, St. Thomas; St. Jan; St. Croix.
RAUWOLFIA LAamMARCKIT A. DC. Hillsides and thickets, St. Thom-
aS; ot. Jal; ot Croix.
CERBERA THEVETIA L. [Thevetia Thevetia Millsp.; JT. neritfola
Juss.] Hillsides and thickets, St. Thomas; St. Jan; St. Croix.
ECHITES AGGLUTINATA Jacq. [F. circinalis Sw.] Forests and
thickets, St. Thomas; St. Jan; St. Croix at Cane Bay (according to
Eggers).
URECHITES LUTEA (L.) Britton. [Vinca lutea L.; Echites sub-
erecta Jacq.; E. barbata Desv.; E. neriandra Griseb.] Thickets, St.
Thomas; St. Jan; St. Croix (according to Eggers).
NERIUM OLEANDER L. Persistent after cultivation; planted for
ornament; St. Thomas; St. Jan; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 81
ASCLEPIADACEAE
ASCLEPIAS CURASSAVICA L. [A. mnivea curassavica Kuntze.]
Fields, hillsides and banks, St. Thomas; St. Jan; St. Croix.
ASCLEPIAS NIVEA L. Collected on St. Thomas by Krebs.
ASCLEPIAS FRUTICOSA L. Cultivated on St. Croix (according to
West).
CALOTROPIS PROCERA (Ait.) Ait. f. [Asclepias procera Ait.] Fields
and hillsides, St. Thomas; St. Jan; St. Croix. Naturalized.
METASTELMA PARVIFLORUM R. Br. [M. Schlechtendalu of Eggers
and of Millspaugh.] Thickets, St. Thomas; St. Croix.
METASTELMA ALBIFLORUM Griseb. St. Thomas (according to
Schlechter). :
METASTELMA DECIPIENS Schlechter. St. Thomas (according to
Schlechter).
METASTELMA GRISEBACHIANUM Schlechter. St. Thomas. Other-
wise known only from Porto Rico.
METASTELMA DECAISNEANUM Schlechter. Hillsides, St. Thomas;
pila; St. Croix.
The above-listed species of Metastelma much resemble each other.
It is possible that Schlechter has recognized too many species in the area.
OXYPETALUM CORDIFOLIUM (Vent) Schlechter. [Gothofreda cordi-
folia Vent.; O. riparium H.B.K.] St. Thomas (according to Schlech-
ter).
PHILIBERTELLA CLAUSA (Jacq.) Vail. [Asclepias viminalis Sw.;
Sarcostemma Brownei Meyer], recorded as from St. Thomas by West
and by Eggers, has not been observed there by recent collectors.
FISCHERIA CRISPIFLORA (Sw.) Schltr. [Cynanchum crispiflorum
Sw.; F. scandens DC.| Forests, Spring-gut, St. Croix (according to
Eggers). Known otherwise only from Cuba and Jamaica; the
determination is doubtful.
IBATIA MARITIMA (Jacq.) Dene. [Asclepias maritima Jacq.;
Ibatia muricata Griseb.| Rocky hillsides and thickets, St. Thomas;
Peefan; ot. Croix.
Hoya carnosa (L. f.) R. Br. Cultivated for ornament.
STEPHANOTIS FLORIBUNDA A. Brongn. Cultivated for ornament.
CUSCUTACEAE
CUSCUTA AMERICANA L. On trees and shrubs, St. Thomas; St.
jan; St. Croix.
CONVOLVULACEAE
EVOLVULUS NUMMULARIUS L. Dry, shaded situations, St. Thomas;
mt. lan; ot. Croix.
7
82 BROOKLYN BOTANIC GARDEN MEMOIRS
EVOLVULUS GLABER Spreng. [E. mucronatus Sw.] Moist, grassy
situations, St. Vhomnias; St. Jan; st. Croi,
EVOLVULUS LINIFOLIUS L. St. Thomas and St. Croix (according
to Schlechtendal); moist localities, all islands (according to Eggers).
Not found by us, and not known on Porto Rico.
JACQUEMONTIA NODIFLORA (Desr.) G. Don. [Convolvulus nodi-
florus Desr.; C. albiflorus West (hyponym).] Thickets, St. Thomas;
St, Jans7St. Crom,
JACQUEMONTIA JAMAICENSIS (Jacq.) Hall. f. [Convolvulus jamai-
censis Jacq.] Coastal thickets, St. Thomas; St. Croix.
JACQUEMONTIA PENTANTHA (Jacq.) G. Don. [Convolvulus pen-
tanthus Jacq.; C. violaceus Vahl; J. violacea Choisy.] Thickets, St.
Thomas: St. Jany) StaCrom,
CONVOLVULUS MATUTINUS West and C. VENENATUS West, described
from St. Croix, are not further identified.
THYELLA TAMNIFOLIA (L.) Raf. [Ipomoea tamntfolia L.; Jacque-
montia tamnifolia Griseb.]| Banks, hills and thickets, St. Thomas;
St. Jan (according to Eggers); St. Croix. :
Exogonium solanifolium (L.) Britton. [Ipomoea solantfolia L.;
I. filiformis Jacq.; Convolvulus filiformis Desr.; Exogonium filiforme
Choisy; Ipomoea eustachiana of Millspaugh.] . Coastal thickets, St.
Thomas: St. Crotx.
EXOGONIUM REPANDUM (Jacq.) Choisy. [Ipomoea repanda Jacq.]
Woods and forests, St. Thomas; St. Jan.
EXOGONIUM ARENARIUM Choisy. [Ipomoea arenaria Steud.; I.
Steudelu Millsp.; I. Eggersiana Peter; E. Eggersit House.| Thickets,
St. suomas; St: Jan: ot, Crop.
IPOMOEA DISSECTA (Jacq.) Pers. [Convolvulus dissectus Jacq.;
Merremia dissecta Hall. f.; Operculina dissecta House; Ipomoea sinuata
Ort.] Thickets, fences and woods, St. Thomas; St. Jan; St. Croix.
IPOMOEA AEGYPTIA L. [Convolvulus pentaphyllus L.; Ipomoea
pentaphylla Jacq.; Merremia aegyptia Urban.] Fields, fences and
thickets; St. Thomas; St. Jan; St. Croix.
IPOMOEA QUINQUEFOLIA L. [Merremia quinquefolia Hall. f.]
Thickets, St. Thomas; St. Croix.
IPOMOEA POLYANTHES R. & S._ [Convolvulus umbellatus L.;
Ipomoea umbellata Meyer, not L.; Merremia umbellata Hall. f.; Ipomoea
mollicoma Miq.; Convolvulus sagittifer H.B.K.] Fields, banks and
thickets, St. Thomas; St. Jan; St. Croix.
Ipomora Nit (L.) Roth. — [Convolvulus Nil L.; Pharbitis Nil
Choisy; JI. hederacea barbata of Kuntze; ? Convolvulus hederaceus of
Schlechtendal.]_ Banks, waste and cultivated grounds, St. Thomas;
st. Jan: St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 83
* IPOMOEA CATHARTICA Poir. [Convolvulus acuminatus Vahl; Ipo-
moea acuminata R. & S., not R. & P.; Pharbitis cathartica Choisy;
P. acuminata Choisy.]| Woods and thickets, St. Thomas; St. Jan;
St. Croix.
IPOMOEA ASARIFOLIA (Desr.) R. & S. [Convolvulus asarifolius
Desr.| Danish Islands (according to Grisebach).
IPOMOEA PES-CAPRAE (L.) Roth. ([Convolvulus Pes-caprae L.;
C. brasiliensis L.; C. maritimus Lam.] Coastal sands, St. Thomas;
Beeyan; St. Croix.
IPOMOEA STOLONIFERA (Cyr.) Poir. [Convolvulus littoralis L.;
C. arenarius Vahl; Ipomoea littoralis Boiss.; Convolvulus stoloniferus
Cyr.; Ipomoea acetosaefolia R. & S.] Coastal sands, St. Croix (ac-
cording to West).
IPOMOEA HEPTAPHYLLA (Rottl. & Willd.) Voigt. [Convolvulus
heptaphyllus Rottl. & Willd.; Ipomoea pulchella Griseb., not Roth.]
St. Thomas (according to Urban).
IPOMOEA CARNEA Jacq. St. Croix (according to West).
IPOMOEA TRILOBA L. [Convolvulus Sloanet Spreng.; I. parviflora
Vahl; I. triloba eustachiana of Eggers; I. triloba quinqueloba Kuntze. ]
Fields, banks and thickets, St. Thomas; St. Jan; St. Croix.
IPOMOEA TILIACEA (Willd.) Choisy. [Convolvulus tiliaceus Willd.;
C. fastigiatus Roxb.; I. fastigiata Sweet; I. Batatus fastigiata Kuntze.]
Thickets and hillsides, St. Thomas; St. Jan; St. Croix.
IpomorA Batatas (L.) Lam. [Convolvulus Batatas L.; Ipomoea
pandurata cuspidata Kuntze.] Persistent after cultivation, St.
Thomas; St. Jan; St. Croix. Much planted for food.
IPOMOEA TRICOLOR Cav. [J. violacea Grisebach, and of Eggers
and Millspaugh.] Woods and thickets, St. Thomas; St. Jan; St.
Croix.
IPOMOEA PURPUREA (L.) Lam. [Convolvulus purpureus L.] St.
Croix (according to West). Planted for ornament.
IpoMoEA LEARII Paxton. Planted for ornament.
IPpoMOEA PEs-TIGRIDIS L. [Convolvulus Pes-tigridis L.] St. Thom-
as (according to Schlechtendal).
IpoMOEA HorsFALLIAE W. Hook. Planted for ornament.
IPOMOEA QUINQUEPARTITA (Vahl) R. & S. [Convolvulus quinque-
partitus Vahl; C. ovalifolius West, not Vahl,] of St. Croix, is not further
identified.
IPOMOEA LEUCANTHA Jacq., a South American species, is recorded
by Eggers from St. Thomas and St. Croix.
OPERCULINA TRIQUETRA (Vahl) Hallier f. [Convolvulus triqueter
Vahl; Ipomoea triquetra R. & S.] St. Thomas; St. Croix.
OPERCULINA TUBEROSA (L.) Meissn. [Ipomoea tuberosa L.|
Forests, St. Thomas; St. Croix (according to Eggers).
84 BROOKLYN BOTANIC GARDEN MEMOIRS
CALONYCTION ACULEATUM (L.) House. [Convolvulus aculeatus L.;
Ipomoea Bona-nox L.] Spontaneous after cultivation, St. Thomas.
Planted for ornament.
CALONYCTION TUBA (Schlecht.) Colla. [Convolvulus tuba Schlecht. ;
Ipomoea tuba G. Don.] Coastal thickets, St. Thomas; St. Jan; St.
Croix.
QuAMOCLIT QuAmoc.itT (L.) Britton. [Ipomoea Quamoclit L.;
Quamoclit vulgaris Choisy.] Banks, thickets and cultivated grounds,
St: Thomas). St. Croix:
QUAMOCLIT COCCINEA (L.) Moench. [Ipomoea coccinea L.; I.
hederaefolia L.; I. sanguinea Vahl.] Banks, thickets and cultivated
grounds, St. Thomas; St. Jan; St. Croix.
RIVEA TILIIFOLIA (Desr.) Choisy. [Convolvulus tilifolius Desr.;
Aregyreia tiliifolia Wright; Convolvulus melanostictus Schl.| Thickets,
St. thomas; St. Jan St. Crom,
POLEMONIACEAE
PHLtox DrumMonpiI Hook. Grown in flower gardens.
HY DROPHYLUAGEAE
MARILAUNIDIUM JAMAICENSE (L.) Kuntze. [Nama jamaicensis L.;
Hydrolea jamaicensis Vahl.] Dry, rocky situations, St. Thomas;
St: Crom.
CORDIACEAE
CERDANA ALLIODORA R. & PP. [Cordia Gerascanthus Jacq., not L.;
C. Gerascanthus subcanescens of Eggers.]| Woods and forests, St.
Thomas; St. Jan.
SEBESTEN SEBESTENA (L.) Britton. [Cordia Sebestena L.; C.
Rickseckert Millsp.| Coastal thickets, hillsides, and planted for orna-
ment, St bhomas: St. Jan‘ St.Croi: ‘
SEBESTEN BRACHYCALYX (Urban) Britton. [Cordia Sebestena
brachycalyx Urban.] Rocky hillside, Buck Island, St. Thomas.
Known otherwise only from Porto Rico.
CorDIA ALBA (Jacq.) R. & S. [Varronia alba Jacq.] Thickets
and hillsides, St. Thomas; St. Croix. Sometimes planted.
Corpia coLtLococca L. [C. micrantha Sw.| Woods, forests and
hills Sts Thomas-:St fan; -st..Croix.
CorpIA NITIDA Vahl. [? C. laevigata of Schlechtendal.] Forests
and hills, St. Themas; St! Jan; St. Croix.
CorpIA SULCATA DC. [C. macrophylla R. & S.| Forests and
hills, St. Thomas; St. Jan; St. Croix (according to West).
VARRONIA CORYMBOSA (L.) Desv. [Lantana corymbosa L.; Cordia
BRITTON: FLORA OF THE VIRGIN ISLANDS 85
ulmifolia Juss.; C. ulmifolia and varieties of Eggers.| Thickets,
fields and hillsides, St. Thomas; St. Jan; St. Croix.
VARRONIA ANGUSTIFOLIA West. [Cordia angustifolia R. & S.;
C. cylindrostachya and varieties of Eggers; C. cylindrostachya of
Millspaugh.| MHillsides and thickets, St. Thomas; St. Croix (type
locality).
CORDIA MARTINICENSIS R. & S. is accredited to St. Croix by
Grisebach (Fl. 481); the record probably refers to V. angustifolia.
VARRONIA GLOBOSA Jacq. [Cordia globosa H.B.K.| Thickets, St.
Thomas and St. Croix (according to West, Schlechtendal and
Eggers).
BOURRERIA SUCCULENTA Jacq. [Ehretia Bourreria L.| Forests,
hillsides and thickets, St. Thomas; St. Jan; St. Croix.
ROCHEFORTIA ACANTHOPHORA (DC.) Griseb. [Ehretia acantho-
phora DC.; (2?) Ehretia spinosa Jacq.| Thickets, St. Thomas; St.
Jan; St. Croix (according to West and to Eggers).
BORAGINACEAE
TOURNEFORTIA FILIFLORA Griseb. [TJ. foetidissima DC. and of
Eggers, not L.] St. Jan (according to Eggers); St. Croix (according
to West).
TOURNEFORTIA HIRSUTISSIMA L. Thickets, banks and hills, St.
imemas;: St. Jan; St. Croix.
TOURNEFORTIA BICOLOR Sw. [T7. laevigata Lam.] Among rocks,
Crown, St. Thomas.
TOURNEFORTIA LAURIFOLIA Vent., attributed to St. Thomas by
Ventenat and by de Candolle, is known to us only from Porto Rico.
TOURNEFORTIA VOLUBILIS L. Thickets, St. Thomas; St. Jan; St.
Croix (according to Eggers).
TOURNEFORTIA MICROPHYLLA Bert. [T. volubilis microphylla DC.;
T. volubilis microcarpa of Millspaugh.] Thickets, St. Thomas; St.
Wan; St. Croix.
MALLOTONIA GNAPHALODES (L.) Britton. [Heliotropium gnapha-
lodes L.; Tournefortia gnaphalodes R. Br.| Coastal sands, St. Thomas;
Bee yan; St. Croix.
HELIOTROPIUM CURASSAVICUM L. Saline soil, St. Thomas; St.
an. ot. Croix.
HELIOTROPIUM PARVIFLORUM L. Banks, hillsides, waste and
cultivated grounds, St. Thomas; St. Jan; St. Croix.
HELIOTROPIUM INDICUM L. Waste and cultivated grounds, St.
a aomas; St. Jan; St. Croix.
HELIOTROPIUM TERNATUM Vahl. [Heliotropium fruticosum L.,
86 BROOKLYN BOTANIC GARDEN MEMOIRS
in part, and of Eggers and Millspaugh.] Rocky thickets and hillsides,
St. Thomas; St. fang ot. Crom.
HELIOTROPIUM PERUVIANUM L. Cultivated in flower gardens.
VERBENACEAE
LANTANA CAMARA L. [L. scabrida Ait.] Thickets and hillsides,
>t, Thomas; St. Jans St. Croix
LANTANA ACULEATA L. [L. polyacantha Schauer.| Waste
grounds, St. Thomas; St. Croix.
LANTANA INVOLUCRATA L. [L. odorata L.; Camara involucrata
Kuntze.] .Thickets and hillsides. St. Thomas; St. Jan; St. Croix.
LANTANA RETICULATA Pers. Stony ground, King’s Hill, St. Croix
(according to Eggers).
LipPIA REPTANS H.B.K. [Lippia nodiflora of Eggers and of
Millspaugh.] Wet ground, St. Croix.
LIpPIA TRIPHYLLA (L’Her.) Kuntze. [Aloysia citriodora Ort.]
Cultivated in gardens.
BOUCHEA PRISMATICA (L.) Kuntze. [Verbena prismatica L.; B.
Ehrenbergit Cham.] Waste and cultivated grounds, St. Thomas;
Stwecoix.
VALERIANODES JAMAICENSIS (L.) Medic. [Verbena jamaicensis L.;
Stachytarpheta jamaicensis Vahl; Valerianodes jamaicensis indicus
Kuntze.] Fields, hills, banks and in cultivated grounds, St. Thomas;
Si: janeSr..Crotx.
VALERIANODES STRIGOSA (Vahl) Kuntze. [Stachytarpheta strigosa
Vahl.| Thickets and hillsides, St. Thomas; St. Jan.
PRIVA LAPPULACEA (L.) Pers. [Verbena lappulacea L.; V. mext-
cana of West; P. echinata Juss.]| Waste and cultivated grounds,
St.. (hemas:-St. Janz St: Croix:
CITHAREXYLUM FRUTICOSUM L. [C. cinereum L.; C. villosum
Jacq.] Woods, hills and thickets; St. Thomas; St. Jan; St. Croix.
CITHAREXYLUM SPINOSUM L. [C. quadrangulare Jacq.] Forests
and slopes, St. Thomas; St. Croix. Planted for shade.
DURANTA ERECTA L. [D. Plumiert Jacq.; D. Ellisia Jacq.; ? D.
repens L.| Thickets and hillsides, St. Thomas; St. Jan; St. Croix.
Sometimes planted for ornament.
CALLICARPA RETICULATA Sw., accredited to St. Croix by West, is
a little-known species of Jamaica; what plant West had in mind is
not further recorded.
AEGIPHILA MARTINICENSIS Jacq. Forests, St. Thomas; St. Croix,
common (according to Eggers).
PETITIA DOMINGENSIS Jacq. In forests, St. Croix, not common
(according to Eggers).
BRITTON: FLORA OF THE VIRGIN ISLANDS 87
VITEX DICARICATA Sw. Forests, St. Thomas; St. Jan; St. Croix
(according to Eggers), and recorded from St. Croix by Swartz.
Vitex AGNus-castus L. Planted for ornament.
VOLKAMERIA ACULEATA L. [Clerodendron aculeatum Schlecht. ;
C. aculeatum grandifolium and parvifolium Kuntze; ? C. longicollis of
Borgesen and Paulsen.| Thickets, St. Thomas; St. Jan; St. Croix.
CLERODENDRON FRAGRANS Vent. [C. fragrans pleniflora Schauer. |
Waste grounds, St. Thomas.
SIPHONANTHUs INDICUS L. [Clerodendron Siphonanthus R. Br.]
Woods near Grove Place and at Crequis, St. Croix. Apparently
naturalized. Planted for ornament.
AVICENNIA NITIDA Jacq. [A. tomentosa Jacq.; A. officinalis nitida
Kuntze.] Coastal swamps and lagoons, St. Thomas; St. Jan; St.
Croix.
PETRAEA VOLUBILIS Jacq. Planted for ornament.
HOL MSKOLDIA SANGUINEA Retz. Planted for ornament.
VERBENA ICHAMAEDRIFOLIA Juss. Planted for ornament.
LAMIACEAE
LEONOTIS NEPETIFOLIA (L.) R. Br. [Phlomis nepetifolia L.]
Waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
LEONURUS SIBIRICUS L. Waste and cultivated grounds, St.
Thomas; St. Jan; St. Croix.
MOLUCccELLA LAEVIS L. St. Croix (according to West).
LEUCAS MARTINICENSIS (Jacq.) R. Br. [Clhinopodium martinicense
Jacq.] Waste and cultivated grounds, St. Croix.
SALVIA THOMASIANA Urban. [S. tenella of Schlechtendal and of
Eggers.] St. Thomas. Endemic. Known only from a_ specimen
collected long ago by Ehrenberg.
SALVIA OCCIDENTALIS Sw. LS. occidentalis bicolor Kuntze.| Banks,
fields and thickets, St. Thomas; St. Jan; St. Croix.
SALVIA SEROTINA L. [S. dominica Sw.; S. micrantha Vahl.]
Banks, fields and hillsides, St. Thomas; St. Jan; St. Croix.
SALVIA COCCINEA B. Juss. [.S. coccinea ciliata Griseb.; .S. coccinea
pseudococcinea Kuntze.] Hillsides, waste and cultivated grounds,
St. Thomas; St. \Croix; St. Jan.
MENTHA AQuATICA L. Naturalized along rivulets, Caledonia, St.
Croix, not seen flowering (according to Eggers). Perhaps, if the
flowers were known, referable to some other species.
HyptTis cCAPITATA Jacq. [Mesosphaerum capitatum Kuntze.]}
Moist grounds, St. Thomas; St. Jan; St. Croix.
HypTiIs SUAVEOLENS (L.) Poir. [Ballota suaveolens L.; Meso-
sphaerum suaveolens (Kuntze.] Thickets, waste and cultivated
grounds, St. Thomas; St. Jan; St. Croix.
88 BROOKLYN BOTANIC GARDEN MEMOIRS
HypTis PECTINATA (L.) Poir. [Nepeta pectinata L.; Mesosphaerum
pectinatum WKuntze.] Waste and cultivated grounds, St. Thomas;
St. Jang St.cCroix.
HypTIs VERTICILLATA Jacq. [Mesosphaerum verticillatum Kuntze. |
St. Thomas (according to Bentham and to Grisebach).
COLEUS AMBOINICUS Lour. Banks and hillsides, naturalized, St.
Thomas; St. Jan (according to Eggers); St. Croix.
OcIMUM MICRANTHUM Willd. Banks, fields and hillsides; St.
Thomas; St: Jan St; Croix:
Ocimum BasiLicum L. Grown as a garden herb.
ROSMARINUS OFFICINALIS L. Grown as a garden herb.
THYMUS VULGARIS L. Grown as a garden herb.
ORIGANUM Majorana L. Grown as a garden herb.
SOLANACEAE
PHYSALIS ANGULATA L. [P. ramosissima Mill.; P. Linkiana
Griseb., not Nees.] Fields, banks, waste and cultivated grounds, St.
Thomas; St.-lan;) St. Crow:
PHYSALIS PUBESCENS L. [P. angulata dubia Kuntze.] Dry soil,
waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
PHYSALIS TURBINATA Medic. Dry soil, St. Jan; St. Croix.:
PHYSALIS EGcErRsiII O. E. Schulz. Water Island, St. Thomas.
Endemic. A species known only from the type specimen. We
searched Water Island for it in 1913, but could find nothing answering
the description.
PHYSALIS PERUVIANA L., recorded by Eggers as found in fields at
Rapoon, St. Thomas, prior to 1879, was, apparently, erroneously
determined (see Schulz in Urban, Symb. Ant. 6: 149).
CAPSICUM FRUTESCENS L. Roadsides and woods, St. Thomas; St.
Jan;*St2C toi:
CapsicuM BACCATUM L. [Capsicum annuum baccatum Kuntze.]
Thickets, banks and woods, St. Thomas; St. Jan; St. Croix (accord-
ing to Eggers).
CAPSICUM ANNUUM L. Cultivated for food.
CAPSICUM DULCE Dunal. Cultivated for food.
SOLANUM LANCEIFOLIUM Jacq. King’s Hill, St. Jan.
SOLANUM JAMAICENSE Mill. St. Thomas, collected by Richard
(according to Poiret).
SOLANUM NIGRUM L. [S. americanum Mill.; S. nigrum nodi-
florum A. Gray; S. nodiflorum Dunal, not Jacq.| Thickets, waste
and cultivated grounds, St. Thomas; St. Jan; St. Croix.
SOLANUM SEAFORTHIANUM Andr. Spontaneous after cultivation,
St. Thomas; St. Croix. Planted for ornament.
BRITTON: FLORA OF THE VIRGIN ISLANDS 89
SOLANUM VERBASCIFOLIUM L. Hillsides and thickets, St. Thomas;
Sean; St. Croix.
SOLANUM CONOCARPUM L. C. Rich. Coral Bay, St. Jan. En-
demic.
SOLANUM MUCRONATUM O. E. Schulz. St. Thomas; St. Jan;
otherwise known only from Porto Rico.
SOLANUM MAMMOsUM L. Waste grounds, St. Croix.
SOLANUM PERSICIFOLIUM Dunal. [S. persicifolium Bellor O. E.
Schulz; S. persicifolium parvifolium (Vahl) O. E. Schulz.] Hillsides
and thickets, St. Thomas; St. Jan; St. Croix.
SOLANUM RACEMOSUM LL. [S. tgnaeum L.; S. bahamense of Eggers.]
Whrckets, St. Thomas; St. Jan; St. Croix.
SOLANUM TORVUM Sw. Hillsides, woods and waste grounds, St.
Thomas; St. Jan; St. Croix.
SOLANUM POLYGAMUM Vahl. [S. tnclusum and S. inclusum albi-
florum of Eggers; S. polygamum thomae Kuntze; S. hirtum of Borgesen
& Paulsen.] Thickets, St. Thomas; St. Jan; St. Croix.
SOLANUM ACULEATISSIMUM Jacq. Naturalized by mules from
Montevideo at Frederiksted, St. Croix (according to Eggers).
SOLANUM TUBEROSUM L. Cultivated for food.
SOLANUM MELONGENA L. [S. insanum L.] Cultivated for food.
SOLANUM MACROCARPUM L. Cultivated on St. Croix (according
to Schulz).
SOLANUM PSEUDOCAPSICUM L. Cultivated for its fruit.
LycorErsicuM Lycopersictm (L.) Karst. [Solanum Lycopersi-
cum L.; Lycopersicum esculentum Mill.; L. cerasiforme Dunal.]
Spontaneous after cultivation for food, St. Thomas; St. Jan; St. Croix.
DATURA STRAMONIUM L. [Datura Tatula L.] Waste and culti-
vated grounds, St. Thomas; St. Jan; St. Croix.
DaturA METEL L. Waste and cultivated grounds, St. Thomas;
Srlan: St. Croix.
DATURA FASTUOSA L. Spontaneous after cultivation, St. Thomas;
Bieon;-ot. Croix.
DATURA SUAVEOLENS H. & B. Cultivated for ornament.
CESTRUM NOCTURNUM L. Forests, Rogiers and Joshee Gut, St.
Jan (according to Eggers). Planted on St. Croix.
CESTRUM LAURIFOLIUM L’Her. [C. diurnum of West and of
Eggers; C. laurifolium neglectum Kuntze.| Forests and thickets, St.
Thomas; St. Jan; St. Croix.
CESTRUM ALTERNIFOLIUM (Jacq.) O. E. Schulz. [Ixora alternifolia
Jacq.; Cestrum vespertinum L.] St. Thomas (according to O. E.
Schulz).
NIcoTIANA TABACUM L. Spontaneous after cultivation, St.
Thomas; St. Jan; St. Croix.
90 BROOKLYN BOTANIC GARDEN MEMOIRS
BRUNFELSIA AMERICANA L. [B. americana pubescens Griseb.]|
Thickets and hillsides, St. Thomas; St. Jan; St. Croix. . Sometimes
planted for ornament. .
Petunias are planted in gardens.
SCROPHULARIACEAE
MECARDONIA PROCUMBENS (Mill.) Small. [Evinus procumbens
Mill.; Herpestis chamaedryoides H.B.K.; Lindernia dianthera Sw.;
Monniera dianthera Millsp.| Wet grounds, St. Croix.
HERPESTIS STRICTA Schrad., accredited to St. Thomas by Bentham,
according to Eggers, is probably an error in record.
BramiA Monniera (L.) Drake. [Gratiola Monniera L.; Herpestis
Monniera H.B.K.; Monniera Monniera Britton; M. calycina Kuntze.]
Wet sandy or muddy situations, St. Thomas; St. Jan; St. Croix.
CAPRARIA BIFLORA L. [C. biflora pilosa of Eggers.] Fields,
banks, waste and cultivated grounds, St. Thomas; St. Jan; St. Croix.
VANDELLIA DIFFUSA L. St. Croix (according to Eggers).
ScopARIA DuLcIs L. [Capraria dulcis Kuntze.] Wet or moist
situations and in cultivated grounds, St. Thomas; St. Jan; St. Croix.
RUSSELLIA EQUISETIFORMIS Schl. & Cham. [R. juncea Zucc.|
Cultivated for ornament.
MAuRANDYA BARCLAYANA Lindl. Cultivated for ornament.
BIGNONIACEAE
MACcCRODISCUS LACTIFLORUS (Vahl) Bureau. [Bzignonia lactiflora _
Vahl; Distictis lactiflora DC.| Thickets and roadsides, St. Croix.
Cultivated on St. Thomas (according to Eggers).
CyDISTA AEQUINOCTIALIS (L.) Miers. [Bignonta aequinoctialis L.;
B. spectabilis Vahl.] Forests and thickets, St. Thomas; St. Jan; St.
Croix.
BatocypiA Uncuis (L.) Mart. [Bignonia Unguis L.] Forests,
st, Phomas:’St. Jan; St. Croix.
MacrOCcATALFA LONGISSIMA (Jacq.) Britton. St. Thomas (accord-
ing to Grisebach). The record is probably erroneous.
TABEBUIA HETEROPHYLLA (DC.) Britton. [Raputia (?) hetero-
phylla DC.; Tecoma Berteriit of Eggers, not DC.; Tecoma triphylla of
Kuntze; Tecoma Leucoxylon Mart.; Tecoma pentaphylla Leucoxylon
Kuntze.]| Dry thickets, especially near the coasts, St. Thomas; St.
Jan.
TABEBUIA PALLIDA Miers. [Bignonia pentaphylla L.; Tecoma
pentaphylla Juss., not Tabebuia pentaphylla Hemsl.; T. Leucoxylon
of Eggers.}] Forests and hills and much planted for shade, St. Thomas;
St. Jang StigUrar,
BRITTON: FLORA OF THE VIRGIN ISLANDS 91
TECOMA STANS (L.) Juss. [Bzignonia stans L.; Stenolobium stans
Seem.; Gelseminum stans Kuntze.| Thickets and hillsides, St. Thom-
as; St. Jan; St. Croix. Sometimes planted for ornament.
TECOMARIA CAPENSIS (Thunb.) Spach. [Bignonia capensis Thunb.;
Tecoma capensis Lindl.] Roadsides, St. Thomas. Planted for orna-
ment, ot. Thomas; St. Croix.
ENALLAGMA LATIFOLIA (Mill.) Small. [Crescentia latifolia Muill.;
Crescentia cucurbitina L.; C. cucurbitina heterophylla Kuntze; E.
cucurbitina Baill.| Forests near rivulets, St. Thomas; St. Jan; St.
Croix.
CRESCENTIA CUJETE L. Forests, hillsides and much planted for
weioit, ot. Lhomas; St. Jan; St. Croix.
CRESCENTIA LINEARIFOLIA Miers. Collected by Oersted on St.
Thomas (according to Miers); coastal hill, Lamosure, St. Jan.
PEDALIACEAE
SESAMUM ORIENTALE L._ [S. indicum L.] Spontaneous after culti-
vation, St. Thomas. Cultivated for its seeds.
MARTYNIACEAE
MARTYNIA ANNUA L. [M. diandra Glox.] Waste and cultivated
grounds, St. Thomas; St. Croix.
ACANTHACEAE
THUNBERGIA FRAGRANS Roxb. [T7. volubilis Pers.| Hedges and
thickets and along ditches, St. Thomas; St. Jan; St. Croix,
THUNBERGIA ALATA Bojer. Banks and waste grounds, St. Thomas.
St. Jan; St. Croix. Planted for ornament.
BLECHUM Browne! Juss. [B. Brownet subcordatum and (?) laxum
Kuntze.] Fields, banks, woods and thickets, St. Thomas; St. Jan;
Due Croix.
RUELLIA TUBEROSA L. [R. clandestina L.] Grassy situations, St.
/mmomas; ot. Jan; St. Croix.
RUELLIA COCCINEA (L.) Vahl. [Barleria coccinea L.; Stemona-
canthus coccineus Griseb.] Thickets, St. Thomas; St. Jan; St. Croix.
RUELLIA STREPENS L., recorded by de Candolle as found on St.
Croix by Isert, must be an error in locality; the specimen is preserved
in the Willdenow herbarium.
GERARDIA TUBEROSA L. [Stenandrium tuberosum Urban; _ S.
rupestre Nees.| Rocky woods and thickets, St. Thomas; St. Jan.
ANTHACANTHUS SPINOSUS (Jacq.) Nees. [Justicia spinosa Jacq.;
A. microphyllus and A. jamaicensis of Eggers; Jasminum coeruleum
a2 BROOKLYN BOTANIC GARDEN MEMOIRS
Kuntze.] Woods, hillsides and thickets, St. Thomas; St. Jan; St.
Croix.
ANTHACANTHUS ACICULARIS (Sw.) Nees, attributed by West and
by Lindau to St. Croix, is known to me only from Jamaica. [Justicia
acicularis Sw.]
ODONTONEMA NITIDUM (Jacq.) Kuntze. [Justicia nitida Jacq.;
Thyrsacanthus nitidus Nees.] St. Thomas and St. Croix, at least
formerly.
DREJERELLA MIRABILOIDES (Lam.) Lindau. [Justicia mirabiloides
Lam.; Beleperone nemorosa of Eggers.] Shaded situations, St.
Thomas: StoCroix
SECHIUM EDULE (Jacq.) Sw. Cultivated for its fruit.
CITRULLUS CITRULLUS (L.) Karst. [C. vulgaris Schrad.] Culti-
vated for its fruit.
CoccINIA CORDIFOLIA (L.) Cogn. [Cephalandra indica Naud.]
Cultivated; recorded by Eggers as naturalized in shaded valleys, St.
Croix,
LOBELIACEAE
ISOTOMA LONGIFLORA (L.) Presl. [Lobelia longiflora L.| Moist
banks, fields and hillsides, St. Thomas; St. Jan; St. Croix (according
to Eggers).
GOODENIACEAE
SCAEVOLA PiLumrierRit (L.) Vahl. [Lobelia Plumierti L.] Coastal
sands, St: Thomas; St.‘Croix.
CICHORIACEAE
SONCHUS OLERACEUS L. Waste and cultivated grounds, St.
Thomas: -St-jan;est. Grom.
LACTUCA INTYBACEA Jacq. [Brachyramphus intybaceus DC.]
Waste and cultivated grounds, St. Thomas; St, Jan; St. Croix.
LacTuCcA SATIVA L. Cultivated for salad.
AMBROSIACEAE
XANTHIUM LONGIROSTRE Wallr. [X. orientale of Schlechtendal:
X. macrocarpum of Eggers; X. strumarium of Millspaugh; X. echi-
BRITTON: FLORA OF THE VIRGIN ISLANDS 97
natum of Urban.] Waste and cultivated grounds, St. Thomas; all
islands (according to Eggers).
AMBROSIA CUMANENSIS H.B.K. [A. artemisiaefolia trinitensis
Griseb.] Waste places, St. Croix (according to Eggers).
CARDUACEAE
STRUCHIUM SPARGANOPHORUM (L.) Kuntze. [Ethulia spargano-
phora L.; Sparganophorus Vaillant Crantz.] Moist grounds, St.
Thomas (according to Eggers).
VERNONIA SERICEA L. C. Rich. ([Lepidoploa phyllostachya Cass.;
Vernonia arborescens Swartziana, Lessingiana and dwaricata of Eggers;
V. arborescens of Schlechtendal and of Millspaugh; V. phyllostachya
Gleason; Cacalia arborescens Lessingiana Kuntze.] Thickets, St.
imomas; St. Jan; St: Croix.
VERNONIA ALBICAULIS Pers. [V. longifolia Pers.; V. Vahhiana
Less.; V. thomae Benth.; V. punctata of Eggers and of Millspaugh;
Cacalia thomae Kuntze; ? Conyza fruticosa of West.] Thickets, St.
feremas- St. Jan; St. Croix:
VERNONIA CINEREA (L.) Less. [Conyza cinerea L.| Waste and
cultivated grounds, St. Thomas; St. Croix.
PIPTOCOMA RUFESCENS Cass. Thickets, Water Island, St. Thomas;
Stan.
ELEPHANTOPUS MOLLIS H.B.K. [E. tomentosus of Millspaugh.]
Banks, fields and hillsides, St. Thomas; St. Jan; St. Croix.
PSEUDELEPHANTOPUS SPICATUS (Juss.) Rohr. [Elephantopus spi-
catus Juss.; Distreptus spicatus Rohr.| Banks, fields, hillsides and
cultivated grounds, St. Thomas; St. Jan; St. Croix.
AGERATUM CONYZOIDES L. [Carelia conyzoides robusta Kuntze.]
Banks, fields and roadsides, St. Thomas; St. Jan; St. Croix.
EUPATORIUM MACROPHYLLUM L._ [Hebeclinium macrophyllum DC. |
Forests, St. Thomas; St. Croix (according to West and to Eggers).
EUPATORIUM ODORATUM L. [E. conyzoides Vahl.] Banks, hill-
sides and thickets, St. Thomas; St. Jan; St. Croix.
EUPATORIUM CUNEIFOLIUM Willd., cited by Eggers from DeCandolle
(Prodr. 5: 177) as from St. Thomas, was not from our island St.
Thomas.
EUPATORIUM ATRIPLICIFOLIUM Lam. [E. repandum Willd.; Erig-
eron atriplicifolium of Millspaugh.] Hillsides and coastal thickets,
St Lnomas; St. Jan; St. Croix.
EUPATORIUM SINUATUM Lam. [E. canescens Vahl.] Rocky thick-
ets, St. Thomas (according to DeCandolle); rocky hillsides, St. Jan;
mt, Croix.
8
98 BROOKLYN BOTANIC GARDEN MEMOIRS
EUPATORIUM TRIPLINERVE Vahl. [£.AyapanaVent.] Cultivated
on) St.sGron.
EUPATORIUM CAPILLIFOLIUM (Lam.) Small. [E. foeniculaceum
Willd.] Cultivated on St. Croix (according to Millspaugh).
MIKANIA CORDIFOLIA (L. f.) Willd. [Cacalia cordifolia L. f.;
? Eupatorium denticulatum of Schlechtendal; Mikania gonoclada DC.;
Willughbaea cordifolia Kuntze; W. gonoclada Millsp.] Thickets, St.
Thomas--St..jan; St. Croix:
ERIGERON CUNEIFOLIUS DC. Grassy places and banks on the
higher hills, St. Thomas; St. Jan.
ERIGERON SPATHULATUS Vahl. Grassy situations, St. Thomas;
St: Jans. St Crom.
LEPTILON PUSILLUM (Nutt.) Britton. [Evrigeron pusillum Nutt.;
Erigeron canadense of Schlechtendal and of Eggers; Leptilon canadense
of Millspaugh.] Grassy places, waste and cultivated grounds, St.
Thomas: St. Jans"St Croix.
LEPTILON LINIFOLIUM (Willd.) Small. [Evigeron linifolium Willd.;
Conyza ambigua DC.| Waste grounds, St. Thomas.
PLUCHEA PURPURASCENS (Sw.) DC. [Conyza purpurascens Sw.;
P. camphorata of Millspaugh.] Wet grounds, St. Thomas; St. Croix.
PLUCHEA ODORATA (L.) Cass. [Conyza odorata L.; C. carolinensis
Jacq.; P. odorata normalis Kuntze.] Thickets and hillsides and in
cultivated grounds, St. Thomas; St. Jan; St. Croix.
BACCHARIS DIOICA Vahl. [B. Vahlit DC.] Coastal rocks, St.
Croix.
EGLETES PROSTRATA (Sw.) Kuntze. [Matricaria prostrata Sw.;
Pyrethrum simplicifolium Willd.; EE. domingensis Cass.] Sandy
shores, St. Thomas.
PTEROCAULON VIRGATUM (L.) DC. [Guaphahum wirgatum L.;
Conyza virgata L.; Pluchea virgata Schl.] Hillsides and banks, St.
Thomas: St.-Jan> St. Crorx:
NoccA MOLLIS (Cav.) Jacq. ([Lagascea mollis Jacq.| Waste
grounds, St. Thomas.
MELAMPODIUM DIVARICATUM (L. C. Rich.) DC. [Dysodium divar-
icatum L. C. Rich.; M. paludosum H.B.K.] Ditches, St. Croix.
PARTHENIUM HysTEROPHORUS L. Waste and cultivated grounds,
St. Thomas; St. fans St: Croix.
CRASSINA MULTIFLORA (L.) Kuntze. [Zinnia multiflora L.]
Roadsides and banks, St. Thomas; St. Jan.
CRASSINA ELEGANS (Jacq.) Kuntze. Grown in flower gardens.
VERBESINA ALBA L. [Eclipta alba Hassk.; FE. punctata L.; £.
erecta L.] Wet grounds, St. Thomas; St. Jan; St.. Croix.
ACANTHOSPERMUM HISPIDUM DC. [A. humile of Eggers.] Waste
and cultivated grounds, St. Thomas; St. Croix.
BRITTON: FLORA OF THE VIRGIN ISLANDS 99
BORRICHIA ARBORESCENS (L.) DC. [Buphthalmum arborescens L.;
B. argentea DC.| Coastal rocks and sands, St. Thomas; St. Croix.
WEDELIA TRILOBATA (L.) Hitche. [Silphium trilobatum L.; W.
carnosa L. C. Rich.] Moist grounds, St. Thomas; St. Croix.
WEDELIA CALYCINA L. C. Rich. [Buphthalmum helianthoides of
West.| Thickets, St. Thomas; St. Croix.
WEDELIA PARVIFLORA L. C. Rich. [W. buphthalmoides of Eggers
and of Millspaugh; W. affinis DC.; W. acapulcensis of Schlechtendal ;
W. brachycarpa of Millspaugh, St. Thomas; Sereneum frutescens of
Kuntze.| Dry hills and thickets, St. Thomas; St. Jan.
WEDELIA CRUCIANA L. C. Rich. [W. buphthalmoides of Mill-
spaugh.] Dry rocky soil, St. Croix. Endemic.
ELEUTHERANTHERA RUDERALIS (Sw.) Sch. Bip. [Melampodium
ruderale Sw.; Ogiera ruderalis Griseb.; Wedelia discoidea Less.]
Banks, fields and waste grounds, St. Thomas; St. Jan; St. Croix.
MELANTHERA CANESCENS (Kuntze) O. E. Schulz. [Amellus asper
canescens Kuntze; M. Linnaei of Schlechtendal; MM. deltoidea of
Eggers.] Hillsides and thickets, St. Thomas.
TEPION ALATUM (L.) Britton. [Verbesina alata L.] Waste and
cultivated ground, St. Thomas; St. Croix.
SCLEROCARPUS AFRICANUS Jacq. Waste grounds, St. Thomas.
SYNEDRELLA NODIFLORA (L.) Gaertn. [Verbesina nodiflora L.;
Ucacou nodiflorum Hitche.] Waste and cultivated grounds, St.
Thomas; St. Jan; St. Croix.
BIDENS PILOSA L. [Coreopsis leucantha L.; B. leucantha Willd.;
B. pilosa dubia O. E. Schulz; ? B. pilosa subbiternata Kuntze.| Waste
and cultivated grounds, St. Thomas; St. Jan; St. Croix.
BIDENS CYNAPIIFOLIA H.B.K. [B. bipinnata of West, of Eggers
and of Millspaugh.| Waste and cultivated grounds, St. Thomas; St.
Wem; ot. Croix.
Cosmos CAUDATUS H.B.K. [Bidens Berteriana Spreng.| Grassy
fields, banks, and in waste grounds, St. Thomas; St. Jan; St. Croix.
POROPHYLLUM PoROPHYLLUM (L.) Kuntze. [Cacalia Porophyllum
L.; Porophyllum ellipticum Cass.| Waste grounds, St. Thomas.
PECTIS HUMIFUSA Sw. [P. serpyllifolia Pers.| Stony banks,
fields, and hillsides, St. Thomas; St. Jan; St. Croix.
PECTIS LINIFOLIA L. [Pectis punctata Jacq.; Pectidium punctatum
Less.| Rocky hillsides, banks and thickets, St. Thomas; St. Jan;
ar Croix.
PECTIS FEBRIFUGA H. van Hall. [P. Swartziana of Borgesen and
Paulsen.] Grassy places, St. Thomas; St. Croix.
NEUROLAENA LoBATA (L.) R. Br. [Conyza lobata L.] Woodlands,
St. Thomas.
100 BROOKLYN BOTANIC GARDEN MEMOIRS
ERECHTHITES HIERACIFOLIA (L.) Raf. [Senecio hieracifolius L.;
E. praealta Raf.; E. hieracifola cacalioides of Eggers and of Kuntze.]
Banks, fields, waste and cultivated grounds, St. Thomas; St. Croix.
EMILIA SONCHIFOLIA (L.) DC. [Cacalia sonchifolia L.; E. sonchi-
folia sagittata of Kuntze.] Banks, fields, waste and cultivated grounds,
St. Phomas; St. Croix; St. ant
EMILIA SAGITTATA (Vahl) DC. » Grown in flower gardens.
CHAPTALIA NUTANS (L.) Polak. [Tussilago nutans L.; Leria nutans
DC.] Woods and thickets, St. Thomas; St. Jan; St. Croix.
CHRYSOGANUM DICHOTOMUM Vahl, of St. Croix, is unknown to me;
it is certainly not a Chrysoganum.
HELIANTHUS ANNUUS L. Grown in gardens.
CHRYSANTHEMUM INDICUM L. [Pyrethrum indicum Cass.| Grown
in gardens.
ASTER CHINENSIS L. Grown in gardens.
TAGETES PATULA L. Grown in gardens.
TITHONIA TAGETIFLORA Desf. [T. speciosa Hook.] Grown in
gardens.
GEORGINA VARIABILIS Willd. Grown in gardens (according to
Eggers).
TARCHONANTHUS CAMPHORATUS L. Cultivated on St. Croix (ac-
cording to Eggers).
PINACEAE
THUJA ORIENTALIS L. Planted for ornament.
JUNIPERUS BERMUDIANA L. Planted on St. Croix (according to
West).
CYCADACEAE
CyCAS REVOLUTA Thunb. Planted for ornament.
PTERIDOPHYTA
CYATHEACEAE
CYATHEA ARBOREA (L.) J. E. Smith. [Polypodium arboreum L.;
? C. Serra of Kuhn.] Forests, high hills, St. Thomas.
POLY PODIACEAE
DRYOPTERIS PATENS (Sw.) Kuntze. [Polypodium patens Sw.;
Aspidium patens Sw.] Forests, high hills of St. Thomas; St. Croix.
DRYOPTERIS OLIGOPHYLLA Maxon. [Polypodium invisum Sw.]
St. Thomas (according to Christensen).
DRYOPTERIS INCISA (Sw.) Kuntze. [Polypodium incisum Sw.]
St. Croix (West, according to Eggers).
BRITTON: FLORA OF THE VIRGIN ISLANDS 101
DRYOPTERIS SPRENGELU (Kaulf.) Kuntze. [Dryoteris Balbisii
Urban; Polypodium Balbistt Spreng.] St. Thomas (according to
Kuhn).
DRYOPTERIS SERRA (Sw.) Kuntze. [Polypodium serra Sw.;
Tectaria incisa Cav.; Dryopteris serra incisa Kuhn.] St. Thomas
(according to Kuhn).
DRYOPTERIS MOLLIS (Jacq.) Hieron. [Aspidium molle Sw.]
Forests, Signal Hill, St. Thomas; St. Jan; St. Croix (according to
Millspaugh).
DRYOPTERIS TETRAGONA (Sw.) Urban. [Polypodium tetragonum
Sw.] Forests, St. Thomas; St. Jan; St. Croix. ;
Dryoptertis PorrEANA (Bory) Urban. | Polypodium crenatum Sw.,
not Forst; Lastrea Poiteana Bory.] St. Thomas and St. Croix (ac-
cording to Eggers); St. Croix (according to West).
CYCLOPELTIS SEMICORDATA (Sw.) J. Smith. [Polypodium semi-
cordatum Sw.; Aspidium semicordatum Sw.| Shaded localities,
Virgin Islands (according to Eggers).
GYMNOPTERIS NICOTIANIFOLIA (Sw.) Presl. [Acrostichum nicoti-
anum Sw.], attributed by Swartz to St. Thomas, was probably from
Porto Rico.
NEPHROLEPIS EXALTATA (L.) Schott. [Polypodium exaltatum L.]
Forests, St. Thomas; St. Croix. Cultivated on St. Croix (according
to Millspaugh).
NEPHROLEPIS RIVULARIS (Vahl) Mett. [Polypodium rivulare
Vahl.| St. Thomas (according to Kuhn).
NEPHROLEPIS BISERRATA (Sw.) Schott. [Aspidium biserratum Sw.;
Aspidium acutum Schk.; Nephrolepis acuta Presl.; Aspidium punctu-
latum Sw.| Forests, St. Thomas (according to Eggers); St. Croix.
ODONTOSORIA ACULEATA (L.) J. Smith. [Adiantum aculeatum L.;
Davallia aculeata J. E. Smith.] Pastures on high hills, St. Thomas.
ODONTOSORIA CLAVATA (L.) J. Smith, is doubtfully attributed to
St. Thomas by Fée.
ASPLENIUM SERRATUM L. Forests, Signal Hill, St. Thomas.
ASPLENIUM PUMILUM Sw. Forests and wet banks, St. Thomas;
St. Jan.
ASPLENIUM ABsciIssuM Willd. [A. firmum Kunze.] St. Thomas
(according to Grisebach).
BLECHNUM OCCIDENTALE L. Banks, fields and forests, St. Thomas;
et. jan; ot. Croix.
PITYOGRAMMA SULPHUREA (Sw.) Maxon. [Gymnogramme sulphurea
Desv.]. Cultivated in gardens.
PITYOGRAMMA CALOMELANA (L.) Link. [Acrostichum calomelanos
L.; Gymnogramme calomelanos Kaulf.; G. calomelanos pumila Eggers. ]
Banks, hills, walls and thickets, St. Thomas; St. Jan; St. Croix.
102 BROOKLYN BOTANIC GARDEN MEMOIRS
HEMIONITIS PALMATA L. Forests, wet banks and rocky thickets,
St. Thomas; Sts Jan} St. Cron,
DORYOPTERIS PEDATA (L.) Fée. [Pteris pedata L.] Forests
and shaded banks, St. Thomas; St. Jan.
CHEILANTHES MICROPHYLLA Sw. [Adiantum muicrophyllum Sw.]
Rocky slopes, St. Thomas; St. Croix (according to West and to
Eggers).
ADIANTUM VILLosUM L. Forests, St. Thomas; St. Croix.
ADIANTUM LATIFOLIUM Lam. [A. denticulatum Sw.; A. tnter-
medium of Eggers; A. obliquum intermedium of Millspaugh.] Shaded
banks, hills of St. Thomas.
ADIANTUM CRISTATUM L. [?A. microphyllum of Eggers.] Hill-
sides, St. Thomas.
ADIANTUM TENERUM Sw. Shaded banks and ravines on high hills,
St. Thomas; St. Jan (according to Eggers); St. Croix. .
ADIANTUM FRAGILE Sw. Thickets and walls, St. Thomas; St. Jan;
St. Croix:
ADIANTUM FARLEYENSE Moore. Cultivated on St. Croix (A.
foliosum of Millspaugh).
PYCNODORIA LONGIFOLIA (L.) Britton. [Pteris longifolia L.| Along
rivulets in forests, St. Croix; in a water spout, St. Thomas.
PrERIS BIAURITA L. St. Thomas (according to Kuhn).
ANTROPHYUM LINEATUM (Sw.) Kaulf. [Hemionitis lineata Sw.]|
Forest, St. Peter, St. Thomas (according to Eggers).
PALTONIUM LANCEOLATUM (L.) Presl. [Pteris lanceolata L.;
Taenitis lanceolata R. Br.; Heteropteris lanceolata Fée.| On rocks
and trees in forests, St. Thomas; all islands (according to Eggers).
POLYPODIUM POLYPODIOIDES (L.) Hitche. [Acrostichum poly-
podioides L.; P. incanum Sw.| On trees, St. Thomas; St. Jan; St.
Croix (according to Eggers).
PHLEBODIUM AUREUM (L.) J. Sm. [Polypodium aureum L.| On
trees and rocks, St. Thomas; St. Jan; St. Croix.
PHLEBODIUM AREOLATUM (H.&B.).J.Sm. [Polypodium areolatum
H. & B.] On trees, St. Thomas; St. Jan.
LEPICYSTIS PILOSELLOIDES (L.) Diels. [Polypodium piloselloides
L.] In forests among rocks, Signal Hill, St. Thomas.
PHYMATODES EXIGUUM (Hew.) Underw. [Polypodium exiguum
Hew.; P. serpens Sw., not Forst.; P. Swartzit Baker.] On trees,
Bordeaux, St. Jan; St. Croix.
CAMPYLONEURUM PHYLLITIDIs (L.) Presl. [Polypodium Phyllitidis
L.; P. Phyllitidis repens of Eggers.] On rocks and trees in forests,
St. Thomas; St. Jan; St. Croix.
CAMPYLONEURUM LATUM Moore. Shaded rocks, Bethania, St. Jan.
BRITTON: FLORA OF THE VIRGIN ISLANDS 103
ACROSTICHUM AUREUM L. [Chrysodium vulgare Fée.] Borders of
marshes, St. Thomas; St. Croix.
OPHIOGLOSSACEAE
OPHIOGLOSSUM RETICULATUM L. Grassy places among rocks,
Crown, St. Thomas; shaded bank, Bordeaux, St. Jan.
LYCOPODIACEAE
LycopoDIUM CERNUUM L. Among rocks in higher hills, St.
Thomas.
PSILOTACEAE
PsILotuM NupDuM (L.) Griseb. [Lycopodium nudum L.; P. tri-
quetrum Sw.| Shaded places among rocks, Signal Hill, St. Thomas;
Bordeaux, St. Jan; Crequis, St. Croix.
BRYOPHYTA
MUSCI*
DICRANELLA LONGIROSTRIS (Schwaegr.) Mitten. [Zvrematodon
longirostris Schwaegr.| St. Jan.
LEUCOLOMA SERRULATUM Bridel. [Z. Riedlei Besch.] On trees
in wet woods, St. Thomas.
OCTOBLEPHARUM ALBIDUM (L.) Hedw. [Bryum albidum L.] On
roots of Anthurium, near Caret Bay, St. Thomas; St. Croix.
FISSIDENS KEGELIANUS C. Muell. [F. palmatus—of various
authors, not Swartz.]| On banks, St. Thomas; St. Jan; St. Croix.
FISSIDENS ELEGANS Bridel. On rocks and earth, St. Thomas;
Sf. Jan.
SYRRHOPODON FLAVESCENS C. Muell. On rotten wood, St. Jan.
CALYMPERES RICHARDI C. Muell. [C. Breutelia Besch.; C. hexa-
gonum Besch.] On rocks and banks, St. Thomas.
HYMENOSTOMUM BREUTELII (C. Muell.) Broth. [Weitsia Breutelii
C. Muell.; Gymnostomum Breutelii Br. & Sch.| On _ banks, St.
Thomas; St. Jan; St. Croix.
HyopuiLa TortTuLa (Schwaegr.) Hampe. [Gymnostomum Tortula
Schwaegr.] St. Croix.
BARBULA AGRARIA (Sw.) Hedw. [Bryum agrarium Sw.] On
rocks, walls and earth, St. Thomas; St. Jan; St. Croix.
BARBULA CRUEGERI Lond. [Hyophila uliginosa E. G. Britton.]
Bethania, St. Jan.
PHASCUM SESSILE E. G. Britton. On the ground, Cowell Point
- and Water Island, St. Thomas.
4 Contributed by Elizabeth G. Britton.
104 BROOKLYN BOTANIC GARDEN MEMOIRS
BRYUM CRUEGERI Hampe. Bed of stream, Tutu, St. Thomas.
Sterile.
PHILONOTIS SPHAEROCARPA (Sw.) Bridel. [Mnium sphaericarpum
Sw.] Moist banks, high hills of St. Thomas.
PHILONOTIS TENELLA (C. Muell.) Jaeger. [Bartramia tenella C.
Muell.] Wet banks, St. Jan.
PIREELLA CYMBIFOLIA (Sull.) Cardot. [Pzlotrichum cymbifolium
Sull.] On trees near Bethania, St. Jan.
NECKERA DISTICHA (Sw.) Hedw. [Fontinalis disticha Sw.| On
trees, rarely on rocks, St. Peter, St. Thomas.
NECKERA JAMAICENSIS (Gmel.) E. G. Britton. [Hypnum jamat-
cense Gmel.| On trees, Bethania, St. Jan.
CALLICOSTELLA BELANGERIANA (Besch.) Jaeger. [Hookeria Bel-
angeriana Besch.| On stones, Bordeaux, St. Jan.
STEREOPHYLLUM LEUCOSTEGUM (Bridel) Mitten. [Leskea leuco-
stega Bridel.}| On wet or shaded rocks, St. Thomas; St. Jan.
MITTENOTHAMNIUM DIMINUTIVUM (Hampe) E.G. Britton. [Hyp-
num diminutivum Hampe.] On old wood, Bordeaux, St. Jan.
TAXITHELIUM PLANUM (Bridel) Mitten. [Hypnum planum Bridel.]
Wet rocks, logs and tree-roots, St. Thomas; St. Jan.
SEMATOPHYLLUM ADMISTUM (Sull.) Mitten. [Hypnum admistum
Sull.]|| Shaded banks, stones and dead wood, St. Thomas; St. Jan.
HAPLOCLADIUM MICROPHYLLUM (Sw.) Broth. [Hypnum miucro-
“phyllum Sw.] Shaded bank between Pearl and Bonne Resolution,
St. Thomas.
THUIDIUM CYMBIFOLIUM (Dz. & Mk.) Br. Jav. Shaded banks, St.
Thomas.
DENDROPOGON RUFESCENS Schimp., a Mexican species, has been
credited to St. Thomas in Paris, Index, and accepted by Brotherus,
but we have seen no specimens.
HEPATICAE OF ST. CROIX, ST. JAN, ST. THOMAS AND
TORTOLA®
In the Synopsis Hepaticarum of Gottsche, Lindenberg and Nees
von Esenbeck, published in 1844-47, three species of Hepaticae are
listed from St. Croix, one from St. Jan, and one from St. Thomas.
Another species, although listed from St. Kitts, was based in all prob-
ability on material from St. Jan. A seventh species has been listed
from St. Jan by Stephani. These seven species, which are the only
ones so far reported from the islands under discussion, deserve a few
words of comment.
5 Contributed by Alexander W. Evans, Yale University.
>
BRITTON: FLORA OF THE VIRGIN ISLANDS 105
The first species, Radula pallens (Sw.) Dumort., is said to have
been found “‘in St. Crucis insula,’ the record being based on a speci-
men in the Weber herbarium. This specimen was originally referred
to Jungermannia complanata L. (Radula complanata Dumort.) by
Weber,® but the later determination is probably correct.
The second species, Lejeunea Montagnei Gottsche, was based on
material from the Mascarene Islands and is now regarded as a species
of Euosmolejeunea. A specimen from St. Croix is listed in the. Synopsis
but is very problematical and would probably now be referred to
some other species. Since the specimen in question has not been
available for study, and since no later references to it are to be found
in the literature, its status must be left in doubt.
The third species, Lejeunea bethanica Gottsche, is based on material
collected by Breutel and is said to have come from “prope Bethaniam
in Insula St. Christopheri.’”” Many years later Stephani,’ on the basis
of a specimen in the Lindenberg herbarium at Vienna, quoted the
species from St. Jan, referring it to the subgenus Cheilo-Lejeunea.
Still later he apparently changed his ideas regarding the habitat of
the plant, citing it from St. Kitts and redescribing it under the name
Cheilolejeunea bethanica Steph.’ In studying the Lejeuneae in the
Lindenberg herbarium, the writer found two specimens labeled
Lejeunea bethanica, both of which were collected by Breutel at Be-
thania, St. Jan. One of these is very fragmentary but is apparently
referable to Rectolejeunea phyllobola (Nees & Mont.) Evans; the other,
which is the specimen studied by Stephani, is (in the writer’s opinion)
referable to Lejeunea rather than to Cheilolejeunea. The species was
originally described from a specimen in the Gottsche herbarium at
Berlin, not available at the present time, and there is therefore a
possibility that the actual type may have come from St. Kitts. The
evidence, however, is against this view, and it seems permissible to
assume that the specimen in the Lindenberg herbarium is identical
with the type and that it formed a part of the same collection. Un-
fortunately L. bethanica has not again been collected on either St.
Jan or St. Kitts.
The fourth species, Lejeunea epiphyta Gottsche, was described as
“parasitans in Lej. bethanica in Insula St. Johannis prope Bethaniam
(Breutel, Hb. G.).’’ This statement affords further proof that L.
bethanica came from St. Jan. According to Stephani® the specimen
of L. epiphyta in the Lindenberg herbarium should be referred to
6 Prodr. Hist. Musc. Hepat. 59. 1815.
7 Hedwigia 29: 86. 1890.
8 Sp. Hepat. 5: 652. 1914.
° Hedwigia 29: 90. 1890.
106 BROOKLYN BOTANIC GARDEN MEMOIRS
Lejeunea myriocarpa Nees & Mont., now Cololejeunea myriocarpa
Evans.
The fifth species, Anthoceros Breutelii Gottsche, was said to have
been collected near Friedensthal, St. Croix. This species, in 1858,
was transferred by its author'® to the genus Notothylas, where it is
still retained. The following year Milde showed that the type
material of the species did not come from St. Croix but from the
Corallberg, St. Jan. There is likewise a specimen from St. Jan in
the Mitten herbarium, which is presumably a part of the original
collection. ;
The sixth species, Lejeunea linguaefolia Tayl., was found ‘“‘in
Insula St. Thomas (Richard in Hb. Hk. a. 1814).’’ A specimen of
this species in the Lindenberg herbarium is referred by Stephani” to
Brachiolejeunea corticalis (Lehm. & Lindenb.) Schiffn., and the writer
would make the same disposition of a specimen in the Mitten her-
barium.!? Recently, however, Stephani has apparently thrown doubt
on the propriety of this reduction. In the fifth volume of his Species
Hepaticarum (1912), on page 35, he includes L. linguaefolia among the
species of Ptychocoleus, citing it as Pt. linguaefolius Steph., and adds
that he has been unable to see the plant and that his diagnosis is
simply a translation of the original description. On page 127, never-
theless, he again quotes L. linguaefolia among the synonyms of B.
corticalis. Since Stephani’s original reduction was based on the study
of an actual specimen, the writer would regard Pt. linguaefolius as
nothing more than an unnecessary synonym.
The seventh and last species, Riccia Breutelit Hampe, is described
as new by Stephani in the first volume of his Species Hepaticarum
(1898), on page 17, the habitat being given as “‘ Insulae S’Kitts et S.
Juan.”’ Dr. Howe informs the writer that there is some question
about the identity of the St. Kitts and St. Jan plants and it is there-
fore omitted from the following list, pending investigation of authentic
material. ;
In February, 1913, a botanical exploration of the islands was
carried on under the auspices of the New York Botanical Garden, the
Carnegie Institution of Washington and the United States National
Museum.'* The Hepaticae collected by the various members of this
expedition form the basis for the present report. The specimens
from St. Thomas, unless otherwise noted, were collected by Mrs.
Elizabeth G. Britton and Miss Delia W. Marble, those from St. Jan
10 Bot. Zeit. 16 (Anhang): 21. 1858.
11 Bot. Zeit. 17:'50. 1859.
Hedwigia 29: 22. 1890.
18 See Bull. Torrey Club 35:.164. 1908.
44 See Britton, N. L., Jour. N. Y. Bot. Gard. 14: 99. 1913.
BRITTON: FLORA OF THE VIRGIN ISLANDS 107
and Tortola by N. L. Britton and J. A. Shafer, and those from St.
Croix by J. N. Rose. The report records also two specimens collected
by C. H. Ostenfeld in 1914, one on St. Thomas and one on St. Jan.
1. RicciA Britton M. A. Howe.!5
St. THOMAS: on the ground, Water Island, N. L. Britton, E. G.
Britton & J. A. Shafer 148 (a much larger plant than the original).
2. PLAGIOCHILA LUDOVICIANA Sulliv.
St. JAN: on wet rocks, Bethania, 360. St. THOMAS: on rocks,
st. Peter, 7453.
3. RADULA PALLENS (Sw.) Dumort.
Sr. Croix: without definite locality, collector unknown, cited in
the Synopsis Hepaticarum.
4. COLOLEJEUNEA MYRIOCARPA (Nees & Mont.) Evans.
Lejeunea epiphyta Gottsche.
St. Croix: without definite locality. St. JAN: near Bethania,
J.C. Breutel. St. THOMAS: on rotten wood, near Magin’s Bay, 1317.
TorRTOLA: on a rock, Road Town to High Bush, 325 m. alt., 786 (a
trace only). |
5. LEJEUNEA BETHANICA Gottsche.
Cheilolejeunea bethanica Steph.
St. JAN: near Bethania, J. C. Breutel.
6. LEJEUNEA GLAUCESCENS Gottsche.
St. THOMAS: on tree roots, Bonne Resolution, 447.
7. LEJEUNEA MINUTILOBA Evans.
ST. THOMAS: St. Peter, 1251, 1254, 1255; on stones, Crown, 1365
(type).
8. LEJEUNEA PILILOBA Spruce.
ST. Crorx: on bark of a tree, without definite locality. Sr. JAN:
at base of a tree, Rosenberg, 300 m. alt., 306; on a shaded rock,
Bordeaux, 350 m. alt., 577. St. THOMAS: on ridge north of Charlotte
Amalia, 406; Crown, 1365 (a trace only); on rotten wood, St. Peter,
1451.
g. MICROLEJEUNEA LAETEVIRENS (Nees & Mont.) Evans.
St. THOMAS: on Anthurium roots, Pearl to Bonne Resolution,
1340; on fern roots, St. Peter, 1253; on mountain behind Charlotte
Amalia, C. H. Ostenfeld 77. TorTOLA: on a rock, Roadtown to
High Bush, 325 m. alt., 786 (in part).
10. RECTOLEJEUNEA PHYLLOBOLA (Nees & Mont.) Evans.
St. JAN: near Bethania, J. C. Breutel (specimen in the Lindenberg
‘8 Determined by Marshall A. Howe.
108 BROOKLYN BOTANIC GARDEN MEMOIRS
herbarium, labeled L. bethanica); on Clusia roots, Bethania, 355. ST.
THOMAS: on roots of royal palm, Tutu, 423. ToRTOLA: on a rock,
Roadtown to High Bush, 325 m. alt., 786 (in part); shaded rocks,
High Bush, 375 m. alt., 875.
11. EUOSMOLEJEUNEA CLAUSA (Nees & Mont.) Evans.
St. THomas:. shaded bank, Pearl to Bonne Resolution, 7339; on
the ground, St. Peter, 1455.
12. EUOSMOLEJEUNEA DURIUSCULA (Nees) Evans.
St. JAN: shaded rocks, Bordeaux, 330 m. alt., 570. St. THOMAS:
on rocks, St. Peter, 1452; on rocks, Crown, 1454.
13. EUOSMOLEJEUNEA TRIFARIA (Nees) Schiffn.
St. THOMAS: on rocks, St. Peter, 1252; on a rotten log, Crown,
450 m. alt., 1367.
14. TAXILEJEUNEA OBTUSANGULA (Spruce) Evans.
Sr. JAN: on a stone, Bordeaux, 400 m. alt., 582.
15. MASTIGOLEJEUNEA AURICULATA (Wils. & Hook.) Schiffn.
St. JAN: on a tree, Bethania to Rosenberg, 243; on a wet rock,
Bethania 356, 362; on loose blocks, Little Cruz Bay, C. H. Ostenfeld
391. ST. THOMAS: on a rock, St. Peter, 1256; on rocks in ravine
below Tutu, 1290.
16. BRACHIOLEJEUNEA CORTICALIS (Lehm. & Lindenb.) Schiffn.
Lejeunea linguaefolia Tayl.
Ptychocoleus linguaefolius Steph.
St. THOMAS: without definite locality, L. C. Richard.
17. FRULLANIA BRASILIENSIS Raddi.
St. JAN: on wet rocks. Bethania, 363 (mostly, specimens with-
out perianths and therefore somewhat doubtful).
18. FRULLANIA KUNZEI Lehm. & Lindenb.
Sr. JAN: on wet rocks, Bethania, 363 (a trace only); on rocks,
Bordeaux, 300 m. alt., 550.
I9. FRULLANIA SQUARROSA (R. BI. & N.) Dumort.
St. THomAs: on rocks, ridge north of Charlotte Amalia, 407;
on rocks, Bonne Resolution, 440; on rocks, Magin’s Bay to Mafolie,
1313; on rocks, Pearl to Bonne Resolution, 7338; on a stone wall,
Crown, 450 m. alt., 1364.
20. NoroTHyLaAs BREUTELII Gottsche.
Anthoceros Breutelii Gottsche.
Sr. JAN: Corallberg, J. C. Breutel (type); without defiriite locality
or collector’s name (specimen in Mitten herbarium); Bordeaux, 528. -
St. THoMAS: on damp earth, Nisky, NV. L. Britton, E. G. Britton &
BRITTON: FLORA OF THE VIRGIN ISLANDS 109
D. W. Marble 76; hills north of Charlotte Amalia, 409; on the ground,
Magin’s Bay to Mafolie, 1312.
21. ANTHOCEROS PUNCTATUS L.
St. THOMAs: in a mud hole, Crown, 450 m. alt., 7363.
REPORT ON THE LICHENS OF ST. THOMAS AND
ST. JAN’
The following report is based primarily upon a study of 90 numbers
of lichens from St. Thomas and St. Jan Islands; from St. Thomas,
82 numbers collected by Dr. N. L. Britton, Mrs. E. G. Britton, and
Miss Delia W. Marble, unless otherwise stated; from St. John, 8
numbers collected by Dr. N. L. Britton and Dr. J. A. Shafer. To
these have been added a few species recorded by Nylander in Flora
(63: 127. 1880) and two endemic species described by Mueller-
Argau. Altogether, we have represented 30 genera with 69 species
and varieties, of which three species and one variety are here de-
scribed as new. The discussion of the characteristics of the lichen-
flora will be confined to that of St. Thomas.
There is no peculiar element in the lichen-flora of St. Thomas
comparable with the gelatinous rock-lichens of Mona Island (see
Annals Missouri Bot. Gard. 2: 35. 1915), or with the crustose and
gelatinous rock-lichens of Bermuda (see Bull. Torrey Bot. Club 43:
146-155. 1916). Such rock-lichens as occur on St. Thomas belong
to widely distributed groups even where the actual species are more
restricted in distribution. In fact the greater part of the lichen-
flora is made up of species more or less common throughout the West
Indies or even in the tropics of both hemispheres. It is probably safe
to say that there are not more than half a dozen endemic species in
St. Thomas. Of the three new species described in this paper, two
occur elsewhere in the West Indies. The flora is comparatively rich
in the variety of crustose bark-lichens, especially in the genera An-
thracothecium with 5 species, Arthonia with 8 species, and Opegrapha
with 5 species. It is rather surprising that only one Graphis and no
Trypethelium should have been collected. On the whole the lichen-
flora is rather commonplace. In the following list, stations outside
of the Virgin Islands are noted for species, the range of which is
limited or not well known.
LICHENS OF ST. THOMAS
I. DERMATOCARPON HEPATICUM (Ach.) Th. Fr. On soil, without
definite locality, 151.
16 Contributed by Lincoln W. Riddle, Wellesley College.
110 BROOKLYN BOTANIC GARDEN MEMOIRS
2. LEPTORAPHIS EPIDERMIDIS (Ach.) Th. Fr. On bark, Bordeaux,
1381.
3. PORINA DESQUAMESCENS Fée. Without data, 1288a.
4. PoRINA NUCULA Ach. On Erythrina, St. Peter, 1444b, 1445.
5. PYRENULA LEUCOPLACA (Wallr.) Koerb. On bark, Smith’s Bay,
1276a.
6. ANTHRACOTHECIUM BREUTELII Muell. Arg. Flora 68: 339. 1885.
On bark, without definite locality, collected by Breutel. En-
demic.
7. ANTHRACOTHECIUM CANELLAE-ALBAE (Fée) Muell. Arg. On
Cephalocereus, Smith’s Bay, 1274a; on twigs of Guettarda,
Crown, 450 m. altit., 1357. South America.
8. ANTHRACOTHECIUM LIBRICOLUM (Fée) Muell: Arg. On Melicocca,
Tutu, 468a; on same, Smith’s Bay, 1276, 1282; on Erythrina,
St.. Meter, 1444
9. ANTHRACOTHECIUM OCHRACEOFLAVUM (Nyl].) Muell. Arg.
On bark of coconut palm, near Charlotte Amalia, 489; on Trichtlia,
Tutu, 467.
10. ANTHRACOTHECIUM PYRENULOIDES (Mont.) Muell. Arg. On bark,
without definite locality, collected by Dr. Forel. Recorded
by Nylander in Flora 63: 127. 1880.
11. MELANOTHECA FOVEOLATA Muell. Arg. On bark, near Tutu, 1287.
Cuba.
12. PARATHELIUM INDUTUM Nyl. On Cephalocereus, Smith’s Bay,
1274; on bark, Bordeaux, 1382 in part. Porto Rico, Colombia.
13. PARMENTARIA ASTROIDEA Fée. On bark, Bordeaux, 1382 in part.
14. ARTHONIA ANEGADENSIS Riddle Mem. New York Bot. Gard. 6:
579. 1916. On Beontia, Smith’s Bay, 1286. Anegada.
15. ARTHONIA CONFERTA (Fée) Nyl. On Plumeria, Tutu, 1484.
16. ARTHONIA GREGARIA (Weig.) Koerb. On bark, Bordeaux, 1380.
17. ARTHONIA OCHRACEELLA Nyl. On bark of old fallen branch,
Crown, 450 m. altit., 1362. Cuba.
18. ARTHONIA RUBELLA (Fée) Nyl. On Melicocca, Tutu, 468b.
19. ARTHONIA SUBRUBELLA Nyl. Collected by Dr. Forel. Recorded
by Nylander in Flora 63: 127. 1880.
20. ARTHONIA.” On Guilandina, Smith’s Bay, 1281 p.p.
21. ARTHONIA. On Plumeria, Tutu, 463.
22. ARTHOTHELIUM MACROTHECUM (Fée) Mass. On mango, St. Peter,
1247.
23. Opegrapha acicularis Riddle sp. nov.
Thallus epiphloeodes crustaceus effusus determinatus nigrolimitatus, albidus
1 There are included in this paper three species of Arihonia the determination
of which it has not been possible to complete in the time at my disposal. L. W. R.
BRITTON: FLORA OF THE VIRGIN ISLANDS 111
vel albo-cinerascens, tenuis laevis subcontinuus. Apothecia sessilia nuda ad 0.8
mm. lata, primum orbicularia suburceolata, margine crenato, dein orbicularia
oblonga vel subdifformia, disco late aperto plano rugoso atrofusco, margine laciniato
partim stellato-radiato nitido nigro; epithecio fusco; hymenio incolore, 120-130 u
altit.; hypothecio incolore; amphithecio sub lamina deficiente, cetero crasso nigro.
Paraphyses firmae crassae ramosae haud connexae. Asci cylindrices, 8-spori.
Sporae incolores aciculares rectae vel subflexuosae vel rarius contortae, 20-24-
loculares, cellulis cylindricis, 60-100 x 2-3 up.
Fic. 1. Opegrapha acicularis Riddle. Vertical section of apothecium (stippling
indicates dark coloration); paraphysis; ascus with two of the eight spores.
On bark of Erythroxylon, Punta Aguila, Porto Rico, collected by
N. L. Britton, J. F. Cowell, and Stewardson Brown, Feb. 27, 1915,
no. 4682 (type!). Also, on Guilandina, Smith’s Bay, St. Thomas,
1276a, 1279; and on Coccolobis, Great Harbor Cay, Berry Islands,
Bahamas, N. L. Britton and C. F. Millspaugh, no. 2545.
This is a striking and distinct species belonging to the section
Pleurothecium of the genus Opegrapha. Externally it is easily recog-
nizable by the marked tendency for the laciniate margin of the apo-
thecia to spread out in a stellate manner. The spores resemble those
of Lecanactis myriadea (Fée) Zahlbr. and of Opegrapha pleistophrag-
moides Nyl. But both of the species named have the black amphi-
thecium complete at the base, and the spores are almost twice as
thick.
24. OPEGRAPHA AGELAEA Fée. On Crescentia, Tutu, 462. Cuba,
Colombia.
25. OPEGRAPHA ATRA Pers. On Guilandina, Smith’s Bay, 1278a; on
Melicocca, Smith’s Bay, 1285.
112 BROOKLYN BOTANIC GARDEN MEMOIRS
26. OPEGRAPHA BONPLANDI Fée. On Melicocca, Magen’s Bay, 1310;
on bark, Mariendahl Road, 1476.
27. OPEGRAPHA VULGATA Ach. On Melicocca, Tutu, 468.
28. GRAPHIS SCRIPTA (L.) Ach. On bark, Smith’s Bay, 1277.
29. PHAEOGRAPHIS INUSTA (Ach.) Muell. Arg. On Acacia, near
Charlotte Amalia, 486; on Guilandina, Smith’s Bay, 1278;
without locality, 464.
30. CHIODECTON (Sect. Enterographa) sp. On Frythrina, St. Peter,
144 5a.
31. GYROSTOMUM SCYPHULIFERUM (Ach.) Fr. On Plumeria, Tutu,
465; on Acacia, near Charlotte Amalia, 487, 494.
32. BILIMBIA CUPREA Massal. in Lotos (1856) 77.
Lecidea cupreorosella Nyl. Mem. Soc. Sci. Nat. Cherb. 5: 122.
1857.
Biatora cupreorosella Tuck. Syn. N. A. Lich. 2: 34. 1888.
On old brick, St. Peter, 1442. Eastern United States, Europe.
33. BACIDIA ALBESCENS (Arn.) Zwackn. On Erythrina,*St. Peter,’
1445b.
34. CLADONIA PITYREA f. SQUAMULIFERA Wainio. On rocks, slope of
Crown, 1440.
35. LEPTOGIUM CHLOROMELUM (Sw.) Nyl. On bark, Cowell Point,
103, 172; on Pisonia roots, Water Island, 153.
36. LEPTOGIUM TREMELLOIDES (L. f.) S. F. Gray. On _ tree-trunk,
Crown, 1361.
37. LEPTOGIUM TREMELLOIDES var. CAESIUM (Ach.) Hue. On rock
near Bonne Resolution, 447.
38. LECANORA CINEREOCARNEA (Eschw.) Wainio. Without data, 23a;
on Guilandina, Smith’s Bay, 1281.
39. LECANORA GRANIFERA Ach. On bark, Mariendahl Road, 1476a.
40. Lecania euthallina Riddle sp. nov.
Thallus crustaceus uniformis effusus crassus rimoso-areolatus, areolis 0.2-0.4
mm. latis leviter convexis contiguis, cinereus vel sat pallide fuscescens; hypothallo
nullo. Gonidia cystococcoidea. Apothecia 0.6 mm. (0.4-1.0 mm.) lata, numerosa
partim caespitosa superficialia sat elevata regularia, disco concavo castaneo vel
fusco-nigricante nudo, margine proprio tenue disco concolore, margine thallino
integro vel demum crenulato crasso prominente thallo concolore; epithecio fulvo;
hymenio et hypothecio incolore. Asci 8-spori. Sporae incolores oblongae bilocu-
lares haud placodiomorphae, 10-12 x 4-5 u.
On rock, Tutu, St. Thomas, collected by Dr. N. L. Britton, Mrs.
E. G. Britton, and Miss Delia W. Marble, Feb. 8-9, 1913, no. 469
(type!).
Lecania euthallina differs from L. erysibe (Ach.) Th. Fr. in the much
better developed thallus (whence the specific name), it being compact,
BRITTON: FLORA OF THE VIRGIN ISLANDS Hs
thicker, and more continuous. Furthermore, the apothecia are more
concave, with the persistent thalline margin more conspicuous.
41. PARMELIA CETRATA f. SUBISIDIOSA Muell. Arg. On tree-trunk,
Crown, 1441 in part. North Carolina, Florida, Cuba, Jamaica.
42. PARMELIA CONSPERSA (Ehrh.) Ach. On rocks, Crown, 450 m.
altit., 1358.
43. PARMELIA LATISSIMA var. CRISTIFERA (Taylor) Hue. On tree-
trunk, St. Peter, 1249; on twig, Crown, 1441 in part.
44. PARMELIA PERLATA (L.) Ach. On rocks, near Bonne Resolution,
446; on Spondias, Mandal, 1311.
45. RAMALINA COMPLANATA (Sw.) Ach. Without data, 1356a.
46. RAMALINA GRACILIS (Pers.) Nyl. On twigs of Guettarda, Crown,
450 m. altit., 1356.
47. Blastenia nigrocincta Riddle sp. nov.
Thallus crustaceus arcte adnatus sat crassus, ambitu subradiato-laciniatus
effiguratusque, centro 1imoso-areolatus, areolis 0.4-0.8 mm. latis leviter convexa
primum contiguis demum hypothallo nigro dispersis, cinereo-albescens dein fumosus
aut partim luridus. Apothecia 0.3-0.5 mm. lata, superficialia dispersa vel partim
caespitosa nuda, disco plano vel leviter convexa ferrugineo-aurantiaco, margine
proprio sat tenue persistente nigro nitido, margine thallino nullo; excipulo externe
coeruleo-nigro interne incolore; epithecio ferrugineo; hymenio incolore; hypothecio
pallide fuscescente. Asci 8-spori. Sporae incolores ellipsoideae biloculares placodio-
morphae, loculis poro tenue confluentibus, 12-14 x 5-6 yu.
On rock, Tutu, St. Thomas, collected by Mrs. E. G. Britton and
Miss Delia W. Marble, Feb. 8-9, 1913, no. 469a (type!). Also, on
limestone, Montalva, Porto Rico, N. L. Britton, J. F. Cowell, and
Stewardson Brown, March 2-4, 1915, no. 4810.
This species is distinct in the contrasting coloration of the black
margin and the orange disk of the apothecia, a character which
will serve to distinguish it on the one hand from species with similar
thalline characters, such as Bl. Forstroemiana (Fr.) Muell. Arg.;
and on the other hand from BI. ferruginea (Huds.) Koerb., where the
disk and margin are concolorous, and from Blastenia peragrata (Fée)
Muell. Arg., where the margin is black, but the disk is aeruginous-
brown.
48. Caloplaca diplacia (Ach.) Riddle comb. nov.
Lecanora Ach. Synop. Lich. 154. 1814.
On rock, near Charlotte Amalia, 493, 495, 1485 in part. Also
recorded by Nylander in Flora 63: 127. 1880. Apparently confined
to the West Indies.
49. CALOPLACA MURORUM (Hoffm.) Th. Fr. On rock, near Charlotte
Amalia, 1485 in part.
114 BROOKLYN BOTANIC GARDEN MEMOIRS
50. Caloplaca subsequestra (Nyl.) Riddle comb. nov.
Lecanora Nyl. Flora 63: 127. 1880.
On rocks, without definite locality, collected by Dr. Forel. En-
demic.
51. BUELLIA DISCOLOR (Hepp) Koerb. On rock, Tutu, 469b; with-
out definite locality, collected by Dr. Forel, according to
Nylander (1. c.). Europe.
52. BUELLIA PARASEMA var. AERUGINESCENS (Nyl.) Muell. Arg. On
coconut near Charlotte Amalia, 489a.
53. Buellia prospersa (Nyl.) Riddle comb. nov.
Lecidia Nyl. Flora 63: 127. 1880.
On rocks, without definite locality, collected by Dr. Forel. En-
demic.
54. PyXINE cocoEs (Sw.) Nyl. On bark, near Bonne Resolution,
445; on Melicocca, Tutu, 466.
55. PYXINE COCOES var. ENDOXANTHA Muell. Arg. On Guilandina,
Smith’s Bay, 1280; on bark, Mariendahl Road, 1475.
56. PyxINE MEISSNERI Tuck. On coconut palm, without definite
locality, collected by Dr. J. N. Rose, 3198.
57. PHYSCIA ALBA (Fée) Muell. Arg. On Erythrina, St. Peter, 1443.
58. Puyscia CAESIA (Hoffm.) Nyl. On rocks, without definite local-
ity, collected by Dr. Forel. Recorded by Nyl. Flora 63: 127:
1880.
59. PHyscIA cCRISPA (Pers.) Nyl. On Elaphrium, near Charlotte
Amalia, 491; on roots, same locality, 492, 496; on Melicocca,
Tutu, 461.
60. PHYSCIA EROSULA Nyl. Flora 63: 127. 1880. Based on material
growing on rocks, St. Thomas, without definite locality, col-
lected by Dr. Forel. Doubtfully distinct from the widely
distributed Physcia tribacia (Ach.) Tuck.
61. PHyscIA PICTA (Sw.) Nyl. On rock, St. Peter, 1260; on coconut
palm, collected by Dr. J. N. Rose, 3197.
62. PHYSCIA SPECIOSA (Wulf.) Nyl. On rocks, near Charlotte Amalia,
405; on bark, St. Peter, 1248, 1250.
LICHENS OF “ST. JAN
I. PYRENULA MAMILLANA (Ach.) Trev. On bark of Icacorea, Bor-
deaux, 597.
. MELANOTHECA ACHARIANA Fée. On Inga, Bordeaux, 598. Cuba,
Venezuela.
3. MyYCOPORELLUM ELLIPTICUM Muell. Arg. Flora 72: 508. 1889.
On bark, without definite locality, collected by Levier, no. 113.
Endemic.
NO
BRITTON: FLORA OF THE VIRGIN ISLANDS 115
4. ARTHONIA. On bark, Bordeaux, 577.
5. ARTHOTHELIUM MACROTHECUM (Fée) Mass. On Jcacorea, Bor-
deaux, 540.
6. Graphina nitidescens (Nyl.) Riddle comb. nov.
Fissurina Nyl. Lich. Japon. 108. 1890.
On Nectandra, Bordeaux, 581. Florida, Cuba, Porto Rico.
7. OPEGRAPHA VULGATA Ach. On Maytenus, Little St. James Island,
N. L. Britton & J. N. Rose, 1405.
8. Leptogium marginellum var. isidiosellum Riddle var. nov.
Thallus isidiis tenuibus dense tectus; ceterus ut in forma typica apothecia
nulla.
On wet rock, road to Rosenberg, N. L. Britton & J. A. Shafer,
Feb. 5-7, 1913, no. 276 (type!).
The fringed apothecia being such a characteristic feature of
Leptogium marginellum, it is only after some hesitation that this
material has been placed here. The texture and the wrinkling of the
thallus is in exact agreement, however, with typical specimens. And
the relation of this variety to the species is strictly comparable with
the conditions in L. tremelloides, abundantly fruiting in the tropics,
and its variety caesium, with isidia but very rarely fruiting; and with
L. phyllocarpum and its variety isidiosellum.
9g. PARMELIA TINCTORUM Despr. On tree-trunk, Bordeaux, 567.
The following lichens are recorded in ‘Le Végétation des Antilles
Danoises”’ by F. Bérgesen & Ove Paulsen in Revue Générale de
Botanique 12: 507, 508. 1900.
STICTA WEIGELII (Ach.) Wainio. St. Croix; St. Thomas.
GRAPHIS scrIPTA (L.) Ach. St. Croix.
LECIDEA BUELLIANA Muell. Arg. St. Croix.
PERTUSARIA WULFENII (DC.) Fr. St. Croix.
PARMELIA PERLATA (L.) Ach. St. Thomas; St. Jan.
PARMELIA TINCTORUM Despr. [P. coralloides Mey. et Flot.] St. Croix.
PHYSCIA INTEGRATA Nyl. St. Jan.
RINODINA sp. St. Croix.
VERRUCARIA sp. _ St. Croix.
ARTHONIA RADIATA (Pers.) Ach. St. Croix.
SCHIZOXYLON sp. St. Thomas.
FUNGI
During our exploration of St. Thomas and St. Jan in 1913, about
25 species of fungi were obtained, and Dr. Rose collected four others
on St. Croix; manuscript record has been made of these.
Twenty species collected by Mr. Ricksecker on St. Croix are
116 BROOKLYN BOTANIC GARDEN MEMOIRS
listed by Dr. Millspaugh in his “Flora of the Island of St. Croix,” as
determined by J. B. Ellis and F. D. Kelsey.18
Thirty-one species brought by various collectors to Copenhagen,
determined by E. Rostrop, are recorded by Bérgesen and Paulsen in
their ‘‘ Végétation des Antilles Danoises.”’
Seven species, collected on St. Thomas during the voyage of the
“Challenger,”’ are listed by M. J. Berkeley in Journal of the Linnean
Society 14: 352.
These records duplicate each other considerably, indicating a
known fungus flora of somewhat over fifty species only. Inasmuch
as there must be several hundred species on the islands, a list of
fungi is deferred for further mycological field work.
ALGAE!9
“The Marine Algae of the Danish West Indies”’ is the title of a
work, now appearing in parts, in which Dr. F. Bérgesen, of Copen-
hagen, is carefully describing and adequately illustrating the seaweeds
of these islands. His adoption of the English language in this work
makes it immediately serviceable to American students. Volume 1,
including the Chlorophyceae (Green Algae) and Phaeophyceae (Brown
Algae), was published in 1913 and 1914, and, at the date of writing,
the first 240 pages of Volume 2, dealing with the Rhodophyceae (Red
Algae) have appeared. Other papers of importance, dealing with the
algae of the Danish West Indies, are the following:
Borgesen, F. A Contribution to the Knowledge of the Marine Alga Vegetation on
the Coasts of the Danish West Indian Islands. Bot. Tidssk. 23: 49-57.
Figs. I-4. 1900.
—— Et Bidrag til Kundskaben om Algevegetationen ved Kysterne af Dansk
Vestindien. Bot. Tidssk. 23: 58-60. 1go0. [An abstract, in Danish, of
the foregoing article. ]
— Contributions a la connaissance du genre Siphonocladus Schmitz. Overs. K.
Danske Vidensk. Selsk. Forh. 1905: 259-291. Figs. I-13. 1905.
— An Ecological and Systematic Account of the Caulerpas of the Danish West
Indies. K. Danske Vidensk. Selsk. Skr. VII. 4: 337-392. Figs. I-31. 1907.
—— The Dasycladaceae of the Danish West Indies. Bot. Tidsskr. 28: 271-283.
Figs. 1-9. 22 My 1908.
—— The Species of Avrainvillea Hitherto Found on the Shores of the Danish West
Indies. Vidensk. Medd. Naturh. Foren. K¢benhavn 1908: 27-44. pl. 33.
Je 1908.
—— Some New or Little-known West Indian Florideae. Bot. Tidssk. 30: I-19.
pls. 1, 2, Figs. 1-11. 23 O 1909; II. Bot. Tidssk. 30: 177-207. Figs. 1-20.
9 D toto.
— Some Chlorophyceae from the Danish West Indies. Bot. Tidssk. 31: 127-
152, Figs. 1-13. i911; II. Bot. Tidssk. 32: 241-273. Figs. 1-17. one:
'8 The new species were described in Bull. Torr. Club 24: 207-209. 1897.
19 Contributed by Dr. Marshall A. Howe.
BRITTON: FLORA OF THE VIRGIN ISLANDS 17
— The Algal Vegetation of the Lagoons in the Danish West Indies. Biol.
Arbejd. tilegn. Eug. Warming. 41-45. Figs. I-9. IgII.
— Two Crustaceous Brown Algae from the Danish West Indies. Nuova Notar-
isia 23: 123-129. Figs. 1-3. 1912.
— The Species of Sargassum Found along the Coasts of the Danish West Indies,
with Remarks upon the Floating Forms of the Sargasso Sea. I-20. Figs.
1-8. 1914. [No. 32 of a Mindeskrift for Japetus Steenstrup. ]
Cleve, Peter Theodor. Diatoms from the West Indian Archipelago. [Virgin Islands
and St. Bartholomew.] Bih. Svens. Vet. Akad. Handi. 58: 1-22 pl. 1-5.
1878. Annot. list.
Dickie, George. Marine Algae Collected at St. Thomas during the Expedition of
He M. S: “Challenger.” Jour. Linn. Soc, Bot. 14: 312-313. 17 O 1874.
List.
Millspaugh, C. F. Flora of the Island of St. Croix. Field Col. Mus. Bot. 1: 441-
546. 1902. On pp. 467, 468 is a list of 17 species of marine algae, deter-
mined by Professor W. G. Furlow.
Vahl, M. Endeel Kryptogamiske Planter fra St. Croix. Skrivt. Naturh. Selsk.
Reo 7. TOO2:
ENDEMIC SPECIES
The approximate number of species native to the islands as re-
corded, excluding fungi and algae, is 1,052, as follows:
SPCHMAVOOIY LA s.2 arene typ-3 kee ayo 8s 890 4
Pete OPIN. fs Ae oe acd os se ace os AI
| ECS 71 pn 46
IL JLB SEE 2, SONS erat ee cs a ibs
1,052
The numbers of Spermatophyta and Pteridophyta are not likely
to be increased by further exploration, but there are probably some
more Bryophyta and many more lichens to be obtained. As we know
the flora at the present time, the following 27 species are endemic, at
least to the Virgin Island group as a whole.
Valota Eggersit (Hack.) Hitche. & Chase
Agave Eggersiana Trelease
Peperomia myrtifolia (Vahl) A. Dietr.
Pilea Richardi Urban
Coccolobis Klotschiana Meissn.
Zanthoxylum thomasianum Krug & Urban
Galactia Eggersit Urban
Malpighia pallens Small
Malpighia infestissima (Juss.) Rich.
Maytenus cymosa Krug & Urban
Reynosia Guama Urban
Sida Eggers E. G. Baker
Psidium amplexicaule Pers.
Calyptranthes thomasiana Berg.
118 BROOKLYN BOTANIC GARDEN MEMOIRS
Eugenia sessiliflora Vahl
Chrysophyllum Eggersi Pierre
Forestiera Eggersiana Krug & Urban
Salvia thomasiana Urban
Physalis Eggers O. E. Schulz
Solanum conocarpum L. C. Rich.
Wedelia cruciana L. C. Rich.
Phascum sessile E. G. Britton
Anthracothecium Breuteliit Muell. Arg.
Lecania euthallina Riddle
Caloplaca subsequestra (Nyl.) Riddle
Buellia prospersa (Nyl.) Riddle
Mycoporellum ellipticum Muell. Arg.
The endemic elements are, then, only about 2.6 percent. of the
native flora. A few other species are almost endemic, being otherwise
known only on Porto Rico or on some other neighboring island.
There are a few endemic species known on Tortola, and one on Anagada,
but if the native species of these two islands, additional to those of St.
Thomas, St. Jan and St. Croix, were taken into account, the percentage
of endemism would not be increased.
Porto Rico, with a very much greater area and much higher moun-
tains, has about 13 percent of its species of Spermatophyta and Pteri-
dophyta endemic.
WEATHER CONDITIONS AND PLANT DEVELOPMENT
GEORGE P. BURNS
Vermont Agricultural Experiment Station
The effect of weather conditions on plant development has been
one of the chief problems studied during the past few years by the
ecologist, the agriculturalist, the forester and in some cases by the
plant physiologist. The weather, however, is a variable mixture
composed chiefly of different amounts of light—direct, diffuse, white,
yellow, red, etc., or darkness; moisture—precipitation, humidity,
soil-moisture, etc.; heat, temperature of the air and soil; wind, etc.
Each of these component parts varies within short intervals of time
and each has its effect direct or indirect on the living plants. The
problems of the effect of weather conditions, then, is largely a physio-
logical problem and such problems should be attacked only by means
of accurate experiments under controlled conditions.
The ecologists have been attempting to change from the old
descriptive methods in which the results of a more or less accurate
study of the vegetation of a given area were published. Sometimes
this study was accompanied by a few tables of meteorological data
gathered from a nearby U. S. Weather Bureau station. In only a
few cases were attempts made to relate these data to the descriptive
part of the study and one was often at a loss to know why they were
included in the publication. This type of work has served a good
purpose in a preliminary way but is now outgrown. More accurate
methods have been introduced by advanced workers and ecologists
have adopted the plan of gathering their own data with instruments
placed in the field, the attempt being made to place them under the
same weather conditions as those of the plants under consideration.
The largest amount of data has been collected on evaporation rates
by workers with atmometers. This is probably due to the fact that
these instruments are inexpensive as compared with the cost of the
recording instruments necessary for collecting other data. But they
lack standardization, many kinds, shapes and sizes being in use.
Since no atmometer can be made to work exactly as a plant, ecologists
should adopt arbitrarily one type in order that data wherever col-
lected may be compared. Some ecologists have gone deeply into this
phase of the work and are well equipped with field instruments record-
119
120 BROOKLYN BOTANIC GARDEN MEMOIRS
ing soil temperature, air temperature, humidity, number of hours of
sunshine, wind velocity, precipitation, evaporation, etc.
The fundamental problem, however, presents itself after the analy-
sis has been made of the elements which enter into the compound
“weather.”’ It is the experimental determination of the effect of
those elements, singly and collectively, as measured by the data
compiled, upon the physiological activities of the plant under con-
sideration. This effect can only be measured by means of accurately
conducted experiments in which very expensive apparatus is used.
One of our problems is so to outline the work and to set forth its
fundamental importance that those in authority will be moved to
purchase the ecological equipment without which these agricultural
and silvicultural problems cannot be studied.
In the attempt to solve the problem above outlined some workers
have used a “plant instrument.’ A given kind of plant has been
grown by the side of atmometers, etc., at stations established under
different climatic conditions and an attempt has been made to interpret
their effect as registered by the “‘plant instruments.”’ As an illustra-
tion of the attempt to interpret meteorological data in terms of plant
development let us take the work dealing with temperature. One
method contemplates the subtraction of a constant from the tempera-
tures recorded and considers that thermometric degrees in excess of
this constant are available for purposes of plant development. A
second method seeks to express growth-rate in terms of the velocities
of chemical reactions. A third—the physiological method—attempts
to take into account the optimum and maximum temperatures as
related to plant growth, and the attempt has been made to develop
one formula which will express the combined effect of rainfall, evapora-
tion and temperature on plant growth. This represents but little
more than an attempt to show what might be done if we had sufficient
experimental data on the reaction of plants to the complex conditions
known as the weather.
Much of this work has been based on averages—averages for a
month, a year, or a number of years. We read that a large amount
of data assists in “smoothing out the curve’’ or that the “spas-
modically jerky graph may be smoothed.” It is certainly true that
in some cases the curve should not be smoothed out, because it is the
spasmodic graph that shows sudden changes and the extremes. The
burden of this paper is to show that, in some cases at least, averages
for long periods are of little value as compared to the importance of
the data obtained for certain critical periods in the conditions of the
environment as shown by the ‘“‘spasmodic graph.’”’ Data collected
for a short period in the summer may be very important, but are by
BURNS: WEATHER CONDITIONS AND PLANT DEVELOPMENT 121
no means as valuable as those gathered by recording instruments
during long periods of time. These latter data are valuable, however,
primarily because they cover critical periods during which the environ-
mental conditions are most severe for plant development. The limit-
ing factor is not the average for the long period, but the maximum or
minimum for any factor or group of factors during cértain critical
short periods of the longer season under consideration. This may be
made more clear by illustration. The effect of shade on the develop-
ment of white pine seedlings was under study. Lath shades known
as “‘full-shade’”’ and “‘half-shade’’ were used. It was found that
germination took place sooner and that larger numbers of seedlings
were produced in the ‘“‘no-shade’”’ bed than in those partly or fully
shaded. The temperature of the soil was the controlling factor.
The average soil temperatures computed from readings recorded every
two hours with a Friez machine for the 24 days during the germination
period were ‘‘full-shade”’ 47° F., “half-shade”’ 46.8° F., ‘“‘no-shade”’
49° F. These differences are too slight to have been responsible
for the observed differences in germination. If we look at averages
only and shut our eyes to the daily fluctuations we would conclude
that soil temperature was not the controlling factor. When, however,
the records were examined for extremes it was found that on certain
days temperature variations occurred of as much as 20° F. as between
the soils of the different beds, the soil of the ‘‘no-shade”’ bed reaching
73° F. It is easy to believe that such differences may constitute a
controlling factor, in view of the fact that Atterberg has shown that
this temperature is about the optimum for germination of these seeds.
One evening the nursery foreman reported that every seedling in
our nursery was dead. Examination showed that the white pine
leaves which the day before had been a beautiful green were brown
and apparently dead. A closer study showed that the ends of the
leaves including about one third of the leaf were dead. However,
very little of this leaf browning occurred in another nursery located
near a river bank, protected from the prevailing wind and on a richer
soil. In the upper nursery where the browning occurred we had a
number of recording instruments but unfortunately none in the lower
nursery. The records show: that three slight showers and one
heavy rain (1.28 inches) had fallen just previous to the appearance of
the trouble; that for five days the sun had shown from nine to twelve
hours daily; that a very heavy wind blew for three days before,
especially while the sun was shining; that the humidity dropped
daily below 50 percent, one day reaching 35 percent; that the air
temperature was usually below 75° F.; that slightly protected areas
in the upper nursery showed less damage than did the rest of the
nursery.
122 BROOKLYN BOTANIC GARDEN MEMOIRS
It is of course impossible with any degree of certainty to determine
from these data why the trees in one nursery suffered severely, whereas
those in the other nearby location were but slightly affected. How-
ever, if one were to hazard a guess he might say that it was due to
excessive transpiration, the chief immediate factor being differences
in wind velocity. A similar result has been obtained experimentally
when trees which have been shaded were suddenly exposed to sun and
wind. The next day they showed ‘‘tip-burn”’ of the pathologists,
or a “physiological disease’’ whatever that may be. One fact is
clear. The death of the leaves was not due to the average conditions
prevailing during the summer.
During the winter months the average soil temperatures in the
nursery for depths of three, six and twelve inches were 35.8° F., 37.9°
F., and 38.5° F. Each figure is the average of 2,100 readings taken
every two hours from the record made by a Friez machine. They
show little differences in temperature at the various depths given.
When, however, the record is examined for critical periods it is found
that probably the most important season was that from March 28
to April 18. During this period of 21 days the soil three inches deep
froze and thawed sixteen times, at six inches, nine times and at twelve
inches four times. Similar data collected in the adjacent forest
showed that the soil both at six and at twelve inches thawed only
once. We have no experimental data which determine the physio-
logical meaning of these facts, but it is easy to surmise that in studying
the effect of soil temperatures on plants we will not go far afield if
we study carefully conditions obtaining during these critical periods
as well as indeed full more than the general averages for the entire
winter. .
Numerous examples could be given to show that averages extending
over long periods for humidity, sunshine, wind, air temperature, etc.,
not only explain little but, on the other hand, conceal the essential
facts. In all study of the relation of weather conditions to the develop-
ment of plants the importance of critical periods in the environment
must be taken into consideration.
MODERN APPLICATIONS OF BOTANY
MEL ‘T. COOK
New Jersey Agricultural Experiment Station
It is very doubtful if any science is so thoroughly misunderstood
by the public as the science of botany. To the average layman it is
usually a study of flowers which usually involves harmless collections,
classifications and mysterious Latin names; a study for the faddist;
a study without applications of any value whatever. It is strange .
that a subject dealing with organisms upon which we are dependent
for practically all of our food, clothing and fuel, a large part of the
material for building and the manufacture of useful implements of
various kinds, and most of our drug products should be so misunder-
stood. Yet, even the educated layman knows more about the Panama
canal than he does about wheat, more about flying machines than he
does about potatoes, and more about the Woolworth building than
he does about cabbage. The names of great and near great military
leaders, statesmen, ministers, physicians, architects, theatrical stars,
ball players and pugilists are familiar to those millions, while very
few can name a single person who has contributed to the feeding and
clothing of mankind. In fact, few people; even among the educated
classes, realize that agriculture, horticulture and forestry are in reality
specialized branches of botany.
A brief statement of the early history of the subject may offer an ex-
planation of this anomalous position of our science. Botany had its
rise in the development of the medical professions, in the efforts of the
practitioner to determine the uses of plants in the art of healing.
This resulted in the study of local flora of a number of the most
advanced countries and also the search for plants in foreign countries.
Very naturally, the great number of species of plants forced these
early students to formulate some system of classification whereby
their materials might be catalogued. With their increasing knowledge
of these species, it became necessary to devise new systems until finally
this phase of the subject became all important. In the meantime,
the medical profession gradually discontinued the use of the less
important of the medicinal plants for those that were most easily
obtained, most economical in preparation and most efficacious in
use. A little later, we find the physician studying the crude drug and
123
124 BROOKLYN BOTANIC GARDEN MEMOIRS
a little later the prepared drug and paying little or no attention to
its origin. Thus the two professions developed along diverging lines.
In the meantime, the invention and development of the microscope
opened new and interesting fields to the botanists as well as to other
scientists and also resulted in the rise of bacteriology, which has had
such a marked influence on many lines of work, especially medicine.
At the same time the scientific study of agriculture was beginning
to attract attention but, unfortunately, it is not an outgrowth of
botany. Chemistry became the first sponsor for this new field of
research and the first directors of many of our American agricultural
experiment stations were chemists; they studied the soils and de-
veloped formulas for fertilizers—for what? To make plants grow,
to increase plant production, and thus the problem of plant growth
was taken by the chemists instead of the botanists. .
Horticulture was very closely associated with botany and the
developments of horticulture and botany were combined in many of
our agricultural colleges. In many cases these soon came to be
known as departments of horticulture, the botany becoming a vanish-
ing factor; but in later years botany has re-entered these colleges as
an independent, but in many cases a secondary subject. In those
agricultural colleges in which botany has had a continuous existence,
the lines of research were by no means the same. In some cases, they
studied weeds and devised methods for their control; in others, they
co-operated with the horticulturists in the study, introduction and
improvement of valuable food and fiber plants; in others they studied
the causes and methods of controlling plant diseases, but in many
cases the second phase of the subject was quickly taken over by the
now independent departments of horticulture.
It is impossible to tell just what the result would have been if
our botanists of a quarter of a century ago had been as energetic in
the development of the applied side of botany as the chemists were
in the development of the applied side of chemistry. But it is reason-
able to suppose that the results would have been similar, and that we
would have today, not only the applied phases of botany, but we
would also have far more workers on technical problems.
The future of botany in America is brighter that at any time in
its history. It is a recognized subject in our universities, in arts and
in agricultural colleges. It is recognized, both as a cultural subject
of great value and interest and as a science with a direct bearing on
the affairs of mankind. The botany of today means not only tax-
onomy, morphology, cytology and physiology as purely scholastic
subjects but all in their relation to applied plant physiology, plant
breeding and plant pathology with a direct bearing on horticulture,
agronomy and forestry.
COOK: MODERN APPLICATIONS OF BOTANY 125,
Plant growth is no longer a problem for chemists but for the
plant physiologist, who is trained not only in botany, but in chemistry,
physics and geology. Plant physiology has outgrown the expectations
of its most enthusiastic devotees of a decade ago, and no one can
foretell its future. It will doubtless result in important changes in
agricultural methods.
Plant breeding, along the lines of artificial selection, is very old;
in fact, it must have originated with the first steps in civilization.
Many of our valuable economic plants were selected, grown and
used by man before the beginning of written history and many im-
proved varieties have been developed by self-taught, practical workers,
men of great natural endowments and keen powers of observation.
However it is none the less true that they are the products of the
workings of natural laws and that a knowledge of these laws enables
the present generations to work more rapidly than their ancestors.
Many of our modern plant breeders are very properly more interested
in researches leading to a knowledge of these laws than in their appli-
cation. A law fully established and well understood will very soon
be utilized by those interested in increased production. But the
breeder should not loose sight of the very great value of plant breeding
to agriculture. The final and true standard of measure of the value
of any science must be in terms of its contributions to the welfare of
mankind.
Plant pathology is one of the last of these branches of applied
botany to be considered. It had its rise in the taxonomic study of
fungi, many of which were recognized as the causes of plant diseases.
Therefore, this study very naturally led to the study of methods of
control. Indefinite and uncertain methods for the control of plant
diseases have been used from time to time for more than a century.
But a lack of definite knowledge of the causes and the physiology of
these diseases and the actions of the remedies made the results very
uncertain and very soon led to their disuse.
Modern plant pathology had its beginning in the works of de Bary
and Berkeley, but did not make much progress until the latter part of
the last century. The progress during the last decade has been rapid
and has emphasized the necessity of many lines of study, such as a
more thorough knowledge of the life history and taxonomy of the
parasites, a knowledge of the physiological factors influencing both
host and parasite and a knowledge of the physiological effects of the
fungicides. It is also extremely important that we make extensive
investigations on that ever increasing number of diseases which cannot
at this time be attributed to any definite organism.
The prosecution of these lines of investigations means more in-
126 BROOKLYN BOTANIC GARDEN MEMOIRS
tensive researches in the taxonomy, morphology and physiology of
the fungi and other organisms that cause diseases; in the morphology
and physiology of the flowering plants; and plant breeding. The
directing ideal in physiology and plant breeding must be the improve-
ment of the plant for economic purposes, the development of resistance
to disease and the increase in plant production.
Many phases of plant pathology are practically untouched. The
greatest advancement has been made in the study of the diseases of
orchard fruits, much has been done in the study of cereals, shade and
forest trees, and certain truck crops, such as potatoes. While much
work still remains to be done on the diseases of these crops, much
more is necessary on miscellaneous truck crops and on ornamentals.
The fact that truck and ornamental crops are grown under glass
presents new and complicated problems of the greatest economic
im portance.
Many people, even botanists, have the idea that all phases of
applied botany must be restricted to agricultural colleges. This is
an unfortunate error which tends to broaden the gap between the
botany on one side and horticulture, agronomy, forestry, etc., on the
other. Only recently, a well-known government plant pathologist
told the speaker that he had no great difficulty in securing young men
trained in plant pathology but that, unfortunately, many of them
were not trained in botany. Applied botany is in very great need of
workers who have a thorough fundamental training in botany plus a
specialized training in applied botany. Much of this work can be
done to an advantage in our universities provided the proper viewpoint
can be obtained. I use the term ‘‘viewpoint”’ guardedly, for while
it is true that many of our workers in applied botany are poorly
trained in fundamental botany, it is also true that many of our uni-
versity men are about as well fitted for applied botany as the students
of Hebrew. It has been said that no one can apply a science unless
he has learned the science, but it is equally true that some learn a
science that cannot be applied. The suggested applications in some
technical papers compare very favorably with the comic sheet in the
Sunday papers.
But the few lines of work indicated in this paper do not include
all that are open to the botanists. Many of the manufacturing
industries are needing, and will need for years to come, many men
trained in botany and biochemistry. Some time ago the writer was
asked to recommend such a man to make investigations on cellulose.
Failing to find such a man, the company employed a chemist. The
manufacture of rubber is another industry in which the services of a
properly trained botanist can be very useful. And there are many
COOK: MODERN APPLICATIONS OF BOTANY 2,
other lines of work too numerous to mention in the short time avail-
able for this paper. Furthermore, in the very near future, America
may be called upon to furnish botanical workers for the world. A
prominent London journal has already called attention to the necessity
for the development of the agricultural resources of Great Britain’s
colonies, and admitted that the workers must come from America.
The great resources of South America are practically undeveloped.
Thus far, those countries have called on European countries for most
of their workers, but in the near future they will probably turn to this
country. With the close of the great international war, now in pro-
gress, the United States will probably become the great education
center of the world, but we must give educational work largely along
industrial lines. Are the American botanists prepared to meet the
new demands? .
STUDIES IN THE GENUS GYMNOSPORANGIUM—I.
NOTES ON THE DISTRIBUTION OF THE
MYCELIUM, BUFFER CELLS, AND
THE GERMINATION OF THE
AECIDIOSPORE
B. ©. DODGE
Columbia University
‘The sporophytic mycelium of different species of Gymnosporangium
exerts an influence in connection with the growth of the tissues of the
cedar hosts which is manifested in a variety of ways. It is not clear
just why one species will cause the formation of a rather fleshy gall,
while another species will lead to the development of a witch’s-broom
or a hard, woody burl. A study of the interrelationships of host and
parasite, especially the more intimate association of the hyphae and
the host cells may help to solve some of these interesting questions.
I wish to report briefly at this time the results of some studies
made to determine: (1) the distribution of the mycelium in an infected
leaf; (2) the possibility of its spread from leaf to stem; (3) the degree
to which it spreads up and down; (4) its distribution in wood, phloem
and cortex; (5) the distribution of haustoria. Four-nucleated
aecidiospore germ-tubes of Gymnosporangium transformans, and the
formation of buffer cells in the teleutospore sori of G. fraternum and
G. transformans will be noted.
Farlow! determined the general distribution of the mycelium in the
host for a number of American Gymnosporangia and described with
considerable clearness the primary effects of the parasites on the
tissues of the host plants. He found that the burls on Chamaecyparis
infected by G. biseptatum are probably the result of a stimulation of
the cambium by the hyphae mainly distributed in the cambium region.
There appeared to be very little in the nature of a deleterious effect
of the parasite on this host. It was evident to him, however, that
the actual presence of mycelium in a given tissue is not necessary
to account for distortions or abnormalities. In G. Ellisii the fungus
interferes with the normal growth of the host, producing proliferations
and swellings of the stems and branches. This may be due to a dis-
turbance in the nutritive processes, the primary cause of which may
‘Farlow, W. G. The Gymnosporangia or Cedar Apples of the United States.
Am. Mem. Boston Soc. Nat. Hisc. 1-38. pl. 1, 2. 1880.
128
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM§ 129
lie at some distance. Farlow noted that the hyphae of this species are
exceptionally large and that the brown mycelium runs down into the
wood and along the medullary rays and also makes other brown
patches extending some distance in circular areas between the annual
rings. The greater part of the mycelium is found near the cambium
and large masses of it are collected at points in the bark in prepara-
tion for the formation of sori.
Wornle? made an extensive study of the relationships of host and
parasite in nine species, and his report published in a forestry
journal furnishes a valuable contribution on the subject. He en-
deavored to determine the particular tissues with which the mycelia
are associated and stated his conclusions with considerable positive-
ness. G. Juniperinum was of special interest to him inasmuch as he
supposed that the sori found on leaves as well as those on small twigs
belong to the species that produces larger sori on the main stems. He
could see that the mycelium in an infected leaf was connected with
that from a small twig. He also learned that the leaf form is per-
ennial. Four successive cork callus formations were found in one
case, showing that for four years a sorus had been developed at the
same point on the leaf. In the stem-inhabiting type he found that
the mycelium is present in the wood as well as in the bast and cortex.
Radially placed strands of parenchyma accompanied by mycelium
are common in the wood; ‘‘Schlafende Augen”’ he calls them. Hyphae
are intercellular, and he noted in some cases the presence of haustoria.
Although Wornle was not himself clear regarding the relationship of
the three forms of the rust which he called G. Juniperinum, he was
inclined to believe that the fungus gains entrance through the leaves,
the mycelium later running down the twigs and into: the main
stem, where it becomes firmly established. Fischer* has: shown that
this was a false assumption since W6rnle was dealing with at least
two species, but the accuracy of Wornle’s observations is not ques-
tioned.
The mycelium of G. clavariaeforme, according to Wornle, is not
present in the wood, although considerable transformation of tracheid
tissue is to be seen in infected stems; arcs and sectors of this
tissue are replaced by parenchymatous cells. He found, however,
no mycelium in such areas. As the mycelium is generally distributed
in the cortex and bast, he assumes that the cambium is in some way
stimulated to develop more than a normal amount of wood,cells, some
* Wornle, P. Anatomische Untersuchung der durch Gymnosporangium-Arten
hervorgerufenen Missbildungen. Forst. Nat. Zeits. 3: 68-84, 129-172. 1894.
’ Fischer, E. Studien zur Biologie von Gymnosporangium juniperinum.
Zeits. Bot. 1: 683-714. f. I-8. 1909; 2: 753-764. I910.
10
130 BROOKLYN BOTANIC GARDEN MEMOIRS
of which are inhibited in their growth, lacking bordered pits and
having thin walls, that is, are more in the nature of parenchyma.
Wornle was especially fortunate in having the opportunity for con-
sultation with Hartig and Tubeuf in his work, but he was handicapped
in studying American species by being restricted to a limited number
of dried specimens. He agrees in the main with Farlow’s account of
the location of the mycelium in G. biseptatum and G. Ellisti, although
he makes no mention of Farlow’s work. He concludes further that
the mycelium of G. biseptatum is intercellular and is entirely absent in
the wood. The tracheids are somewhat irregular and have thinner
walls than ordinarily. He found that the hyphae of G. Ellisw are
about 8 w in diameter and are present in the wood, bast and cortex.
The brown hyphae are associated with brownish cells which together
make easily recognizable patches. The mycelium here also is strictly
intercellular. His study of a three year old stem of red cedar infected
with G. clavipes disclosed the fact that the tissues of the host are only
slightly affected. The mycelium is distributed not only beneath the
sorus but in the whole periphery of the twig, especially in the bast
region. The wood is entirely free from the fungus. W6rnle pre-
dicted that this rust must develop sori one year after inoculation
because in this three-year-old stem he found traces of two former
sori, one above the other.
Harshberger’s account! of the relationships of hyphae and
host cells deserves special consideration, inasmuch as it does not
agree in certain important particulars with the statements made by
Farlow and Wornle. He finds that in G. biseptatum the mycelium
is quite generally present in the wood region where he states the
hyphae are for the most part strictly intracellular. They run down
through the lumen of a tracheid, pass out through bordered pits,
enter an adjacent tracheid, or move over to medullary ray cells which
they penetrate and thus become established where they receive nour-
ishment sufficient to maintain their perennial growth as the wood of
the burl increases in diameter. He believes that the hypha actually
in the lumen of the cambium cell is responsible for the stimulation of
this cell to produce abnormal amounts of wood! He describes and
figures these intracellular hyphae in much detail, especially the
hyphae in longitudinal sections of wood. The explanations accom-
panying his figures leave no doubt of Harshberger’s opinion regarding
the identification and location of intracellular hyphae. He lays much
stress on the presence of “‘plugged”’ tracheids. He believes that
they are caused by the mycelium with which they are generally asso-
4 Harshberger, J. W. Two Fungous Diseases of the White Cedar. Proc. Acad.
Nat. Sci. Philadelphia 1902: 461-504. pl. 22, 23.
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 131
ciated. Hyphae may sometimes be intercellular. Swollen and
nodular hyphae are not infrequent. Harshberger questions whether
haustoria are ever present. It is well known from the work of Hartig
and others that hyphae of wood-destroying fungi are capable of boring
through lignified cell walls. Such fungi obtain their nourishment by
activities leading to the disorganization of wood cells. The rusts are
highly parasitic and haustoria play an important part in their nutrition.
It would be interesting to find that such trunk parasites as G. biseptatum
and G. Ellisii are more like the common heart rot fungi than they are
like other rusts where the hyphae crowd in between the cells or mass
in the intercellular spaces. My own observations do not support
several statements made by Harshberger.
GYMNOSPORANGIUM ELLISII
I have succeeded in infecting Chamaecyparis by spraying potted
cedars with aecidiospores of G. Ellisii (G. myricatum). Several
cedars naturally infected and bearing brooms of different ages have
also been grown in pots, so that I have had an abundance of material
in all stages of growth for study.
The sorus usually matures about twenty-one months after inocu-
lation. Where young leafy branches have been infected we find
that the sorus may break out either in the leaf axil or through the
leaf itself. At this time there is very little distortion of the twig.
The primordium of the axial sorus is partly in the tissue at the base
of the leaf and partly in the stem cortex beneath. Where the sorus
emerges through the leaf we find that there is an increase in the
number of mesophyll cells and the sorus primordium is not far below
the epidermis. Strands of hyphae’can be traced down to the short
vein and into the woody portion of the stem. Serial sections show
that the mycelium does not travel up and down the stem very rapidly;
in some cases only one or two cm. in the first two years. Where a
rapidly growing main stem is infected the hyphae run as much as
five cm. in the same time. Trunks thirty years old have been cut
and one such shows traces of mycelium for a vertical distance of only
about ten cm., although the fungus had been active during the life
of the tree, thirty years. Sections taken from various parts of a small
artificially infected plant bearing a dozen potential witches’ brooms
skow that each broom will be the result of a separate infection.
The mycelium does not enter at one point and spread through the
entire plant. However, if the original infection should be at the
growing point of the main stem a broom is formed that permanently
dwarfs the plant. The mycelium invades every tissue except the
cork. It is found in patches in all of the annual rings, and is espe-
132 BROOKLYN BOTANIC GARDEN MEMOIRS
cially abundant along some of the medullary rays. It is not evenly
distributed. The hyphae seem to travel in fascicles and they are
everywhere intercellular. Sections of the wood show that there are
strands of parenchyma that, from appearances, would seem to be
burrowing through the wood, thrusting the tracheids aside as though
endowed with great power. These same parenchyma strands are
also found in the cortex. They run in almost every direction. Hyphae
are always associated with them. ‘Tracheids in infected areas of the
wood are considerably modified. The walls are thinner, the cells are
prismatic and in many cases have failed to develop bordered pits.
The walls of such cells frequently appear to be broken down or
crushed in and partially disorganized. It may very well be that the
fungus has some power to disorganize lignified cell walls. Wherever
hyphae occupy the lumen of a cell it is likely to have been the result
of such mass action. ‘There is no boring through the walls nor entering
tracheids through bordered pits. The ‘‘Schlafende Augen,” or
parenchyma strands, in the cortex and along the line of medullary
rays in the wood as well as the patches of abnormal or partially
developed tracheid tissue are the result of the stimuli proceeding
from hyphae that were nearby at the time this tissue was being de-
veloped. It is difficult to understand how a cambium cell harboring
a hypha could divide at all, or how a tracheid could change its form
once it has become lignified.
The cambium reacts in such a way as to cut off by the excessive
development of tracheids certain fascicles of hyphae and thus
check the radial and longitudinal advances of the fungus. The
apparently isolated patches of mycelium found in the heart wood are
nevertheless quite generally connected above or below with some
radially placed strand that ultimately reaches the cortex. This may
be the main reason why one finds living hyphae deeply imbedded
beneath several rings of wood.
Haustoria may occasionally be found in cells of the cortex medullary
rays, but they are not abundant. Some of these haustoria are bi-
nucleated.
There seems to be no question that Wornle was right in stating
that the hyphae of G. Ellisii are intercellular.
GYMNOSPORANGIUM BISEPTATUM
Harshberger and Wornle disagree on a second important point in
their studies of G. biseptatum. This relates to the presence or absence
of mycelium in the wood cylinder of the cedar.
I have as yet been unable to infect the cedar with this species.
I have studied specimens naturally infected and especially one from a
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM § 133
small plant which I was able to transplant and grow in the greenhouse.
In 1915 this small burl bore two sori. The same burl bore six sori in
1916. The branch was six years old when cut. The mycelium spreads
quite evenly through the cortex and is especially abundant beneath a
sorus where we find one or two large haustoria in nearly every cortex
cell. The medullary ray cells of the cortex are likewise attacked and
the mycelium penetrates down to the cambium. The walls of the
tracheids are somewhat thicker than usual, in this respect differing
from the specimens examined by Wornle.
I have been unable to find any intracellular hyphae, and in this
six-year-old branch there are certainly no hyphae inside of the cambium
ring, that is, in the wood cylinder, such as Harshberger describes.
The most striking feature about this fungus is the great abundance
of large haustoria found in nearly every cell of the cortex in the
vicinity of a sorus.
GYMNOSPORANGIUM CLAVIPES
The red cedar may be infected with G. clavipes without difficulty
by spraying with aecidiospores. Plowright® states that it takes two
years for G. clavariaeforme to mature sori, but Tubeuf® found that
sori developed one year after inoculation of the juniper. My experi-
ence with G. clavipes may serve to explain this discrepancy.
On August I, 1915, a small cedar was inoculated with Gymno-
sporangium clavipes. A few sori appeared in 1916 on what was, in
1915, the growing region of the main stem. In 1917 sori burst out
quite generally over the plant. The question has arisen: Is it possible
that from the original point of infection of I915 the mycelium ran
down the main stem out into the branches where further sori formed
in 1917? Inspection showed that the sori were not evenly scattered
along the branches, but appeared in groups with intervening spaces
of some length between, varying from one to several cm. Serial
sections of some of the smaller branches made at points between
groups of sori do not show the presence of mycelium. For several
inches near the top of the main stem the sori are so close together
that mycelium appears to be continuous. It is noteworthy, however,
that there are no sori on those parts of the plant that have grown
since the plant was inoculated in August, 1915. The mycelium is
intercellular and lies for the most part well out in the cortex just
beneath the cork, some hyphal ends even pushing in between the inner
cork cells. It may require only one year for full development at the
5 Plowright, C. B. British Uredineae and Ustilagineae. 1893.
®Tubeuf, C. Mitteilungen iiber einige Pflanzenkrankheiten. Zeitschr.
Pflanzenkr. 3: 201-205. 1893.
134 BROOKLYN BOTANIC GARDEN MEMOIRS
growing point where abundant food is available, or even take two years
in regions less favorably located. The characteristically binucleated
haustoria are of large size and are easily demonstrated.
I have examined several stems three years old but do not find
that the mycelium spreads out through the entire cortex and into the
bast as described by Wo6rnle. G. clavipes brings about less increase
in development of wood tissue than G. biseptatum, but this may be
due to the fact that the mycelium does not approach the cambium as
closely. This species may develop strictly foliicolous sori in which
case the mycelium is very limited in extent. The sorus is then not
deep seated. Haustoria can be found in epidermal cells.
GYMNOSPORANGIUM TRANSFORMANS
I have previously reported’ that two leaf-inhabiting species of Gym-
nosporangium can be distinguished on Chamaecyparis. The account of
the cultures in support of this statement is being published in another
paper. For convenience I shall call one form G. transformans. Its
aecidial form is Roestelia transformans on Aronia. Gymnosporangium
fraternum is an appropriate name for the second leaf form which
infects Amelanchier. The aecidium is very similar to that of Roestelia
Botryapites; I am not prepared to prove that it is R. Botryapites. A
cytological examination of cedar leaves infected with G. transformans
and G. fraternum reveals further characteristics by which they may be
distinguished.
If we section a leaf of Chamaecyparis infected with G. transformans,
we find that the mycelium is especially abundant in the large inter-
cellular spaces of the spongy mesophyll and the hyphae push in be-
tween the cells of the palisade on all sides. No hyphae are to be found
in the epidermis. If the section includes the short vein of the leaf we
see that hyphae are prevented in some way from entering the vascular
tissue. There is an irregular row of large cells surrounding the vein
of the leaf. These pericycle(?) cells do not normally form a per-
fectly closed ring; it is occasionally broken by smaller supporting
cells. In regions where the hyphae reach the large cells one can
find, here and there, that they have been penetrated by one or two
haustoria. Such infected cells are about one third larger than usual.
The cytoplasm is rather dense, including considerable stored food
and takes the gentian violet stain somewhat deeply. The nuclei
appear to be quite normal. Haustoria are also occasionally found in
mesophyll and palisade cells. The cells of the mycelium are bi-
nucleated and the nuclei stand out very clearly, especially where
7 Dodge, B. O. Report on further cultures of Gymnosporangia. Paper read
at the December meeting of the Botanical Society of America, New York, 1916.
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 135
leaves have been fixed at a time when the sorus is fully matured. The
mycelium is confined to the leaf bearing the sorus and does not ordi-
narily invade the stem at any point along the line of attachment.
The large cells surrounding the leaf vein appear to prevent the my-
celium from entering the phloem of the stem.
At the point where the sorus is to be developed, we find a well-
defined pseudo-parenchyma, the cell walls taking the orange stain.
The upper cells of this pseudo-parenchyma are somewhat enlarged
and elongated. These are likewise binucleated. They soon begin
to swell, lose their cytoplasm, and the nuclei degenerate. In order to
show these upper cells in this condition the material must be fixed at
the earliest possible time that an infected leaf can be distinguished,
Fic. 1. Section of a sorus of G. transformans on a leaf of the southern
white cedar at the narrowest portion of the young sorus. The epidermis is broken
up On either side, only traces of the cuticle and fragments of the cell walls being
visible. At the center epidermal cells are still visible. A number of buffer cells
in various stages of degeneration can be seen, and binucleated teleutospore buds
growing through the buffer cells are common.
that is, when a spot appears as a slight, waxy, translucent, light orange
blister. In such cases the epidermis may not have been ruptured
and fixation of the mycelium is not apt to be of the best, unless the
leaf is cut through. The upper cells mentioned become mere bladdery
sacs and during this process of swelling the inner walls of the epidermal
cells, and hypodermal cells when present, are broken down either by
enzyme action or by actual pressure, and the heavily cutinized epi-
dermis is lifted up and split open (Text-fig. 1). The splitting usually
occurs in a line along one side of the leaf, but very often the split
runs longitudinally down through the middle. Sometimes two sori
develop side by side on the same leaf. The bladdery cells evidently
function as buffer cells to disrupt the epidermis. These buffer cells
perhaps represent simply the first series of teleutospore mother cells
136 BROOKLYN BOTANIC GARDEN MEMOIRS
that are sacrificed in order that the epidermis may be broken open
(Text-fig. 2). The true basal cells now grow out through the buffer
cells, forming a club-shaped bud which soon becomes binucleated.
These nuclei divide and the stalk is cut off; later three pairs of nuclei
can be seen and the wall is formed between the two cells of the teleuto-
spore (Text-fig. 3). Quite mature spores can be found along the line
where the epidermis first ruptures and all stages in their development
can be seen further back.
Fic. 2. A portion of a sorus of G. transformans showing two rather pointed
buffer cells forcing epidermal cells aside, fragments of the walls of the epidermal
cells lie just above the buffer cells at the right. Four young teleutospore buds
are visible.
Fic. 3. Teleutospore of G. transformans.
The gametophytic stage on Aronia may sometimes attack the
young stem, giving rise to an irregular herbaceous gall, which becomes
covered with horn-like projections from which the aecidia arise.
If such a plant is kept in the greenhouse all winter, aecidiospores will
continue to be formed in some cases for several months after the
leaves have fallen from the plant. These spores are regularly bi-
nucleated and possess 7 or 8 germ pores irregularly distributed (PI. I,
Fig. 1). The spores germinated on agar or water frequently form a
swollen pouch near the tip of the germ tube. The nuclei of the spore
push through the germ pore, apparently one closely following the other
(Pl. I, Fig. 3). At the next stage we find two nuclei lying in the
germ tube just outside of the pore (Pl. I, Fig. 4). These nuclei then
migrate further out into the tube and come to lie in the swollen pouch
(Pl. I, Fig. 5) where they presumably divide conjugately, since many
cases have been observed where there were four nuclei, in pairs, lodged
in this portion of the tube (Fig. 6). The germ tube now elongates
rapidly and branches freely at the tip (Fig. 8). The four nuclei move
forward and may lie along the tube in a row in the wider portion at
the end or they may be distributed, one nucleus in each branch, or
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 137
two lying further back, the other two occupying separate branches.
A cell wall is finally laid down, cutting off the outer portion of the
germ tube containing the nuclei and most of the cytoplasm (Figs.
8-10). This curious method of germination is in no sense similar
to the development of a promycelium, though four nuclei are pro-
duced in each case. It may be that it is fairly common among the
rusts as Sappin-Trouffy has pointed out. Whether or not it is pos-
sible to find an appropriate artificial medium for the development of
the mycelium of a rust in artificial cultures, it would seem that such
cases as these afford at least a starting point. Four-nucleated germ
tubes are the rule in these cultures, but fully developed tubes with
only two nuclei are not difficult to find (Fig. 11).
In some of my cultures in which the petiole of a leaf had been
infected at the junction with the blade, it was found that the winter
bud was larger than usual. The mycelium must have run down the
petiole and become established in the bud. When such plants were
put in the cold frame over winter and taken out in the spring, these
buds developed small leaves which at once became evenly covered.
with spermogonia and later were transformed into large galls from
which aecidia developed quite normally.
In some cases the mycelium seems to penetrate into the tissues of
the stem where a spindle-shaped swelling or burl is formed. In the
following spring a green gall bursts out through the cork, forming a
nodular swelling outside and from this spermogonia and aecidia are
produced. I have had several cases in which Roestelia transformans
has survived the winter and developed aecidia the following spring
The same is true in my cultures of R. Botryapites. In October, 1915,
winter buds of six Amelanchiers showed signs of being infected. All
of these survived the winter and developed spermogonia and ripened
aecidiospores in the month of June, which is several months earlier
than they can be found in nature. These are not cases where the
formation of an aecidium has simply been delayed. On the con-
trary, an entirely new crop of spermogonia arises from newly formed
tissue, new gall growth, and we find the aecidia developing as in
normal cases of infection with sporidia.
GYMNOSPORANGIUM FRATERNUM
The buffer cells in the teleutospore sori of G. fraternum are much
more striking in appearance, forming as they do a perfectly even
palisade layer that frequently extends entirely across the sorus without
interruption (Text-fig. 4). This is a very characteristic feature of
8 Sappin-Trouffy, P. Recherches histologiques sur la famille les Urédinées.
Le Botaniste 5: 59-244. f. 1-69. 1 D 1896.
138 BROOKLYN BOTANIC GARDEN MEMOIRS
the rust. The difference in the shape of the buffer cells of G. trans-
formans and G. fraternum corresponds roughly with that between the
teleutospores; they are comparatively long in the latter species.
On January 30, 1917 a potted plant naturally infected with G.
fraternum was taken from the cold frame and examined. Several
leaves showed by the presence of slight yellowish spots that they
would develop sori. Sections of one leaf cut on this date showed
Fic. 4. Section of a leaf of the white cedar infected with G. fraternum show-
‘ing the layer of buffer cells at the time when the epidermis has been quite completely
disorganized. At the right a large palisade cell in the process of disorganization,
but the nucleus is still visible and a haustorium is present.
that the pseudo-parenchyma or teleutospore primordium was well
marked. The buffer cells were mostly without granular contents and
nuclei (Text-fig. 5, 4). Ina few cases fragments of the degenerated
nuclei could be seen. By February 2, a sorus taken from the leaf of
the same plant showed great numbers of teleutospore buds in the
2- and 4-nucleated stages (Text-fig. 5, C). A few buds had 6 nuclei,
and the stalk cells of these had been cut off. Buffer-cell walls
were just visible as narrow irregular lines showing most distinctly
at the base of the young teleutospore. Two days later, February 4,
cross-walls had been formed in many spores but nuclear fusion had
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 139
not occurred (Text-fig. 5, D). Remnants of buffer-cell walls were
now difficult to find. A mature spore is shown in Text-figure 5, E.
The mycelium of G. fraternum penetrates through the leaf in every
direction. The mesophyll cells are usually somewhat enlarged and
are packed rather closely together with small intercellular spaces.
Haustoria are quite abundant in such cells. The cells surrounding
the vein are especially affected. They appear to be filled with minute
granules and as many as eight or ten haustoria can be found ina
single cell, the more common number being two to four. A complete
ring of these large cells is formed. This is due to an increase in
number as well as their larger size. This, taken in connection with
the increase in the mesophyll tissue, gives the leaf a slightly thicker
and more compact appearance. The mycelium does not invade the
Fic. 5. a, pseudoparenchyma with buffer cells; 6, 2-nucleated stage of the
young teleutospore; c, 4-nucleated stage; d, 6-nucleated stage with cross walls; e,
small spore after nuclear fusion.
vascular tissue even to the extent of penetrating the phloem of the
stem. Both G. transformans and G. fraternum are capable of producing
sori two or three years in succession, the latter may produce a.sorus
even after the leaf has apparently died. While haustoria are more
numerous and attack the individual cells more vigorously in the case
of the latter species, G. transformans seems in some way to be more
destructive, as infected leaves more frequently die after maturity of
the first sorus. Of the two species, G. fraternum is clearly the more
nearly related to G. biseptatum, both from the nature of their teleuto-
spores and the similarity of the aecidia in the two species. If the
mycelium of G. fraternum is ever able to push in beyond the large cells
surrounding a vein and get into the central cylinder of the stem,
therefore nearer the cambium, we should look for a stimulus such as
140 BROOKLYN BOTANIC GARDEN MEMOIRS
might lead to the formation of a greater amount of wood tissue such
as we find in the burl of G. biseptatum. G. fraternum has almost con-
stantly 2-celled teleutospores, 3-celled spores are exceedingly rare.
In G. biseptatum 3- and 4-celled spores predominate. I should be
highly gratified to learn that the change in environment from the leaf
to the stem, or more exactly from cortex to phloem, by the fungus
could bring about such a decided change in the structure of its spores.
EXPLANATION OF PLATE I
Gymnosporangium transformans
Stages in the Germination of the Aecidiospores. X 750
Fic. 1. A binucleated spore.
Fic. 2. The germ tube has pushed out, the nuclei are still within the spore
and do not show appreciable change in form. :
Fic. 3. One nucleus is crowding through the germ pore, the other lies beneath.
The “‘pouch”’ is formed just back of the tip of the germ tube.
Fic. 4. Both nuclei have escaped from the spore and lie just outside of the
germ pore.
Fic. 5. Two nuclei lie in the expanded portion of the germ tube.
Fic. 6. Four nuclei are plainly visible in the “‘pouch.’”’ The germ tube has
not begun the second stage of its growth.
Fic. 7. Tip end of fully developed germ tube showing four nuclei in a row.
Fics. 8 and 9. Other fully developed germ tubes showing a more pronounced
type of branching.
Fic. 10. The germ tube has made about the maximum growth of which it is
capable under artificial conditions, a cross wall cuts off the main portion of the
granular cytoplasm at the forward end of the germ tube.
BROOKLYN BOTANIC GARDEN MEMOIRS. VoLuMmE |, PLATE I.
DopGE: GYMNOSPORANGIUM TRANSFORMANS
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INTERCROSSES BETWEEN SELF-STERILE PLANTS
‘E. M. EAST
Bussey Institution, Harvard University
The fact that self-fertilization is practically impossible in certain
hermaphroditic plants, although both the pollen and the ovules are
functional in crosses, has been known since the time of K6lreuter
(1760-1765). The oddity of the phenomenon has been a lure for al-
most every hybridist from that time forward. As in the case of most
other genetical problems, however, our knowledge of its cause and
meaning remained in status quo from the time of Darwin until Men-
delian days. Indeed when the writer began his investigations on the
subject in 1910, the only considerable post-Darwinian work had been
done by a zoologist (Morgan, 1904) on the self-sterile ascidian, Ciona
intestinalis. Since 1910 botanical papers have appeared by Correns
(1912), Compton (1913) and Stout (1916), but these investigations
will not be discussed here, as it is proposed to treat in this paper only
certain phases of the work carried on by the author and his associates!
during the past seven years, leaving critical review for another place.
For our purpose it seems essential only to present a hasty sketch of
the subject as left by Darwin.
In addition to the utilization of most of the previous and the con-
temporaneous work, Darwin (1876) carried out several investigations
of his own on the five self-sterile species, Eschscholtzia californica,
Abutilon darwin, Senecio cruentus, Reseda odorata and Reseda lutea.
Darwin’s first important result was that the expression of self-
sterility in Eschscholtzia californica and Abutilon darwintt was influ-
enced by changes in external conditions. Six generations of Esch-
scholtzia californica had been found to be completely sterile in southern
Brazil by Fritz Miiller (1868, 1873). As English plants were self-
fertile, Darwin obtained from Miiller seed of Brazilian plants of known
self-sterility. The plants which they produced in England, while
not wholly self-fertile, tended toward self-fertility, which fact Darwin
attributed to the lower English temperature. A second generation of
seedlings proved to be still more self-fertile. Conversely, seed of
English stock was somewhat self-sterile the first season and one plant
' The author desires to make grateful acknowledgment to Dr. O. E. White and
Dr. J. B. Park for their painstaking aid in this work. Without it, the numerous
experiments undertaken could not have been completed.
141
142 BROOKLYN BOTANIC GARDEN MEMOIRS
wholly self-sterile the second season, when grown in Brazil. One
may assume, [| think, arguing from data of similar character, that this
progressive result was not due to actual inheritance of an acquired
character but rather to the fact that the first generation in each case
passed a portion of its life cycle in the original environment.
Similar results were obtained in the case of Abutilon darwinii,
which though self-sterile in its native Brazil, became moderately self-
fertile late in the first flowering season in Darwin’s greenhouse.
Darwin made more detailed experiments on Senecio cruentus,
Reseda odorata and Reseda lutea and found, as he believed, that each
plant though self-sterile was cross-fertile with every other plant.
His pollination experiments with Senecio cruentus and Reseda lutea
were so inadequate that they may be omitted from consideration; it
was really his experiments on Reseda odorata that were thought to
establish the fact of complete cross-fertility.
DARWIN’S EXPERIMENTS ON Reseda odorata IN 1868
Male Parents
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Only sixteen cross matings were made, however, and this is not
sufficient to prove the point, as is shown by one of our own experi-
ments, where 131 cross-matings were made with only 4 cases of cross-
sterility. From the fertile cross-pollinations Darwin raised four
plants in 1869. Three of these proved to be self-fertile and one self-
sterile. Six more plants were grown in 1870. Of these, two were
almost self-sterile and four were almost completely self-fertile. The
former produced altogether five seeds from self-pollinations, and the
resulting plants proved to be self-sterile like their parents. These
varied results Darwin attributed to a difference in inherited sexual
constitution, but it seems to me that this conclusion should be ques-
tioned. Our own results have proved conclusively that toward the
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 143
very last of the flowering season? self-sterile plants may sometimes
become somewhat self-fertile.
Darwin’s (1876, p. 346) general conclusions are as follows:
“Finally, the most interesting point in regard to self-sterile plants
is the evidence which they afford of the advantage, or rather the
necessity, of some degree or kind of differentiation in the sexual
elements, in order that they should unite and give birth to a new being.
It was ascertained that the five plants of Reseda odorata which were
selected by chance could be perfectly fertilised by pollen taken from
any one of them, but not by their own pollen; and a few additional
trials were made with some other individuals, which I have not
thought worth recording. So again, Hildebrand and Fritz Miiller
frequently speak of self-sterile plants being fertile with the pollen of
any other individual; and if there had been any exception to the
rule, these could hardly have escaped their observation and my own.
We may therefore confidently assert that a self-sterile plant can be
fertilised by the pollen of any one out of a thousand or ten thousand
individuals of the same species, but not by its own. Now it is obvi-
ously impossible that the sexual organs and elements of every indi-
vidual can have been specialised with respect to every other indi-
vidual. But there is no difficulty in believing that the sexual elements
of each differ slightly in the same diversified manner as do their
external characters; and it has often been remarked that no two
individuals are absolutely alike. Therefore we can hardly avoid the
conclusion that differences of an analogous and indefinite nature in
the reproductive system are sufficient to excite the mutual action of
the sexual elements, and that unless there be such differentiation
fertility fails.”’
One cannot but admire these inductions Darwin has so cleverly
drawn from such meager data, nevertheless one cannot aecept them
today just as they stand. The reasons for this statement will be
seen more clearly when our own data have been presented, but a brief
can be submitted with only the support of the work known to Darwin.
In the first place, the seemingly contradictory results that were
obtained in the experiments on Reseda odorata are not necessarily con-
fusing. As reported, self-sterile plants produced varying ratios of
self-sterile and self-fertile plants. Unfortunately, the progeny of the
self-fertile plants was not followed. If it has been, the problem might
have been more easily solved, for, in all probability, the daughter
plants would have been self-sterile. It is my own belief, however,
that the answer can be read in the casual remarks dropped by Darwin
in the midst of his careful descriptions, remarks to which he paid little
attention. Darwin found that both Eschscholtzia california and
Abutilon darwinii, though self-sterile in Brazil tended to become self-
fertile in England,—especially late in the flowering season. Now
2 Cf. Darwin’s observation on Abutilon darwini.
11
|
144 BROOKLYN BOTANIC GARDEN MEMOIRS
these facts together with that mentioned above regarding the in-
constancy of the results obtained from planting the seed of self-sterile
plants, may be interpreted by the assumption that he was dealing
entirely with fluctuations in all of the five species investigated. These
species genetically were wholly self-sterile. The tendency toward
self-fertility was due to conditions. In other words, these plants
genetically self-sterile needed conditions conducive to a fine healthy
growth to bring out their self-sterility. In the lower temperature of
’ England, at a time of decline (the last of the flowering season), they
became phenotypically somewhat self-fertile. In the light of my own
experiences, I believe we can reconstruct a picture of Darwin’s experi-
ments on Reseda odorata with considerable confidence. He isolated
the plants that he desired to test under nets; then came pressure of
other work, and the data were not collected until the plants had ceased
flowering. At that time capsules were found beneath the nets, and
this seemed to prove at least a partial self-fertility. But instead of
this. procedure, suppose that successive self-pollinations had been
made throughout the season. The presumption is that the plants
would have been declared to be self-sterile with the same remark
added which he jotted down in the case of Abutilon darwinii, — .
they “became moderately self-fertile late in their flowering season.”’
Again, Darwin found no cross- sterility i in the plants tested, ‘and
concluded that a self-sterile plant can be fertilized with the pollen
of any one of a thousand or ten thousand individuals of the same spe-
cies. Such a conclusion was less cautious than was Darwin’s wont for
it was made from a total personal experience of some twenty-odd cross-
matings only, unless his records are extremely incomplete. Indeed
this conclusion must have been somewhat of a surprise to himself
since he states that “it is obvious impossible that the sexual organs
and elements of every individual can have been specialized with respect
to every other individual.’’ He surmounted this difficulty by assum-
ing that the sexual elements of each plant differ slightly in the same
manner as their external characteristics, and that this slight difference
is sufficient to excite the mutual action of the sex elements necessary
in order to have fertilization ensue. The kernel in this conclusion,
that differences in the reproductive systems of two self-sterile plants
are necessary in order to promote cross-fertilization, is so similar to
that to which the writer has been forced after seven years of rather
intensive work as to be uncanny, for it seems to have been reached
in spite of rather than because of.the data at hand. This feeling of
surprise at Darwin’s clairvoyancy may seem affected, since he was
usually in advance of. his time, but it is a. fact perhaps worth men-
tioning as a confession of omission that the writer reached his con-
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 145
clusions as the outgrowth of work on heterozygosis and did not refer
to Darwin’s view until recently. Be this as it may, a short com-
parison of Darwin’s main induction with the facts from which it came
will, I think, show a real reason for wonderment. He believed in
universal cross-fertility of self-sterile plants, his basis being the small
number of cross-fertilizations made by Hildebrand, Miiller and him-
self; although Robertson Munro (1868), with whose work he was
familiar, had found cross-sterility in Passiflora alata, and even the
works of Hildebrand and Miiller as published leave the matter in
doubt. Now how much more reasonable the general induction
mentioned above seems if one assumes (1) that self-sterile plants breed
true for self-sterility but may show a slight degree of self-fertility as a_
fluctuation under certain conditions, (2) that a “variable but limited
number of gérminal | “factors”’ influence the success of matings, cross-
fertilization being possible only when two plants differ in these effective
factors, and (3) that when two plants h have the same effective factorial
composition, cross-sterility of the same type as self-sterility exists.
This is what we believe our own work has shown, as we shall try to
demonstrate.
Emphasis must first be laid upon the fact that the behavior of
self-sterile plants among themselves and the relation between self-
fertile and self-sterile plants are distinct problems. Compton (1913)
found the relation between self-fertile and self-sterile plants of Reseda
odorata to be that of a simple Mendelian monohybrid with self-fertility
dominant. The same relation appears to hold in crosses between the |
self-fertile species Nicotiana langsdorffii and the two self-sterile species
with which our work has been done, Nicotiana forgetiana and Nicotiana
alata. ‘There is some single differential between self-fertility and self-
sterility. “Given the proper composition a plant breeds true for self-
sterility. The behavior of self-sterile plants among themselves
therefore must be considered separately.
Our work, as stated before, has been done with the two-self-sterile
species, Nicotiana forgetiana and Nicotiana alata, and largely with
crossés between these species. Both of these species are affected in
their manifestation of self-sterility by certain environmental changes,
Nicotiana alata much more than Nicotiana forgetiana. Self-sterility
is determined by the inheritance received, but it can develop fully
only under environmental conditions which promote a normal healthy
growth, and during the period of intense flowering. Toward the
end of thesflowering period, especially under conditions adverse to
vegetative growth, self-sterility sometimes shows a marked and rather
sudden decline. A few seeds, or even a well-developed seed capsule
may then be obtained. This.is nota common occurrence; indeed, it
146 BROOKLYN BOTANIC GARDEN MEMOIRS
is rare, but it is a possibility. Three cases of seed production out of
over three hundred plants tested have been observed in Nicotiana
forgetiana. A considerably higher percentage of fertility has been
marked in Nicotiana alata. Self-sterility can be restored in such
plants, however, if they are allowed to go through a period of rest and
are then, by proper treatment, brought into vigorous flower again.
This is not the whole evidence that this occasional end-season
fertility is a pseudo-fertility brought about by external conditions—
a fluctuation. Three generations of Nicotiana alata plants have been
grown from selfed seed produced by end-season fertility without the
occurrence of a single plant which behaved in every way like a truly
| self-fertile individual. This phenomenon, therefore, while teaching
' us to test self-sterility only during the main part of the flowering
season, has shown that there is no reason why fusion between gametes
produced by a self-sterile plant may not occur_provided the male
generative nucleus enters the embryo sac. Such unions may take
“place without affecting the self-sterility of the progeny.
What is then the difference in behavior that makes a cross-pollina-
tion effect fertilization while a self-pollination produces nothing?
What occurs is this: After a self-pollination the pollen grains germinate
and the tubes pass down the style at such a slow even rate that they
reach only about half way to the ovary before the flower wilts and
falls off; while the pollen tubes after a cross-pollination, though
starting at the same rate as the others, grow faster and faster until
fertilization is effected in four days or less. The curve of distance
traversed plotted against time is in the case of the self-pollination
nearly a straight line, while in the case of the cross-pollination it
simulates that of an autocatalytic reaction.
From these facts it seems reasonable to suppose that the secre-
/tions in the style offer a stimulus to pollen tubes from other plants
rather than an impediment to the development of tubes from pollen
of the same plant. And we believe that this stimulus is in some way
caused by certain effective differences in the factorial composition
characterizing two compatible plants.and that if two plants do. not
have these effective differences in factorial composition they are by
the same token cross-sterile with each other. It is clear that this
assumption presumes that the pollen grains matured by a given plant
behave as if they are sporophytic as regards that part of their con-
stitution that affects self-sterility and cross-sterility. The pollen
grains of any plant may carry many different hereditary factors, they
may even carry several different factors which function in controlling
the success or failure of particular cross-matings in the next generation,
but in their own action on the stigmas of other plants they behave
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 147
as if each carried the composition of the mother plant from which it
came. In other words, as far as its action in fertilization is concerned,
a pollen grain partakes of the character of its.mother-plant_and_is_
like its sisters; as far as the hereditary.characters carried-on_to the
next generation aré concerned, sister pollen grains may differ both
from their mother and from each other.
A part of our evidence on these points we shall present. For
further details the reader isreferred to a forthcoming paper in Genetics.’
The first experiment to which attention is called is an inbreeding
experiment performed on a cross between Nicotiana forgetiana and
Nicotiana alata. If sister plants are mated in successive generations
after an original mating Aa & Aa, by Mendelian recombination there
results a gradual approach to 1/2 AA, 1/2 aa and o Aa. Expectation
of homozygosis in successive matings is I/2, 5/8, 11/16, 24/32 --- I
(Jennings, 1916). If, therefore, plants of like constitution as far as
effective factors are concerned are cross-sterile with each other,
cross-sterility should become more and more apparent in generations
succeeding F,. To test this possibility, a comparatively small number
of cross-matings was made on the Fo, F3, Fy and Fs; generations. In
the F. generation, out of 131 intercrosses on 20 plants only 4 were
unsuccessful. The percentage of unsuccessful matings increased from
this time on, until in the F; generation about 21 percent of the cross-
matings tried on 20 plants were impossible to make.
In this experiment as well as in all others, results.showed that
reciprocal crosses were alike in their “compatibility. If two plants
Were fertile together, they were fertile reciprocally; if two plants
were incompatible, they were incompatible reciprocally. This is
proof of the sporophytic behavior of the factors affecting the behavior
of self-sterile plants.
The two crosses to be described next are reciprocals made with the
same two individuals. Made with Nicotiana alata and Nicotiana
forgetiana as parents, they are in a sense repetitions of the cross just
described, but it is hardly probable that they duplicate it. Both of
these species must consist of plants which differ among themselves in
the factors which affect self-sterility, hence any crosses in which
different individuals are used may show different results.
All of the individuals resulting from this cross were grown in a
reenhouse as potted-plantss The F, generation came into blossom
during the latter part of the winter. Conditions were extraordinarily
favorable for growth and the pollinations were all made while the
plants were vigorous, hence scarcely any trouble arose over classi-
fication of the results through end-season pseudo-fertility.
*This paper has since appeared. See “Studies on Self-sterility I. The
Behavior of Self-sterile Plants.’’ Genetics 2: 505-609. 1917.
148 BROOKLYN BOTANIC GARDEN MEMOIRS
Our study was made on a population of 53 plants. Pedigree
numbers from 0 to 39 inclusive represent the cross NV. alata « N. for-
getiana; pedigree numbers 40 to 52 inclusive represent cross JN. for-
getiana X N. alata.
Each plant was selfed_one or more times, and all proved abso-
lutely self-sterile. Further each plant was back-crossed with pollen
from a single plant of each of the parent species with complete success
in every case. The plants used in this case were not the individuals
that entered into the cross, however, for unfortunately these were
not available.
TABLE I
RESULT OF MATINGS ON F, PLANTS 0 TO 39
NV. alata X N. forgetiana and on Plants 41 to 52 UN. forgetuana X N. alata
Ped. No. Fertile with Ped. No. Sterile withPed. No.
Ownarre: BAN AOL ae coe ats eee RAE ae ogae eh aa ae ae | 22, 34, 38, 49
1 acter Per el OP, Nl We ee aacle teat, ta See bd bei eer 8
2 ae AP LG SAU SAA 5D er nets areas Creuegeds, Mes pS ete 0,222,223
Bearer ack 2 OyGLA 12 3\ DOr a. caenehe coors. cherie, enc eae te ode 4, 6, 18, 41, 46
Bia tile ce ZOE TO -AA\. vs chewed seats aes A Norse sera te oR Te eyes 18
etary i 2 SION O; LOPS TA OF Gene 9or oy Senet eee 8, 44
(Oey Soe e Tye lO Poy kc Wher: Ws Name AA hy ARR oo Ores creTTO Renee, Oia ce 3; 4,18; 40
Ty. oclawiies PUB D2 AAS. « Ravectysue atin ath costae ante rorvrenete aerators 18, 46
(ohn Hee oats OF Oy LON 395140; AGra coaah cis sence eh serena nee tere de 5, 44
QUE BY TBs A Ar PS Det rharatap tee tok Ate ohare a crate eee Sev eae ef 2, 10, 23; 37qae
LORAr is < AAO WUS,, AO TAA Wale cere token case Gee et eee teat eee 2; 23, 24, 27,034,140
1 a Nee 2 Tore, U5 a4 AAO
Loe ee OPLG; 522) Aer Peay tieie enc OAC eas i cmeunes RL ICIS = 6, 18, 46, 52
Eo ere BAST SO. AA AGr tart pun emia porch te coentne aii tie 2,9). 15, 2a
WA reece DS DO; 43 [rs cies ae dhen Meteo owe Ree vc ete ge Ns eee 10, 34
Litas Te LOn Te LO 20h es cities eneee cee thine rete ames x 9, 13, 14, 23, 44
LOnpaeys MAAS LOS 25, (ABSA Orcas oe one Ten ee ae eel 17, 29
17 ees WATS; 20% 225208 cee tte cit ae eG tee ne 16, 26, 44
Re tnee aa “é DROW 23 NOON ade BO mA nt ry seen wa reise mses 3, 46
TOs cee TJ R2 2 ADO BAS HAL wre ieee, 8 sie e ond aeteceunicveuetucie Sete 18
20. earn Deis (Oooh key its aA Alo weap Koy iik 5 aN oe Oe 43
21 eee APPEQN TOPRNO td Over are aces ete ot ae acy ae eee 2, 9,/22;.25, 270
PIT Eee EOF A SrA ire age ot Mea test AERO. oS AERA rn, sear Oa Re 14, 23, 24, 36, 48
Oa eae ASR an sgh het aaah stele camect ocean cope een One Ait Aen 9, 10, 37, 48
DAN eens BMONRZO a2 Om 2s sAA nanos ea ee Teng ae ee Te Ane LO; 22; 235-s0"ei
PA he CA S53 SrA ASA Oyaet An AEE PEO Ae Le ae oho eee Rie ea 26.0) 23) 27
2) Sa ree Q) ALSO ho sek AOMAG In cata po ects earn, tree 28, 29, 44
747 fea aero 3) LOGS 2 cA AWA Orie many ee hal ees eee es 2, 9, 30, 34, 48
2B WA. By Bee Se SOO 74 One kcmeee ey re Tite ihe Oe. ANT 8, 26, 29, 44
20 suis o. 2 TAS LG 22523, 24125, 30m a4 a7 Laas. 5, 26, 28, 31, 44
BOs. 85729;.9395744s 4 5y AC etn. oe ee eee ace O21, 22527
Se ee 225 B22 ruc yaie ies aeav See RE ate ORR aT ears oie reek 8, 29, 36, 44
22. ahaa hls Dy 215285729; 30s 34 cid 554 Are en ewer enn RE ee 18, 33, 46
S25... Oy EO 22s AGe sh een eee Nee a ERE ae 18, 32
BAMA 5.) 2B FAT Ay. 4.0 Sucve cy eich morgen tae aie. Perot ee eee LO; 23; 245537
Cte ae 8). Op lo els 27 nO a4 sy ce eral Cnet ee 8
Kile ey pope ie bie bs Pine aa ae A TR See we Ue TR 10, 23
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 149
Gyo rey EYRE DOI S GO WAP TAS AO cpe aise. seckehed sielerehon) teteote lence Cees Aly)
jl lrercterene OF Ee Nic eae EEO Se PCR CO IS SOREN Oe REPCRA CSP amar’ Sop rtike 18, 40, 42
m0... BoA ERA AT AQIS cls 5 cole he's ihe atts nega 6, 33, 46
EES aca ea MOP Qa Ae AG aN sera Neots ait SAME Sede = PCPA NER 33, 40, 46
(LENE Beate DOA mere et ety se Wass Fade) OS. cite coe t aay s 39, 41, 45
43..--.. 5, 27, 33, 38, 39, 40, 42, 44, 46, 51
44...... 10, 14, 23, 34, 45
1 a eee lek (il Reichs tapi rth Menai Bic cWAeuO IO Becpur bade Or on ohare 46, 52
AGBerscines : LO po? SANG SMO eons ts ay keene we Mtgose sie horse) Aue 52
17/2 eee BOCADNAAT AS MAGS 5 PPS Ob rac teenie lta iy Heya oO
POR Io is eeroasinns tary wena ene TO 22 2A od
a0), AQAA CASE cid cass apoio mote ce ee ee aoe thee On OW e703 4547)
ESF tals s PSASO a5 E Sete ces aia eaedeae se eee tae eens 9, 27, 37
eae si « OPS 52353 Ose 4s Opn Ore seamen eeeyerg yreuse cee st cs 8, 29
Eee. << TON 23 20302705 Lecchah ha caste iteae ns crt eeen ee BL rit sie 3, 4, 6, 18, 41, 45, 46
The numerous cross-matings made are shown in Table 1. There
were 103 reciprocal matings. Of these 100 gave duplicate results, 39
pairs being fertile and 61 sterile. The three which did not check are:
2 X 3, sterile, 1 pollination
3 X 2, fertile, 1 pollination
6 X 52, fertile, 1 pollination
52 X 6, sterile, I pollination
37 X 21, fertile, 1 pollination
21 X 37, sterile, 1 pollination
I classed as fertile,
classed as sterile,
classed as sterile.
Since but one pollination was made in each of these cases we have
\ made our decision as to fertility or sterility by a consideration of the
‘circumstantial evidence. The behavior of these plants in other crosses
shows conclusively that 3 should be fertile with 2, 6 sterile with 52,
and 21 sterile with 37. They have been classed accordingly. That
this grouping is correct is further shown by the fact that the mating
3 X 2 (classed fertile) was made at the height of the flowering season,
while the matings 6 X 52 and 37 X 21 (classed sterile) were re-
spectively the last and next to the last matings made on those plants.
In spite of the fact that plants 0-39 are from cross N. alata & N.
forgetiana, and plants 40-52 are from cross NV. forgetiana X N. alata,
they behave as one family in intercrosses. The entire population
intraclass sterility. The following explanation may be necessary to
make it clear just how Table II was obtained from Table I. Table I
shows all of the matings, but in the form given it is not easy to see at a
glance every combination in which a particular plant was used, both
as male and as female. It was necessary, therefore, to make a new
table, in which the pedigree numbers in the column at the left were
tabled as males, and the pedigree numbers in the columns headed
“Fertile matings’? and ‘Sterile matings’? were tabled as females.
150 BROOKLYN BOTANIC GARDEN MEMOIRS
Thus plant 2, used as a female, was fertile with pollen from plants 4,
18, 41, 44 and 52, and sterile with plants 9, 22 and 23; but pollen
from plant 2 was fertile on plants I, 3, 4, 5, 7, 11, 18, 20, 28 and 29,
and sterile on plants 9, 10, 13, 25 and 27. It is clear, therefore, that
instead of the 8 matings on plant 2 that Table I appears to show,
there are really 21, the 3 reciprocals of course being counted but once.
These tables were combined for analysis. In the interest of
economy of space only one is shown, however, since the second can
easily be made from the first.
The four exceptions in this huge set of matings are in reality
negligible. Matings 15 X 44 and 31 X 36 were sterile, though they
do not belong to the same class. Plant 15 was sterile to 4 plants of
Class A and fertile to 2 plants of Class B, 3 plants of Class C, and to
the isolated individuals forming classes D and F. It is unquestionably
a member of Class A. Plant 44 was sterile to 7 individuals in Class C
and fertile to 17 plants of Class A, 12 plants of Class B and to the
singletons forming classes D, E and F. This evidence places it un-
mistakably as a member of Class C. Plant 31 is also a member of
Class C as evidenced by 3 sterile matings within that class and by
fertile matings with 1 plant of Class A and 3 plants of Class B. Plant
36 is like plant 15 thrown into Class A by its sterility with 3 others of
that class, and by its fertility with 3 individuals of Class B, with 2 of
Class C, and with the lone plant of Class D. In view of this evidence
and the fact that in these two matings but one pollination was made
in each case, they are much more likely to be errors of record or of
technique than true exceptions to our classification.
The other two exceptions, matings 45 X 18 and 33 X 46, were
fertile where from the evidence of numerous other matings they should
have been sterile. Here again but one pollination was made in each
case; and, coincidence though it may be, each pollination was the last
mating made on that particular plant. What is more probable than
that this is a pseudo-fertility appearing during the wane of the flower-
ing season of the two mother plants, No. 45 and No. 33?
Six groups appear in Table II, but there is proof of the existence
of only five. Groups A, B, C, D and E£ are definitely established.
Plant 11, on the other hand, is an isolated individual rather than a
class. It does not belong to groups A, B or C; but unfortunately it
was not crossed either with Class D (plant 20) or with Class E (plant
43), hence one cannot say that it does not fall into one or the other of
these two classes.
In the three large groups the distribution of individuals is 22, 16
and 12. About all that can be said about the typeof this distribution
is that the classes are not of equal size. On the other hand, it is
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 151
interesting to note that the plants of both cross No. 2 and cross No. 3
fell into the three groups as if they were samples of the same popula-
tion. There were 40 plants of Cross No. I, and 13 plants of the
TABLE, U1
PLANTS OF F; GENERATION OF RECIPROCAL CROSS BETWEEN N. forgetiana AND N.
alata, GROUPED IN ACCORDANCE WITH THEIR BEHAVIOR IN INTERCROSSES
Plants 0-39 are products of the cross; plants 40-52 are products of its reciprocal
Cases Fertile in Group Cases Sterile in Group
Group Ped. No.
Galan | tee e BN GC Dr lem Wer
aS
os
bX
BN
| =e]
el
| =
|
|
COCO OOOO OOOO oO OOOO OO OW MINN POW TUN OUN HO OPAMN OW OP
Le | e+ eS Se |
| |
| |
i]
il Xey I Yertey tel (||
|
|
|
Ci
|
_
|
°
|
Pt
|
i
_
|
i S=t | ih fl
|
|
Pxeye ll ehe [|
|
a]
co
=
_
DOSOWMWOWWAMNWAHAAWHABNHAHAGCCCOCCCCCDCOCOQCACDOCQDGDADODACOO SO
as
lon
_
= | = | =e oe ee |
|
ter Mitek |) tes for tohion |i
|
NFP OHR RF OO COFCO RF COC FC OWWHMNOUNWAHBHR NN AAW NNANN OH
SOCOCHON ND NWWHWWWUO TTA NUN HF HHT NNWNHAMNOUNHANNWHWWH AU
by
=
on
WO
DAWNnFEUAUNHAH ANNWWNHFH WODCCACDCDCGDCCCCGCCCCCOOcCoO oo 8
Si Seon OLOVO OO OFvOvOrOtO ONO) (OO OO: Ol OO FiO) OC Od! ©) (ONONG) eS) ‘ONO OOO
|
°
| o
°
152 BROOKLYN BOTANIC GARDEN MEMOIRS
TABLE I1—Continued
Cases Fertile in Group | Cases Sterile in Group
Ss
B G D E dE
NS
by
G D EB FF
al
YN OWN NWWH AUN NF
as
f
Lol
NABuocOoOcooooo do
|
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|
(@) oJ (e) (©) Taie) Tite) ley (e) (eye)
(oy toy fe) 1s) (el fon elie} te) lon (2) fe)
COONN FP WN HW ND
|
|
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iHoOotHF I
|1ornrmnm |
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elle) tor |
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ies)
wn
WONUOBRPNNFODALWN
reciprocal, Cross No. 2. In the classes A, B and C the proportions
were 18, 10, 10 and 4, 6, 2 respectively. This similar behavior of the
progeny of reciprocals seems to us strong corroboratory evidence in
favor of the conclusion that reciprocal crosses always behave in like
manner as regards self-sterility. So oe ,
~The study on this family is but one of several that have been
made but we believe that the data on it alone show unmistakably that
the behavior of self-sterile plants in intercrosses is governed by a
relatively small number of factors which act through pollen as if the
pollen grain possessed the characters of the sporophyte from which it
jcame, and that the gametes of plants having like constitutions as re-
gards effectivé factors are incompatible in the sense that they do not
make a normal pollen-tube growth and hence do not reach the ovary
in time for fusion to occur. This interpretation shows both why
plants are self-sterile and cross-sterile. It accords completely with
the fact that a population of plants may be divided into groups on the
basis of their mating proclivities and that each member of any group is
cross-sterile with every other individual of that group although it is
fertile with every individual of every other group.
These assumptions being true, it ought to be possible by con-
tinuous self-fertilization, utilizing end-season pseudo-fertility, to
obtain ultimately a population in which every individual possesses
the same effective self-sterility factors. In such a population all of
the plants will not only be self-sterile, but will be cross-sterile. Sucha
population has been obtained.
EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 153
REFERENCES CITED
Compton, R. H. Phenomena and Problems of Self-sterility. New Phytologist 12:
197-206. I913.
Correns, C. Selbststerilitat und Individualstoffe. Festschr. d. mat.-nat. Gesell.
zur 84. Versamml. deutsch. Naturforscher u. Arzte Miinster i.W. pp. 1-32.
1912.
Darwin, Chas. Effects of Cross- and Self-fertilisation in the Vegetable Kingdom.
Bde 1878. N.Y. DD) Appleton. 1876,
Hildebrand, F. Bastardierungs Versuche an Orchideen. Bot. Ztg. 23: 245-249.
1865.
— Ueber die Nothwendigkeit der Insektenhilfe bei der Befruchtung von Cory-
dalis cava. Jahrb. wiss. Bot. 5: 359-363. 1866.
— Ueber die Bestéubungsvorrichtungen bei den Fumariaceen. Jahrb. wiss.
Bot. 7: 423. 1869.
Jennings, H.S. The Numerical Results of Diverse Systems of Breeding. Genetics
P53 —89. LOL:
K6lreuter, J. G. Vorlaufige Nachricht von einigen das Geschlecht der Pflanzen
betreffenden Versuchen und Beobachtungen, nebst Fortsetzungen I, 2 u. 3.
Pp. 1-266. Ostwald’s Klassiker, Nr. 41. Leipzig: Engelmann. 1761-6.
Morgan, T. H. Some Further Experiments on Self-fertilization in Ciona. Biol.
Bull. 8. 313-330. 1904.
Miiller, Fritz. Notizen iiber die Geschlechtsverhaltnisse brasilianischer Pflanzen.
Bot. Ztg. 26: 113-116. 1868.
— bBestéubungsversuche an Abutilon-Arten. Jen. Ztschr. f. Naturwiss. 7:
22-45, 441-450. 1873.
Munro, Robertson. On the Reproduction and Cross- fertilization of Passifloras.
Bot. Soc. Edin. 9: 399-402. 1868.
Stout, A. B. Self- and Cross-pollinations in Cichorium intybus with Reference to
Sterility. Mem. N: Y. Bot. Gard. 6: 333-454. 1916.
BINARY FISSION AND SURFACE TENSION IN THE
DEVELOPMENT OF THE COLONY IN VOLVOX
R: A. HARPER
Columbia University
In Volvox we have an incipient metaphyte with a many-celled body
of definitely organized form and the fundamental differentiation of
soma and germ cells fully established. Klein’s (’89, ’90) argument
that Volvox differs from true metaphytes in that the differentiation
of the germ cells does not take place until after cell division is com-
pleted is not very illuminating. The germ cells do not appear in
Oedogonium until a considerable series of undifferentiated cells have
been formed. Falkenberg’s comparison of the entire colony to a
zoosporangium also overlooks the very important fact that in the
development of the Volvox colony growth regularly alternates with
cell division (at least after the first few divisions) just as it does in
the development of the soma of one of the higher metaphytes. Vol-
vox is frequently referred to as one of the best known algae. There is
general agreement as to the order of cell divisions in the formation of
the colonies, both for the egg and the asexual germ cells. The litera-
ture has been frequently summarized.
To proceed further with the study of Volvox from the standpoint
of evolution and morphogenesis, we may compare it with such a
simple coenobe as Gonium. In addition to their more obvious struc-
tural characteristics there are two essential differences between the
colonies in such forms as Gonium and Volvox. First, the adhesion
between the daughter cells is much more firm in the latter. In
Gonium, as all observers testify, the adult colonies break up into their
component cells with the greatest readiness. I have figured such
broken-up colonies (’12, Pl. V, Fig. 23). Slight pressure, change in
the chemical composition of the medium in which they are, etc., lead
to almost explosive separations. One marked difficulty in getting
good photographs of the colonies is due to their tendency to go to
pieces. On the other hand, Volvox shows almost no tendency to
separate into its component cells. The colonies may be crushed into
formless masses without isolating a single cell and I know of no chemi-
cal or other stimulus which will cause their cells to fly apart as do
those of Gonium.
154
HARPER: BINARY FISSION AND SURFACE TENSION 155
The second difference lies in the very fundamental fact that, as
noted, in Volvox the germ cell grows to relatively large size before
dividing and the daughter cells grow in size between the successive
cell divisions. This is a very long step toward the full metaphytic
habit in ontogeny. It marks a return to the habit of the simple
protophyte like the bacteria and the appearance of a new point of
departure in the development of the morphogenesis of a metaphytic
plant body out of the primitive habit of reproduction by swarm-
spores which is seen in Chlamydomonas and Sphaerella. In these
protophytes, the cell having reached maturity forms from four to
eight swarmspores by rather rapidly succeeding divisions of the mother
cell. They escape by breaking of the mother cell wall and then as
free individuals proceed to grow to the size of the parent.
Swarmspore formation in Chlamydomonas and Sphaerella is a step
beyond the conditions in Euglena, for example, where in ordinary
reproduction each cell division is followed at once by the individualiza-
tion of the daughter cells and their independence as separate organisms.
In these particulars we may distinguish three steps in the evolution
of the metaphyte from the typical protophyte.
1. Cell division, in simple, direct alternation with growth, reproduction
and individualization, practically simultaneous and identical
processes. Euglena.
2. Cell divisions at unequal intervals, reproduction multiple and in
alternation with growth. Individualization delayed. Chlamy-
domonas, Sphaerella.
3. Cell division and growth in direct alternation, reproduction multiple
and individualization delayed by intercalation of a true em-
bryonic period. Volvo.
In Volvox individualization is already in essence the complex
process of differentiation and maturing which we find in the highest
plants and animals. In Gonium individualization of the daughter
colony, as I have shown in a former paper (’12), is accompanied merely
by certain gliding movements of the cells upon each other by which
an approximation to a least surface configuration is achieved so far
as is possible for sixteen ovoid cells arranged in a flat plate.
In Volvox, with the retention of multiple or colony reproduction
as in Sphaerella, we have growth intercalated again between each
successive cell division and also a specialization in function between
germ and somatic cells. Complete individualization is delayed till
the colony has become very many-celled. What may be called a
pseudo-growth comparable to the elongation of the cells just back of
the root tip by absorption of water and the formation of large central
vacuoles is also represented in Volvox by the formation from the cell
156 BROOKLYN BOTANIC GARDEN MEMOIRS
walls, as Meyer has most fully described, of the large masses of slime
by which the protoplasts in the adult colony are surrounded. This
slime, about whose nature Cohn, Klebs, Blochmann, and others have
differed so widely, is, as is now generally recognized, the gelatinized
cell wall comparable to the secondary thickenings in collenchymatous
tissues through which extend the broad strands which provide for the
again much disputed intercellular protoplasmic connections which
are so conspicuous in the adult colonies.
The firm adhesion of the daughter cells to each other and the re-
establishment of the primitive cell division-growth rhythm are two
further conditions to be reckoned with in the development of the
colony of Volvox as compared with that of Gonium.
We may turn now to the morphogenetic processes involved in the
reproduction of Volvox. Braun (’75) first clearly recognized the
“division by the wheel-forming type” in Eudorina as distinguished
from the ordinary successive bipartitions at right angles in Palmella,
etc. Braun refers the readjustments of rounding up and rearrange-
ment of the cells in forming the globular colony to the pressure of
the developing slime envelopes.
It has not been sufficiently emphasized that in the two-cell stage
and in the four-cell stage, as in the cleavage of the animal egg, the
halves and the quadrants respectively will tend by surface tension to
round up and give us a plate-shaped instead of a globular mass. In
Eudorina Goebel shows that the four cells tend to round up and are
shortened. He further represents them as tending to remain at one
end of the mother-cell cavity and to adhere to the surface of the
mother-cell wall. This leads to a divergence of their major axes and
(’82, p. 36, Fig. 17) gives already at this stage a polar opening. Over-
ton (89, Taf. II, Fig. 10, a, b, c) figures evidence of this divergence of
the four cells and it has been observed by others. The third division
by the wheel type, or radial type, gives us with the rounding up of
the cells a disk consisting of four interior and four peripheral cells
alternating with them, the familiar cross figure. The four inner cells
may appear much larger and are commonly so figured.
Overton (’89, Taf. II, Fig. 12, a and b) has shown very clearly
that the apparent relative size of the central and peripheral cells
varies with the level at which they are observed. The peripheral
cells have slipped out of the plane of the central four so that the
group of eight is already markedly concave. Two factors are pri-
marily concerned in this displacement. First, the fact that we have
by binary fission in two planes at right angles a group of eight rounded
bodies which can form no stable least surface configuration in one
plane and, second, the disk-shaped group formed by the rounding
HARPER: BINARY FISSION AND SURFACE TENSION 157
up of the eight cells by surface tension tends to conform to the shape
of the cavity of the mother cell. The’ whole is an expression of the
incompatibility of the principles of surface tension and binary fission
complicated still further by the rather firm adhesion of the cells to
each other. If division had produced seven instead of eight cells
and if they were free to adjust their interrelations in accord with their
capacity to achieve a position in which their pressure relations were
as nearly as possible mutually compensatory, we might have the
typical least surface group of one surrounded by six in one plane.
If ten cells were produced by division and if, as in Hydrodictyon, the
rounded form of the mother cell were a dominating factor we might
get one cell surrounded by five in the form of a saucer conforming to
the curved surface of the mother cell. A further series of five added on
the margin of the saucer and the figure could be closed by the re-
maining cell. If the eight cells produced by binary fission were free
as in Pediastrum, we might get a group like the typical eight-celled
colony of P. Boryanum with two inversely bilaterally symmetrically
placed central cells and two groups of three peripheral cells also
inversely bilaterally symmetrically placed with reference to each
- other (16). Pressure of the mother cell might make the group slightly
concave.
With eight cells produced by bipartition from four strongly ad-
herent mother cells and themselves rather firmly adherent the familiar
concave cross figure is the best approximation to a least surface
configuration.
I have noted that in Volvox there is growth of the daughter cells
intercalated between the divisions. This is very slight at first. In
the early stages, as has been generally noted, the mass of the young
colony seems little larger than that of the mother cell. In the prepara-
tion for the third division, however, there is a marked elongation of
the four cells.
In the two-celled stage the halves appear symmetrical, or one cell
may be slightly oblique (Fig. 1, Pl. II). In the four-celled stage the
sectors at first appear quite symmetrical and uniform in appearance
(Figs. 2, 3, Pl. I1) but with the preparation for the third division a char-
acteristic change in the form of the cells is observed. This growth
period intercalated between the cell divisions is an essentially differ-
entiating metaphytic character and makes possible in Volvox as in
higher types the formation of the indeterminately large and many-
celled colony as contrasted with the fewer-celled colonies of Gonium,
Pediastrum, etc., in which the cell-division stages are sharply separated
from the cell-growth stages. The growth in the four-celled stage of
Volvox is quite specific in that it is not a mere swelling of the cell in
158 BROOKLYN BOTANIC GARDEN MEMOIRS
all its dimensions. It results rather in a characteristic elongation in
one axis of the cell, the axes of elongation of the four cells tending to
be tangential to the general outline of the four-celled group. Klein
figures this elongation of the cells very clearly for Eudorina in the
four- and again in the eight-celled stage (’88, Taf. VI, Fig. 61, 63).
Overton shows it more crudely (’89, Taf. II, Fig. 10) and I have been
able to photograph it for one of the four cells (Fig. 4, Pl. II). It may
take place successively rather than simultaneously in the four cells and
apparently proceeds in either direction around the group. Biitschli’s
figure (83, Taf. XIV, 1 g.) shows rather crudely the resulting arrange-
ment of the cells just after the third division. The division seems to
be nearly simultaneous in all four cells and the wheel-formed group of
eight results. This characteristic growth and elongation of the cells
at this stage leads naturally to the oft-noted fact that the plane of the
third division cuts that of the second obliquely rather than at right
angles. A determining factor is, however, obviously the tendency to
bisect the elongated cell at right angles to its major axis as well as
the direct relation between the second and third cleavage planes. The
elongation of the cells during division reminds one at once of the
familiar elongation of the egg cell and other free globular cells at the
time when the bipolar karyokinetic spindle figure is at its climax of
development. We have no good figures of karyokinetie division in
Volvox but Overton’s figure from a 200-celled colony (’89, Taf. III, 18)
shows telophase stages with the cells all elongated and the spindles
in every case in the long axes of the cells.
It seems obvious that such a cell form in division implies a spindle
figure with polar asters and justifies the assumption that the same
internal forces are operating in the elongation of the Volvox cell as in
the dividing egg. Typical polar asters may be expected to be found
at such a stage as that shown in Overton’s figure, like those shown by
Swingle (’97) for a corresponding stage of division in Sphacelaria.
We may conclude then that the adhesion of the four mother cells
makes it necessary that the movement of material preparatory to
the production of two equivalent rounded daughter cells should take
place upon their free surfaces and the result is the characteristic
bulging and elongation of the four cells during division. That this
change of form is associated with the production of the karyokinetic
figure with two polar asters seems clear from the figures of division
in other alga cells with centrosomes. In any one of the cells of the four-
celled group (Fig. 3, Pl. II), for example, if the third spindle figure has
its axis 90° from that of the second division and in the same plane it is
obvious that one of the asters will not have space for its full expression
and if the adhesion of the quadrants is strong the yielding will be on
HARPER: BINARY FISSION AND SURFACE TENSION 159
the free surface of the cell and tend to give us the oblong cell form
shown in the figures. That elongation precedes or accompanies divi-
sion is indicated by the number of oblong cells in the older colony
shown in figure 7, plate II.
If the four cells should remain together and flattened upon each
other the successive divisions at right angles would give very variously
shaped and far from rounded daughter cells, as shown in Biitschli’s
diagram (’83, Fig. 1).
In effect, as viewed from the pole, the eight cells come to lie in an
up-and-down zigzag line instead of forming an in-and-out zigzag,
thus enabling each cell to remain as nearly as possible isodiametric
without reducing the compactness of the group.
It has been generally agreed that each of the eight cells of the wheel
figure divide to give a sixteen-celled stage and that the spherical or
ellipsoidal form of the colony may be achieved (as it obviously is in
sixteen-celled colonies of Eudorina) in this sixteen-celled stage.
Goroschankin (’75) held that the 16- and even the 32-celled stage
in Eudorina is a plate-shaped disk and that the transformation to a
globular form came rather suddenly with the gelatinizing of the cell
walls. He has not been followed in this view by later writers, though
they have very generally been inclined to accept his account of the
order of the cell divisions. Braun (’75), Biitschli (83), Overton (’89),
Klein (’90), and others hold that the spherical or rounded form of the
colony is practically achieved by the sixteen-celled stage.
The fourth division is then a binary fission of all eight of the cells
giving the sixteen-celled stage. The radial elongation of the four
central cells on their free surfaces is followed naturally according to
Hertwig’s law by their transverse division. (See Klein’s figure of
Eudorina, ’90, Taf. V, Fig. 63.) The four peripheral cells of the
eight-cell stage also elongate before dividing, just as in the four-cell
stage and divide transversely, giving the sixteen-celled group.
In Gonium the cells glide upon each other so as to form a series of
groups of three with the central square opening, as I have pointed
out elsewhere (’12). The greater adhesion of the four central cells in
Volvox and the elongation of the cells before division prevent the
Gonium configuration. The same situation develops as in the pre-
ceding third bipartition. There are eight new cells formed and these
tend to more than fill the space on the margin of the curved disk of
eight. Such a group is in very unstable equilibrium. The rounded
shape of the mother-cell cavity influences the direction in which the
cells. glide upon each other, and adhesion tends to develop the com-
pact groups of three. Of the eight new marginal daughter cells the
four coming from the original group of four form a part of the equa-
12
160 BROOKLYN BOTANIC GARDEN MEMOIRS
torial series, while the four coming from the peripheral four cells of the
eight-celled stage (Text-fig. 2) form a new polar group of four whose
cells alternate with those of the original polar group. Henfrey (’56)
described the sixteen-celled colony of Eudorina as consisting of two
polar groups of four and an equatorial circle of eight cells.
Biitschli (’83) describes the sixteen-celled colony as consisting of
four upper central cells, a ring of eight equatorial cells and four lower
cells which alternate with the upper and gives a figure (Taf. XLV,
Fig. 1, 2). Overton’s (’89) description of the arrangement of the
cells in the eight-celled stage is essentially like that of Biitschli,
though as Meyer notes, he ascribes the origin of the central cavity
of the colony to the divergence of the cells in the four-celled stage
and the subsequent hollowing out of the cells toward the center of the
Fics. I AND 2. For explanation see text.
colony. Neither of these authors gives any very clear account of the
arrangement of the eight equatorial cells or their relationship to the
polar groups.
It is not easy to obtain satisfactory photographs of these sixteen-
celled stages since, as noted, they are practically spherical and the
number of cells is so small that no characteristic groups can be ob-
tained in one focus. From a study of a large number of cases and
photographs of various views I have been able to obtain data for the
model made of marbles held together with wax and shown in two views
in text-figures I and 2. Text-figure I shows the anterior pole of the young
colony with the four cells forming a square, as has been observed by
all students of the group. The opposite posterior pole of the colony
would show a similar group of four, each cell alternating with the
four at the posterior pole.
HARPER: BINARY FISSION AND SURFACE TENSION 161
These are the relations of the two groups of four, as described by
Biitschli and Overton. As noted, it is with reference to the position of
the eight equatorial cells that clear description has been lacking. Text-
figure 2 shows an equatorial view of the model and it is obvious at
once that a least surface configuration requires the eight equatorial
cells to form a zigzag belt, each cell alternating with two cells of the
polar groups of four. With this arrangement the colony is made up
of eight groups of three cells, the most compact arrangement possible,
each of the four cells of each polar group appearing in two of the groups
of three. The groups of three are so placed with reference to each
other as also to form the groups of four seen in both the polar and
equatorial views. The whole colony would consist of eight of these
groups of four, since each of the eight equatorial cells is a member of
three such groups and each cell of the polar group is a member of
two such groups.
The whole forms as perfect an approximation to a least surface
configuration as can be achieved by sixteen cells arranged in as nearly
as possible a spherical group. Such a configuration may be regarded,
it seems to me, as the type configuration and illustrates the inter-
action of surface tension, adhesion and binary fission as morphogenetic
factors. In the actual colonies the cells are, of course, flattened upon
each other and, as I have pointed out, elongate during division. As the
photograph (Fig. 7, Pl. Il) shows, they are quite variable in both form
and size. As is shown clearly by Klein also in all figures of the so-
called polar openings, phialopores, the posterior group of four do not
form any such definite square group as is maintained by the anterior
group of four, yet the tendency to the formation of the groups shown
in the model is, it seems to me, obvious. The abundance of pentagons
and hexagons in the cell outlines of the adult colonies is good evidence
of a tendency to the most compact configuration possible.
I have not attempted to make the model with any great accuracy.
We lack data as to the relative efficiency of adhesion and surface
tension in the cells, which would be necessary for the exact determina-
tion of their interrelations. The actual relations of fission, adhesion
and surface tension in the processes just described are perhaps brought
out more clearly in the two diagrams (Text-figs. 3 and 4).
If the protoplasm of the mother cell were a mass which grows
merely by imbibition and swelling, and if the cutting up of the mass
into cells were merely a secondary phenomenon following the principle
of the rectangular intersection of the cleavage planes and surface
tension, as Hofmeister (’67), Sachs (’78), and other critics of the cell
theory have assumed, we should expect a configuration of the cells like
that shown in text-figure 4. This is the sort of configuration Magnus
162 BROOKLYN BOTANIC GARDEN MEMOIRS
(13) has obtained in his very interesting and suggestive experiment »*
with paraffine wax cooling over mercury, a configuration determined
entirely by the molecular forces operating in the system. It would
be possible to form a sphere out of these nineteen units, but there is
nothing in their number to favor the change from the discoid to the
spherical grouping. To make a hollow or solid sphere out of such a
group would involve very fundamental rearrangements.
On the other hand, if we make a diagram of the arrangement of
the sixteen cells produced by binary fission, assuming for the sake of
simplicity that each pair of daughter cells instead of remaining flattened
upon each other with the resulting lateral displacement (Klein, Taf.
VI, Figs. 61-63) rounds up completely after the third division and
that the four cells first formed remain fixed by adhesion, we get the
3 4
Fics. 3 and 4. For explanation see text.
configuration shown in text-figure 3. Here it is obvious that as con-
trasted with the arrangement in text-figure 4 it is a simple matter for
the group to become cup-shaped and spherical simply by folding in
the radial series and that the four outermost cells will come together
ina group of four with its members alternating with those of the
original group of four.
The group produced by binary fission in two planes with elongation
of the cells upon their free surfaces and strong adhesion tends naturally,
especially in the cavity of a spherical mother cell, to produce a globular
young colony at the sixteen-celled stage.
As noted, all authors agree in maintaining that all the cells of the
colony divide up to the sixteen-cell stage when the spherical form is
practically complete. The later stages have not been followed.
HARPER: BINARY FISSION AND SURFACE TENSION 163
Biitschli regards it as an open question whether all the cells divide in
the later stages. It is, however, obvious that successive bipartition
of all the cells is the natural method of maintaining the globular form
already achieved. Any excess or deficiency of the number of divi-
sions in any considerable group of cells would manifest itself at once
as a bulge or depression in the surface of the expanding sphere unless
compensatory divisions elsewhere and far-reaching, gliding move-
ments of the cells among one another were possible. There is no
evidence either of the occurrence or the possibility of such movements.
As has been many times observed, the daughters of the original group
of four can be recognized late in the life of the colony in their original
positions with respect to each other and the colony as a whole.
Oltmanns ('04) follows Goebel (’82) and Goroschankin (’75) in
asserting that this original polar group of four cells does not divide
after the sixteen-cell stage but does not give any very positive evidence
on the point. Overton (’89, Taf. III, Fig. 18) in a colony of about
200 cells shows a group of eight cells, six of which are dividing. It is,
of course, obvious that a failure of the original group of four to con-
tinue dividing after the sixteen-celled stage would not prevent the
maintenance of the rounded form of the colony in case there were
compensating divisions in the adjacent cells.
It is interesting to note that Kirchner (’79) finds the development
of the colony from the fertilized egg of V. aureus essentially like that
of the asexual germ cell. His figures give some indication of the
elongation of the cells before division and he describes the cup-shaped
form of the colony in the eight-cell stage.
It seems to me that we are justified in concluding that Volvox,
though showing deep-seated specialization of somatic and germ cells
in which it contrasts markedly with Eudorina, Pandorina and Gonium,
still like them shows vegetative totipotency and equivalence of its
cells in the growth of the colony. This is an important consideration
in view of the question as to the origin of the differentiation of germ
and somatic cells which is so conspicuous in the adult colony of Vol-
vox, and entirely lacking in the simpler members of the series. This
differentiation is such that the germ cells are distributed solely in a
single half or three fourths of the colony, the remaining portions re-
maining persistently somatic-sterile. The fertile area of the colony is
regularly the posterior half or three fourths as the colony swims.
It would be of great theoretical interest in this earliest appearance of
the differentiation of soma and germ plasm if it could be shown that
the cells bearing the germ plasm were different in cell lineage, age
since last division, relative maturity as indicated by total number of
divisions undergone, or in any other way, from the remaining cells of
164 BROOKLYN BOTANIC GARDEN MEMOIRS
the colony which show no capacity for reproduction and apparently
undergo senile degeneration. If, for example, as is so commonly
and loosely stated in textbooks, the colony were formed by marginal
growth and cell division, forming first a curved plate and finally a
sphere, so that the cells at one pole would be ontogenetically younger
than those at the other, we might expect this to be the basis for
differentiation of germ plasm and soma. The evidence is, however,
that all the cells of the adult colony are of the same generation and
ontogenetically equivalent.
The difference in their behavior is to be sought, then, in their
relative environment and internal development as the colony grows.
Their position in the posterior portion as the colony swims and around
the pole which is nearest the point of connection between daughter
colony and mother colony are obvious epigenetic factors in their
environment which may be of significance. The distribution of the
parthenogonidia at relatively equal intervals may be due to diffusion
phenomena affecting nutrition directly or as stimuli, the whole com-
plex perhaps suggesting analogy with Liesegang phenomena.
The attempt to differentiate the eight daughter colonies commonly
formed in asexual colonies of V. globator as descendants of the eight
cells produced by the third division seems to me wholly artificial.
This third division is not essentially different from the other divisions.
In V. aureus also the number of daughter colonies varies.
Meyer (95) notes that the protoplasmic connecting strands
between the cells are more numerous in this region of the germ cells
than in the anterior part of the colony. They are especially well
developed between the germ cells and the sterile cells as Janet’s
diagrams show so strikingly (12, Fig. 4). It is also in this region,
as noted, that the young colony maintains its connection with the
mother colony through protoplasmic strands from the cells around
the posterior polar opening which connect with the adjacent cells of
the parent as shown by Overton (’89, Taf. III, Fig. 16), and Janet
(12, Fig. 1). The germ cells are borne then in that region of the
colony which up to birth was most directly connected with the mother
colony and perhaps received from it a large amount of food materials
in the early growth stages.
The antheridia of V. globator form the so-called packets of anthero-
zoids consisting of bundles of sixteen to thirty-two gametes. These
are formed by binary fission of the mother cell in two planes. The
eight-cell stage shows the wheel figure. The cells instead of forming a
globular colony ordinarily form a flat plate like the simpler Goniwm.
Whether this is really due to a tendency to recapitulation retained in
the sexual germ-cell formation or whether it is due to the elongation
HARPER: BINARY FISSION AND SURFACE TENSION 165
of the male gametes as they form, is not clear, though it is generally
agreed that the cell arrangement in the bundle of male gametes is
homologous with that of the young vegetative colony.
Klein (’97) describes colonies of antherozoids in the form of hollow
spheres. Stein (’78) had also observed them. Chodat (’02) reports
plate-shaped individuals of Pandorina. These cases indicate how
closely the plate-shaped and the spherical colonies are related physi-
cally. The globular antherozoid bundles in Volvox may well be
regarded as tending toward the vegetative and away from the sexual
condition. That such changes should involve changes in surface
tension and adhesion is quite conceivable, as the elongation of the
antherozoid cell body must of course be a factor in determining the
form of colony. The further study of the formation of these abnormal
male colonies, especially, should throw light on the whole series of
morphogenetic problems here involved. The facts as known are
certainly quite in harmony with the view that such presumably easily
influenced factors as adhesion and surface tension, combined with the
more fundamental and ever-present incompatibility between the
principles of binary fission and least surfaces, may be of determining
significance in the transition from the plate-shaped to the three-
dimensional globular form of colony with all its evolutionary sig-
nificance.
INDEX OF LITERATURE
Cohn, F. Untersuchungen ueber die Entwickelungsgeschichte der mikroskop.
Algen u. Pilze. Nova acta Leop.-Carol. 24 (1): ror. 1854.
Henfrey, A. Notes on Some Fresh-water Confervoid Algae. Trans. Mic. Soc.
Eondon, N: S:, LV, p. 49." 1856:
Hofmeister. Die Lehre von der Pflanzenzelle. Leipzig. 1867.
Braun, Al. Bemerkungen zu Cohn’s Schrift ueber Volvox. Sitz.-Ber. d. Ges.
naturf. Freunde zu Berlin, Bot. Z. 190. 1875.
Braun, Al. Ueber einige Volvocineen. Sitz.-Ber. d. Ges. naturf. Freunde, Bot. Z.
189. 1875.
Cohn, F. Die Entwickelungsgeschichte der Gattung Volvox. Beitr. z. Biologie
der Pflanzen 3: 93. 1875.
Goroschankin. Die Genesis bei den Palmellaceen. Versuch einer vergl. Morpho-
logie der Volvocineae. Nachr. d. Kais. Ges. f. Naturw. usw. Moskau, 16.
1875.
Sachs, J. Ueber die anordnung der Zellen im jiingsten Pflanzenteilen. 1878.
Stein, Fr. v. Organismus der Infusionstiere. 3: 1. 1878.
Kirchner. Ueber die Entwickelungsgeschichte von Volvox minor. Cohn's Beitrage
3:95. 1879.
Goebel, K. Grundziige der Systematik und speziellen Pflanzenmorphologie. 1882.
Falkenberg. Die Algen. Schenk’s Handbuch, p. 284. 1882.
Biitschli. Protozoa. Bronn’s Klassen u. Ordn. d. Tierreichs. 1. 1883.
Klebs, G. Organization einiger Flagellatengruppen und ihre Beziehungen zu Algen
und Infusorien. Arb. d. bot. Inst. Tiibingen 1: 339. 1883.
166 BROOKLYN BOTANIC GARDEN MEMOIRS
Klein, L. Morphologische und biologische Studien ueber die Gattung Volvox.
Pringsh. Jahrb. 20: 133. 1889.
Klein, L. Neue Beitrage zur Kenntnis der Gattung Volvox. Ber. d. d. bot. Ges. —
7: A2. 1889.
Overton, E. Beitrag zur Kenntnis der Gattung Volvox. Bot. Centralbl. 39: 65.
1889.
Klein, L. Vergl. Untersuchungen ueber Morphologie und Biologie der Fortpflanzung
bei der Gattung Volvox. Ber. d. naturf. Ges. zu Freiburg i. B. 5: 1890.
Meyer, Arthur. Ueber den Bau von Volvox aureus Ehrb. und V. globator Ehrb.
Bot. Centralbl. 63: 225. 1895.
Swingle, W. T. Zur Kenntnis der Kern- u. Zellteilung bei den Sphacilariaceen.
Prings. Jahrb. 30: 299. 1895.
Kofoid, L. A. Plankton Studies. II. On Pleodorina illinoisensis, a New Species
from the Plankton of the Illinois River. Bull. Ill. State Lab. Nat. Hist.
5: 273. 1898.
R. Chodat. Algues vertes de la Suisse. Berne. 1902.
Oltmanns. Morphologie u. Biologie der Algen, Jena, 1904.
Harper, R. A. The Structure and Development of the Colony in Gonium. Trans.
Am. Mic. Soc. 31: 65. 1912.
Janet, C. Le Volvox. Limoges, 1912.
Magnus, W. Ueber zellenférmige Selbst-differenzierungen aus flussiger Materie.
Ber. d. d. bot. Ges. 31: 290. 1913.
Harper, R. A. On the nature of types in Pediastrum. Mem. N. Y. Bot. Gard. 6:
gI. I916.
DESCRIPTION OF PLATE II
All figures of asexual reproduction in Volvox taken with Zeiss apochrom, ob-
jectives and the compensating eye-pieces.
Fic. 1. Two-celled stage of young colony. X about 600.
Fic. 2. Second division just completed. X about 600.
Fic. 3. Four-celled stage of young colony. X about 350.
Fic. 4. Four-cell stage, one cell elongating preparatory to third division.
x about 350.
Fic. 5. Ejight-cell stage showing the wheel or radial arrangement of the cells.
x about 350.
Fic. 6. Same stageaslast. The young colony photographed after being teased
out of the mother cell. X about 400.
Fic. 7. Young colony not yet set free from mother colony and cell walls not
yet gelatinized. X about 400.
VOLUME I, PLATE Il.
BROOKLYN BOTANIC GARDEN MEMOIRS.
IN VOLVOX
BINARY FISSION
HARPER:
FURTHER STUDIES ON THE INTERRELATIONSHIP
OF MORPHOLOGICAL AND PHYSIOLOGICAL
CHARACTERS IN SEEDLINGS OF
PHASEOLUS!
J. ARTHUR HARRIS
Station for Experimental Evolution, Cold Spring Harbor, N. Y.
INTRODUCTORY REMARKS
In a series of papers published during the past several years I have
emphasized the importance of investigations of the relationship be-
tween the morphological and the physiological characteristics of the
organ and of the organism.
The structural variations of the organs of which the organism is
made up are the resultant of intrinsic and extrinsic factors—of heredity
and environment, or of nature and nurture. Morphogenetic processes
must, therefore, be investigated by physiological methods, and be
interpreted in physiological, and ultimately in physical and chemical,
terms.
The purpose of this paper is to supplement and extend the results
of an earlier study? in which it was shown that in bean seedlings char-
acterized by certain morphological variations from type, the develop-
ment of primordial leaf tissue is less than in normal controls grown
under conditions as nearly as possible identical. The data then
available indicated that a reduction of the volume of primordial leaf
tissue is associated with abnormalities of all the types studied, but
that the type of variation influences, in some degree, the amount of
reduction.
In these first experiments the conclusions were based on primordial
leaves only.
The use of such leaves has the obvious disadvantage that they
are completely formed in the seed, and undergo merely an enormous
expansion (and an undetermined amount of differentiation) in the
1 Studies on the Correlation between Morphological and Physiological Char-
acters, V. Studies I-IV of the series are to be found in Genetics 1: 185-196. 1916;
Peeloo—2i20) TOL7e2, 282-200, IOr7.
* Harris, J. Arthur. Studies on the Correlation of Morphological and Physio-
logical Characters: The Development of the Primordial Leaves in Teratological
Bean Seedlings. Genetics 1: 185-196. 1916.
167
168 BROOKLYN BOTANIC GARDEN MEMOIRS.
germination of the seed and the development of the plantlet to the
stage at which measurements were made.
Since the development of the primordial leaves during the germina-
tion and establishment of the seedling is relatively great, it seemed
quite legitimate to use the weight of green tissue produced by these
leaves as a measure of the physiological capacity of seedlings of various
types. The fact that these leaves are differentiated in the seed, does,
however, constitute a valid objection against their use as a measure
of the physiological capacity of the seedling. For such purposes a
constant based upon some organ developed later in the life of the
individual is desirable.
One of the purposes of this paper is to present the results of deter-
minations upon a later developed organ. The one chosen is the
first trifoliate leaf. Sa |
' This leaf was used because groups of plants of a higher degree of
uniformity can be selected at the time of maturity of this leaf than
at any later stage in the development of the plant, and because the
first compound leaf reaches a degree of maturity sufficient for the
purposes of the present study before the primordial leaves are too
old to be used for a series of determinations. It is, therefore, possible
to repeat, at a slightly later stage of development of the plant, the
determinations made on the primordial leaves in the first study as a
basis of comparison with the work already done and with the series
of constants to be obtained for the first compound leaves of the same
plants.
In the first investigation the green weight of the leaf tissue served
as the fundamental measurement. In addition to this character
certain measurements on the sap properties were also made. In the
study of the saps some difficulties were encountered, and it seemed
most desirable to discontinue that phase of the study temporarily
and to carry out determination of dry weight and water content
instead. These new measurements have, therefore, been added to
these for green weight.
MATERIALS AND METHODS
The materials upon which this study is based are the same as
those previously employed—a mixture of slightly different strains
of navy beans. The seeds which were germinated in the fall and
winter months of 1916 were grown in field cultures in 1915.
Seeds from individual plants were germinated in sand. In sorting,
the morphologically aberrant seedlings were laid aside with a normal
plant to serve as a check for each abnormal. An abnormal and a
control seedling from the same seed plant and germinated in the same
HARRIS: INTERRELATIONSHIP IN PHASEOLUS 169
seed flat were potted side by side in a three inch pot and allowed to
grow to the proper stage of maturity under conditions as favorable
as we were able to give them.
Before the samples were taken, the plants were carefully inspected
and all pairs, one member of which had died, had been injured or
which showed in its subsequent development any abnormality in addi-
tion to these specified were discarded. Note that there was no direct
selection for the characters of the abnormal plantlets in this process,
since both abnormal and control were discarded if either was unsuited
for the purposes of the experiment.
There probably was a fairly stringent indirect selection, since the
death rate and the mutilation rate of the variant individuals was
probably greater than that of the normals. Thus more pairs were
probably discarded because of an injury to or the death of the ab-
normal member of the pair than because of the death or injury of a
normal member.
The probability that the materials were somewhat selected before
the physiological measurements discussed in this paper were carried
out renders the findings of greater significance than they would
otherwise be.
After the pairs of seedlings had grown until the first compound
leaf- had attained its full size, and the second compound leaf was
developing, but before the primordial leaves had materially deteri-
orated, samples of leaves were taken by nipping off the laminae only,
or the laminae and the single petiolule of the terminal leaflet in the
case of the compound leaf. These samples of tissues, each from_
1oo_ plants, were enclosed in flasks, weighed, and dried to constant
weight 1 in a bath surrounded by boiling water.
Thus the. technique employed was exceedingly simple. Because
of the size of the samples dealt with, the relative infrequency of the
abnormalities, and the large number which had to be discarded, the
routine has been excessively laborious. For example, the weighings
of the 23 samples and checks discussed in the present paper involve
13,800 leaves gathered from 4,600 plants which were secured by
germinating and classifying nearly half a million seedlings.
The structural variation in the bean seedling which is probably the
simplest, and the most frequent, is a slight vertical separation of the
cotyledons which are normally sensibly opposite in insertion. The
amount of the separation is difficult to express quantitatively, since
it is in some degree dependent upon the length of the axis. In our
‘studies of seedling variation in Phaseolus, three grades of separation
of the ‘cotyledons have been recognized. The line of demarcation
between these grades is a quite arbitrary one. This is also true of
170 BROOKLYN BOTANIC GARDEN MEMOIRS
the line between ‘‘normal,’’ and ‘‘abnormal”’ as applied to the dis-
tinction between plants which have cotyledons inserted on the same
level and those which have one of the pair sensibly higher on the axis
than the other. “Slightly but distinctly separated,’ has been the
descriptive term used in our classification schedules. The cotyledons
range in position from those which are just perceptibly not inserted on
the same level to those which are perhaps two or three or four milli-
meters apart. So imperceptible is the line of distinction between nor-
mal and abnormal plants that in the classification of the seedlings
frequent discussions arose concerning the normality or abnormality of
individual plants.
In the present paper I am considering only the simplest type of
abnormality. This course has been followed for two reasons.
First, the proof of the existence of a physiological differentiation
associated with a very slight structural variation is of far greater in-
terest than the demonstration of measurable physiological differentia-
tion associated with great morphological variation. Second, other
types of abnormality with which I have dealt are so difficult to secure
in satisfactorily large series that the number of samples as yet avail-
able is not sufficient to justify detailed comparisons between the
different types of abnormality. I hope ultimately to be able to meet
these difficulties. For the present the one type of structural devia-
tion dealt with serves to illustrate the method and one phase of the
results of the investigations.
PRESENTATION OF DATA
Consider first of all the green weight of the organs selected.
The average green weight of the primordial and of the first com-
pound leaves for plants which are normal except for slight separation
of their cotyledons is shown in Table I.
With one single exception, the average weight of the primordial
leaves of the normal plants is higher than that of the abnormal plants.
In the single exception to the rule, the difference is small in amount.
The average weight of the first compound leaf produced by abnormal
plants of this class is in every case but one lower than the weight
produced by the sensibly normal individuals. The exception to the
rule is the same sample as in the case of the primordial leaves.
The average weight of primordial leaf tissue in the abnormal plants
is .5873, the average weight for normal plants is .6680, and the average
difference —.0807. The differences in mean weights range in the
individual samples from +.0074 to —.1286. For the first compound
leaf of the same plants the average weight of the tissues from abnormal
individuals is .4797, from a normal plant it is .5610, while the average
HARRIS: INTERRELATIONSHIP IN PHASEOLUS 171
difference between the sample and the control is —.0813. The differ-
ences in average weight vary from +.0368 to —.2492.
TABLE I
Mean Green Weight per Plant of Primordial Leaves and of First Compound Leaf
Primordial Leaves First Compound Leaf
papple Percentage) Percentage
Abnormal | Control Difference | Difference | Abnormal} Control | Difference | Difference
32 .6034 .7096 | —.1062 15.0 .5132 .5929 | —.0797 13.4
35 .5648 .6767 —-I1T19 16.5 -5444 .6188 —.0744 12.0
36 .5951 .6361 —.0410 6.4 -5931 162545) 20323) 5.2
39 .56019 .6277 —.0658 10.5 .5160 .5549 — .0389 7.0
40 .6096 O52) |= 00503 1350 .5179 .6138 | —.0959 15.6
41 .6068 -7304 E230 16.9 .4877 61405 | —.1263 20.6
42 .5879 6141 — .0262 4.3 .4712 -7204 | —.2492 | 34.6
43 .6222 -7508 —.1286 17.1 | .5008 6115 —.1107 | ~ 18.1
46 5956 -7160 —.1204 TO.8" | 4645 .6019 | —.1374 | 22.8
47 .7058 | .6984 | +.0074 II | .5841 | .5473 | +.0368 6.7
48 .6389 .7272 —.0883 12.1 -5593 -6395 —.0802 12.5
49 .5902 16674-0772 11.6 | .4960 65851 | —.0891 15.2
53 +5402 5990 | —.0588 9.8 -4491 4948 | —.0457 9.2
54 .5720 .6530 | —.08I0 12.4 .4091 4547 | —.0456 10.0
56 | .5380 .5921 —.0541 g.I .3994 .4646 | —.c652 14.0
Gi e513 .5827 — .0634 10.9 4443 .4811 — .0368 7.6
Of | 5853 .7052 —.1199 17.0 .4530 .5848 —,1318 22.5
OS) VW 25747 .6938 —TLOr Wee -4402 577 —, £315 23.0
66 .5886 .6790 | —.0904 mane .5246 .5960 | —.0714 12.0
70 .6853 .7066 —.0213 3.0 -4794 .4998 | —.0204 4.1
71 -5639 .6059 —.0420 6.9 4132 4534 | —.0402 8.9
72 25505 6744 | —.1179 1725 |) -3799 .4882 | —.1083 | 22.2
73. |_-5033 | .6140 | —.1107 | 18.0 | .3933 | .4887 |'—.0954 | 19-5
If these differences be reduced to percentages by using the weight
of the normal plants as a base, as shown in the final columns of each
section of the tables, it appears that the primordial leaves of the
morphologically aberrant plants are from 3.0 to 18.0 percent lighter
than the leaves of the normal plants in the 22 samples in which this
relationship between the two types of plants holds for the primordial
leaves. Thus the percentages are highly variable. The average for
the 23 determinations is 11.95 percent. In the case of the first com-
pound leaves, the percentage reduction ranges from 4.1 to 34.6 with
an average of 14.06 in the.23 samples. Note that the percentage
shows that the difference between the abnormal and the control sample
is far less in the case of the single exception, sample 47, than it is in
the average series. Thus it is only I.1 as compared with an average
_value of 11.95 for the primordial leaves and only 6.7 as compared
with the average of 14.06 percent in the compound leaves.
I now turn to a consideration of dry weight.
172 BROOKLYN BOTANIC GARDEN MEMOIRS
The primordial leaves of the abnormal plants in which the two
cotyledons are slightly separated are, as shown in Table II, lighter
TABLE II
Mean Dry Weight per Plant of Primordial Leaves and of First Compound Leaf
Primordial Leaves First Compound Leaf
Sample
, Abnormal | Control | Difference Beamer Abnormal | Control | Difference patties
32 .0445 .0537 | —-0092 70 .0442 105072 |) =-0075 14.5
35 .0366 SOAS Oly, 24.2 .0465 .0530 | —.0065 12.3
36 .0422 .0457 — 0035 Well .0476 .0499 —.0023 4.6
39 .0409 .0467 | —.0058 12.4 .0430 .0470 | —.0040! 8.5
40 .0438 .O511 —.0073 14.3 -O415 .0496 | —.0081 | 16.3
41 .0431 .0526 | —.0095 18.1 .0406 .0494 | —.0088 | -17.8
42 .0416 .0504 | —.0088 17.5 .0383 | .0519 — .0136 26.2
43 .0429 .0532 | —.O103 19.4 -0391 .0493 | —-0102 20.7
46 .0408 .O501 — .0093 18.6 .0400 .0492 — .0092 18.7
47 .0442 .0446 | —.0004 9 .0442 .0433 | +.0009 23K
48 | .0420 .0464. | —.0044 |; 9.5 -0444 .0525 —.0081 15.4
49 0381 .0436 | —.0055 12.6 .0397 (047210 — 0075 15.9
53 .0365 .O410 | —.0045 II.0 .0399 .0427 | —.0028 6.6
54 .0384 | .0445 —.0061 13.7 .0339 0412.5 |, =.0073°|) Aigeg
56 .0349 .0491 — .O142 28.9 0331 | .0395 ‘| —.0064 16.2
61 .0356 .0402 —.0046 A! 20383 | 20407 — .C034. 8.2
64 | .6354 C438 | —.0084 | 19.2 0341 | .0435 | —.0094 | 21.6
65 .0357 .0410 | —.0053 12.9 0344 | .0398 | —.0054 | , 13.6
66 .0354 | .0395 —.0041 10.4 0381 | .0439 | —.0058 13:2
70 .0426 .0465 | —.0039 | 8.4 0407 | .0438 | —.0031.| — 7-8
7ilee a \eeO27.9 .0303 | —.0024 | 7.9 .0274. 0299, | —.0025 |? (8-4
72 .0273 .0407, | —.0134' 32.9 .0265 £0392 || —.O127, | .2
72) 0208 .0378 | —.0080 21.2 .0315 .0408 | —.0093 |- 22.8
than those of the normal controls in every instance. The average
dry weight of the abnormal is .0382, that of the control .0452 and the
average difference is —.0070 grams. If the differences be expressed
as a percent of the control value as a base, they range from less than
I to nearly 33 percent, with a general average of 15.21 percent.
The results for the first compound leaf are very similar. In 22 of
the 23 cases the primordial leaves of normal plants yield a greater
weight of dry substance than those of the abnormal plants. The
exception to the rule is again sample 47. The average dry weight
of the first compound leaf of abnormal plants is .0385, that of normal
plants is .0452 and the average difference is —.0067 grams. If the
differences be expressed as percentages of the control constants they
are seen to range from 3.2 to 26.2, for the 22 series in which the ab-
normal plants produce a smaller amount of dry substance. The differ-
ence in the single exceptional series is small, only 2.1 percent as com-
pared with a general average of 13.36 percent in the 23 samples.
HARRIS: INTERRELATIONSHIP IN PHASEOLUS 173
Having shown that the abnormal plants produce both a smaller
green weight and a smaller dry weight in both the primordial and in
the first compound leaves, the problem of the relative quantities of
water and dry materials in the leaves of the two types of plants
naturally presents itself for consideration.
The results have been expressed in terms of the percentage of dry
substance in the leaves, 7. e. (dry weight X 100)/green weight. The
constants appear in ‘Table III.
TABLE III
Percent of Dry Matter in Primordial Leaves and in First Compound Leaf
Primordial Leaves First Compound Leaf
Sample l
Abnormal | Control Difference | Abnormal Control Difference
32 mes7as | &7.507 — Oe 8.612 8.703 — ,OO1
35 6.480 | TELAT = Moz 8.541 8.564 —".023
36 ee OoL J e7- Los — 003 8.025 7.978 + .047
39 | 7.278 7.439 = ui 8.333 8.469 = AS
40 | 7-185 | 7.246 — .06r | S073 8.080 — .067
41 7.103 e201 — 098 | 3.324 8.045 Se eae
42 7.076 8.207 Teak ain 8.128 7-204 | + .924
ae | 6.804 7.085 = OL” | B7280g 8.062 | — .255
ae | = 6.850 G.097* "| —_ 147 8.611 8.174 + .437
el) (6.262 6.386 | — .124 7.567 7.912 SI AGG
48 6.574 6.381 + .193 | 7.938 8.210 SS aE
49 | 6.455 6.533 — .078 | 8.004 8.067 | — .063
53 6.757. | 6.845 | — .c88 | 8.884 8.629 + .255
54 femO7zls | OS LS = oe 8.286 9.061 ifs
56 6.487. | 8.293 —1.806 | 8.287 8.502 .215
61 6.855 6.899 — .044 | 8620 | 8.668 — .048
64 61048). |) G21 — .163 | 7.528 7TAZSG | ==> G90
oo) 6.272. |" 5.909 + .303 | 7.815 6.962 + .853
66 Georg | 5-517. +} * size G7 7.263 | 7.300) S| 2. —- 103
wae) 6.216 "| .6.581° | = .265 7)? “Bi490 8.764 | — .274
fee A048 | 5.001 |° — .053 | 61631 6.595 + .036
72 4.906 6.035 | —I.129 | 6.976 | 8.029 SLOSS
WS mRo2r | 6.156 | 7235) S000) Vy areas — .340
green weight and dry weight. This condition is to be expected for
two reasons. First, the abnormal plants show lower values of both
green weight and dry weight than the normal controls. One cannot,
therefore, expect such large differences in the indices calculated from
these constants as if both measures did not differ in the same direction
between abnormal and control series. Second, two sets of technical
operations are involved in the indices, only one in each of the con-
stants used in calculating these ratios. While every effort to avoid
error was made, the probabilities of error in an index are clearly
twice as great as in either of the constants upon which it is based.
174 BROOKLYN BOTANIC GARDEN MEMOIRS
Notwithstanding these two sources of difficulty in basing con-
clusions on relative amount of dry substance, there seem clear evi-
dences that the abnormal plants produce relatively as well as abso-
lutely less dry matter than the normals. era’
In the case of the primordial leaves, there are 20 samples in which |
the relative dry weight is lower in the abnormal plants as against 3 in|
which it is higher. In the first compound leaf there are 15 samples_
in which the relative weight in the abnormal plants is lower, as com-
pared with 8 in which it is higher than the normals.
The average percentage content of dry substance in the primordial
leaves of the abnormal seedlings is 6.509 as compared with 6.779 in
the normal controls, or a difference of —0.270. The average percent
of dry matter in the first compound leaf is 8.030 in the abnormal as
compared with 8.080 in the normal, or a difference of —.o50 percent.
CONCLUDING REMARKS
The constants recorded in this paper are the results of one of the
phases of an attempt to determine the nature of the relationship
between morphological and physiological variations in plants.
The results of the criteria applied are beautifully clear and con-
sistent.
Seedlings of Phaseolus which show one of the smallest definite
structural variations, the slight vertical separation of the two coty-
ledons in their insertion on the axis, are differentiated from the struc-
turally apparently normal individuals in their physiological as well
as in their morphological characteristics.
This is shown by the facts that the cabulolceieule abnormal
plants produce a smaller weight of green leaf tissue, a smaller actual
weight of dry substance in the leaf tissue, and a smaller relative weight
of dry substance. This is true for both the primordial leaves and
the first trifoliate leaf.
AMERICAN HEATHS AND PINE HEATHS
JOHN W. HARSHBERGER
University of Pennsylvania
One of the attempts of modern phytogeography and ecology has
been to establish an exact nomenclature and to correlate the existing
knowledge of the fundamental units of vegetation the world over.
The thought of the leading phytogeographers has been to limit the
use of descriptive terms to an exact meaning, following the lead of the
earlier morphologists, who, like Linnaeus, made an exact science of
morphology out of a chaos, or jumble, of inexactly applied descriptive
terms. The earlier morphologists had somewhat of an advantage,
because they were applying names to parts of plants which had no
recognition as such by the laity, and where in common usage, the
concept of such words as bract, carpel, ovule, and the like, had no
application. The phytogeographers, however, find that their con-
cepts parallel those of the botanically uninitiated, who refer in every-
day speech to forest, to meadows, to prairies, to marshes, to swamps
and to heaths. Elsewhere! the writer has shown that country people
have a keen appreciation of the fundamental differences in the native
vegetation for they have applied names in a general and unscientific
way to plant formations recognized by ecologists. Graebner? has
emphasized this fact in his “ Die Heide Norddeutschlands.” If the
phytogeographer adopts a local descriptive term he should attempt to
strictly limit the use of the term to the same unit of vegetation. If
this is done, then there can be no objection to the adoption of the
descriptive name from the common speech of the people, who roughly
distinguish a certain association of plants. Professor Diels on the
other hand considers the use of vernacular names in plant geography
very questionable. He maintains that such terms are ambiguous,
even in the language to which they belong, that to persons of foreign
birth they are either meaningless or liable to misunderstanding, that
even if such terms are strictly defined, they will become confused
again, and that they are permanently confusing to people unfamiliar
1 Harshberger, John W. The Vegetation of South Florida. Wagner Free Inst.
of Sci. Trans. 73: 146. 1914. The Vegetation of the New Jersey Pine-barrens,
Dis, 1016.
2Graebner, Paul. Die Heide Norddeutschlands. Die Vegetation der Erde
Pema leipzig. 901.
13 175
176 BROOKLYN BOTANIC GARDEN MEMOIRS
with phytogeography. He believes that there should be universal
expressions in Latin, or Greek, and to have these alone. I can heartily
agree with the general opinion of Diels about the necessity of a stable
nomenclature in plant geography, but it would be unwise to abolish
vernacular terms, even if these are used with some confusion. As
teachers in the class-room, in our published papers, in our conversa-
tion and in our encyclopedic work, if called upon to contribute articles
to dictionaries and encyclopedias, we should try and clarify the ideas
of the public on these essential points.
For example, the word forest is a nomen confusum. In its use, in
England, a forest may signify any wild, open, uncultivated tract of
land, not necessarily a tract of woodland, though historic documents
prove that parts of the ancient forests of the British isles were covered
with trees. The term forest in the United States fortunately is applied
more exactly and properly to a tree-clad area. The same confusion
is seen in the application of the words swamp, marsh and moorland.
The natives of the island of Nantucket, and the visitors who have
learned the name from the habitant, call typic heathland by the cog-
nomen moor, and similarly in England, where the word heath is in
common use, it is applied very inexactly. Heath to the Britisher is
usually a heather-clad tract of land, yet in eastern England, the word
is also used to denote a calcareous pasture, with no heather, as New-
market Heath and Royston Heath, and in Somerset, it is used to
designate tracts of deep and often wet peat.
As the research investigations of the writer have led him to believe
that certain types of vegetation in America correspond with the true
heath and pine-heath of Europe, it becomes necessary to see if we
can correlate the different usages of the word heath so as to unsnarl
the tangle into which the use of the word seems to have fallen.
Tansley in the introduction to ‘‘ Types of British Vegetation ” (p. 2)
states that heathland nearly always involves a relatively poor and dry
soil. Under the climatic conditions of the British Isles, heath is found
on shallow, dry, peaty soils dominated by the common ling (Calluna
vulgaris) and occurs in regions of medium rainfall in the center, south
and east, and on similar sandy soil in Belgium, Holland, Denmark and
northwest Germany. The surface of the soil of such heaths is covered
with dry peat (Trockentorf) with the general absence of deep peat.
Where in hollows of true heathland with an impervious substratum,
true moor peat is found, heath passes imperceptibly into moor, and
hence there has often been confusion of the two kinds of phyto-
geographic concepts.
In ‘Types of British Vegetation’’ (pages 98-99) is given a state-
ment as to the character of the heath formation of northwest Europe.
HARSHBERGER: AMERICAN HEATHS AND PINE HEATHS 177
It is typically developed on relatively poor sandy and gravelly soils,
whose climate is wetter than that which gives rise to steppe, the
climate of which is too dry for tree growth. Heath may exist side by
side with woods and may represent a degeneration of woodland.
Heath occurs in Europe in regions with an annual rainfall between
25 and 40 inches (60 to 100 cm.), but the Cornish heath and those of
the eastern Highlands of Scotland often receive a rainfall of between
40 and 60 inches (100 to 150 cm.) in the year. The Scottish heaths
develop a deeper layer of relatively pure acid humus, up to 8 or 12
inches (20 to 30 cm.), according to Hardy. The East Anglian heaths
have a rainfall of 25 inches, or less, and a minimum of dry peat forma-
tion, while the heaths of the southeastern counties have a layer of dry
peat seldom more than a fraction of an inch in thickness passing down
into sand darkened by humus. The surface layer of dry peat is
formed by lichens and mosses, which are pioneers on denuded soils.
Drude in his comparison of the flora of Great Britain with that
of Central Europe?’ believes that the lowland heaths, the ‘heath asso-
ciation” or ‘‘Callunetum arenosum”’ of Tansley, for the most part
correspond with those of northwestern Germany in the region of the
Weser and the Ems, and on the English heaths one would often feel
oneself transported to Germany, if it were not for the sudden oc-
currence of Erica cinerea between Erica tetralix and Calluna vulgaris,
or of Ulex minor, or Ulex gallii with masses of Schoenus nigricans,
Myrica gale, Narthecitum and Hypericum elodes, which indicate the
west European conditions.
If there is a physiognomic similarity in the heathland of England
and northwest Europe, then we must determine the essential character
of the heath vegetation and the kinds of soils on which it is found, for
by extension we can apply these characters as a test of heathland in
other parts of the world.
Although the soils of the North German plain are the same in dhe
east as in the west, according to the researches of Graebner and others,
yet the vegetation of the two areas is quite distinct. In the west, in
Hanover, Oldenburg and Schleswig-Holstein, are great stretches of
heathland, whilst in the east these are entirely absent and are replaced
by thin pine woods (pine-heath = Kiefern-heide) and a steppe-like
flora. This difference is due in part to the different climate, for the
main heathland is west of the Elbe, where the rainfall rarely falls
below 24 to 28 inches per annum, whereas in the east the rainfall often
does not exceed 20 inches per annum. In other words, heathland is
developed with an oceanic climate, while pine-heath is found where
the climate is continental. The seasonal changes of temperature of
3 The International Phytogeographic Excursion in the British Isles (1911), p. 93.
178 BROOKLYN BOTANIC GARDEN MEMOIRS
an oceanic climate are slight. The relative humidity is higher. There
is a larger amount of cloudiness and a heavier rainfall than is found
over continental interiors. Marine climate is equable. With a
continental climate the annual temperature ranges increase, as a
whole, with increasing distance from the ocean. The regular diurnal
ranges are also large. The winters are cold and the summers are hot.
Cloudless skies are more frequent and the air is dustier and drier.
In Europe, where heath plants grow, there is a formation of fibrous
and slightly earthy humous layers, the undecomposed elements of
which are deposited in dense masses to form the so-called ‘‘ vaw-humus,”
which in some cases is deposited as a thin peat layer. The humic
Fic. 1. Nantucket heath with pearly everlasting (Anaphalis margaritacea)
and bayberry (Myrica carolinensis). August 20, 1914.
acid, according to Ramann, acts on the unweathered silicate, de-
composing it energetically, bringing into solution alkalis and alkaline
earths, leaches the soil, 7. e., the soluble substances are carried down
to greater depths. If raw humus lies on sandy soils, the sand of the
upper layers appears bleached and becomes light gray in color, being
in this condition called lead sand. Below this light-colored layer is
found a yellowish to brownish soil due to the grains of sand being
mixed with ferric-oxid or ferric hydrate. The precipitated organic
substances cement the separate sand grains into compacted layers
HARSHBERGER: AMERICAN HEATHS AND PINE HEATHS§ 179
below the lead sand and meadow ore, and a humus sandstone is formed.
This is called Ortstein and forms an impervious layer. The roots of
pine trees are unable to penetrate this layer and flatten out on it.
The pine trees become stunted, their branches gnarled and the tree
finally dies to be replaced by others which pass through the same cycle
unless the tap-root manages to pass through a hole into the subsoil.
The phytogeographic investigations of the writer have shown that
on the island of Nantucket we have true heaths, called moors by the
Fic. 2. Bear oak (Quercus nana) in heathland, Nantucket. September 7,
IQI4.
inhabitants. Typic heather plants grow on the sandy and gravelly
glacial soils of the island and there is a formation of dry peat on the
surface of the soil. The low, rolling hills are covered with plants in
different groupings, so that in places the western prairies are suggested,
while in other places the Roman Campagna, especially where the
vegetation has been browsed by cattle. The plain-like character of
the vegetation is suggested in those areas where the beard grass
(Andropogon scoparius) covers the ground in exclusive growth, except
where patches of heath plants are found such as the huckleberry
(Gaylussacia resinosa) and the bearberry (Arctostaphylos uva-ursi)
(Figs. 3 and 4) which forms mats, or carpets of large extent. Another
common plant, which suggests the Scotch heather (Calluna vulgaris),
180 BROOKLYN BOTANIC GARDEN MEMOIRS
is a plant with bright yellow flowers (Hudsonia ericoides), while with
it we find associated the grayish patches of the reindeer moss (Cladonta
rangiferina), hair moss (Polytrichum), goat’s rue (Tephrosia virginiana),
extensive masses of trailing arbutus (Epigaea repens), and patches of
Anaphalis margaritacea (Fig. 1). Clumps of the bayberry (Myrica
carolinensis) of a dark green color break the monotonous level of the
heath (Figs. 1 and 3). Patches of an irregular rounded form of the
huckleberry (Gaylussacia resinosa) and sweet fern (Comptonia aspleni-
folia) are common. A few other flowering plants relieve the flat
Fic. 3. Granite boulder (crow-stone) surrounded with carpet of bearberry
(Arctostaphylos uva-ursi), Nantucket heath. August 20, 1914.
green tones of the rolling surface, such as the wild indigo (Baptisia
tinctoria), golden aster (Chrysopsis falcata) and white-topped aster
(Sericocarpus asteroides).
Some parts of Nantucket, notably the southeastern, are covered
by two low oaks, namely, the bear oak (Quercus nana) (Fig. 2), and
the dwarf chestnut oak (Quercus prinoides). In some places on
Nantucket and in central Marthas Vineyard, this growth of oaks
and associated plants suggests the dwarf elfin wood, or chaparral, of
the California coast. The other constituents of these low oak thickets
are the bayberry (Myrica carolinensis), sweet fern (Comptonia aspleni-
folia) and the low spreading carpets of the bearberry (Arctostaphylos
HARSHBERGER: AMERICAN HEATHS AND PINE HEATHS 181
uva-urst) (Figs. 3 and 4). The growth of low oaks found over a
large part of Nantucket and the central part of Marthas Vineyard
may be looked upon as an oak-heath, because many of the associates
of Quercus nana and Q. prinoides belong to the heath family. The
soil conditions are such that a dry peat is formed over a subsoil of
sand and gravel, just as we find in the heathland of European countries.
The island of Nantucket is also characterized by the presence of
the broom-crowberry (Corema Conradii), which grows rather plenti-
fully in the central part of the island (Fig. 5), and along Tom Never’s
Bluff. It forms on Nantucket round-headed clumps, which vary in
Fic. 4. Bearberry (Arctostaphylos uva-ursi) spreading over a denuded gravel
slope, Nantucket. August 20, 1914.
size from a few feet across to many feet in diameter. Along Tom
Never’s Head, owing probably to the undermining of the bluff by
the action of the surf, we find the plant mingling with the true beach
plants of the upper sea beach. In New Jersey, the most local and
peculiar plant of the Lower and Upper Plains is the broom-crowberry
which grows in two general forms. The first type of plant is one
which grows in dense cushions (Fig. 6) about 3 decimeters (1 foot)
tall, its color varying from light green through dark green to a rich
brown color, distributed in clumps between the prostrate pine trees.
That this is the typic form is indicated by the fact that it is the type
182 BROOKLYN BOTANIC GARDEN MEMOIRS
found on Nantucket and elsewhere. The second type is the diffuse
one, where the stems are scattered widely.
In New Jersey, Corema Conradii (Fig. 6) is associated with rounded,
basket-like, dwarf pitch pines (Pinus rigida) along with bearberry
(Arctostaphylos uva-ursi), huckleberry (Gaylussacia resinosa), sweet
fern (Comptonia asplenifolia), trailing arbutus (Epigaea repens) and
the like, so that this type of heath I have called a Corema]. Some
facts may be learned about the vegetation of the New Jersey Coremal
by contrasting it with the Coremal on the Island of Nantucket. The
physiognomy is slightly different, owing to the absence of the dwarf
Fic. 5. Cushion of broom-crowberry (Corema Conradit), central part of Nan-
tucket. September 5, 1914.
pines in the true Coremal of Nantucket. In both districts, there is a
low vegetation of shrubby oaks, Quercus nana and Q. prinoides. The
bearberry (Arctostaphylos uva-urst) is more abundant on Nantucket
(Figs. 3 and 4) than on the plains of New Jersey, while the broom-
crowberry is found in both districts (Figs. 5 and 6). A comparison of
the Nantucket vegetation gives a clue to the origin of the plains
(Coremal) and the pine-barren vegetation of New Jersey. Heath-
land is the result of the factors which are summed up under the general
term oceanic climate, and Nantucket, isolated far out at sea, has an
oceanic climate. The strong winds that blow and the pervious
HARSHBERGER: AMERICAN HEATHS AND PINE HEATHS 183
glacial soils prevent the deciduous trees of large size from spreading
out of the valleys, where they are protected, and even the pitch-pine,
recently introduced into Nantucket, follows the valleys and protected
slopes of the hills. In all probability, when the pine-barren region of
New Jersey was an island, an oceanic climate prevailed. The typic
heathland of which the Coremal is a part was left as a relict in the
plains of New Jersey. The pine forest in those early times probably
filled the valleys and later spread over the hills until all of the region
was covered with pine forest except the areas represented by the
Upper and Lower Plains, where edaphic conditions prevented the
growth of tall pine trees. The heathland of this early time was
Fic. 6. Broom-crowberry (Corema Conradii) along cart road, Warren Grove,
N. J. In flower, April 7, 1917.
finally converted into the present pine-heath. Graebner has detailed
a similar conversion of heath into pine forest by the invasion of
pines, and in such German pine forests the undergrowth consists of
characteristic heath plants, hence the term Kiefern-heide applied to
this pine forest with an undergrowth of heath plants.
The pine trees have been unable to grow to tall size in the New
Jersey Coremal because of a-hard layer of soil immediately below the
upper sandy layer (Fig. 6). This layer corresponds to the caleche of
Mexico, the plow-sole of agriculturists and the Ortstein of the Germans.
184 BROOKLYN BOTANIC GARDEN MEMOIRS
Klebahn‘ figures an Ortstein Kiefer where the tap-root striking the
hardpan is bent over, being unable to penetrate that soil layer.
Graebner narrates how such pine trees grow for a time, but finally,
after reaching a certain age, begin to go back, or decline in vigor, until
they succumb, and he describes how certain pine trees more fortu-
nately situated by natural planting over holes through the Ortstein
(Ortsteintépfe) are able to send their tap-roots into the deeper soil
layers.» Under such conditions tall thrifty pine trees will be scattered
here and there over the surface of the heathland, while the majority
of the trees, that become established in the region, are dwarf and
Fic. 7. Pine-barrens (pine-heath) near Lake Ronkonkoma, Long Island.
July 20, 1913.
languishing. Similar conditions are found in the plains of New
Jersey where the low, dwarf pine trees live for a number of years and
finally succumb, to be replaced by other trees that pass through a
similar existence. Hence the dwarf basket pines of the New Jersey
Coremal are all short-lived. Thus hardpan and fire are the two most
important factors which have perpetuated the heath vegetation of the
New Jersey plains (Coremal), while the surrounding region with more
‘Klebahn, H. Grundziige der allgemeinen Phytopathologie, p. 14.
*Graebner, P. Die Heide Norddeutschlands. Die Vegetation der Erde
5: 125.
HARSHBERGER: AMERICAN HEATHS AND PINE HEATHS 185
pervious soil, although similarly fire-swept in later years, has been
preserved as a pine forest, or pine-heath (Kiefern-heide). Remove
the pines and the conditions as they exist in the Coremal of Nantucket
are duplicated. The New Jersey pine-barrens with the removal of
the pines represent such an oak-heath as we have described for the
islands of Marthas Vineyard and Nantucket and of similar physiog-
nomy with such oaks as the bear oak (Quercus nana) and dwarf chest-
nut oak (Quercus prinoides) forming the main ground cover.
Similarly, as in Germany, the pine trees in the Long Island and
New Jersey regions have become dominant and the heath plants in
Fic. 8. Rounded clumps of pine-barren heather (Hudsonia ericoides) in full
flower growing one mile south of Shamong, N. J. May 27, 1916.
the form of the bearberry (Arctostaphylos uva-ursi), sand myrtle
(Dendrium buxifolium), huckleberry (Gaylussacia resinosa), laurel
(Kalmia latifolia), and the oaks become subordinate to the pines and
form the characteristic undergrowth of the pine forest (Figs. 7 and 8).
Graebner distinguished several types of heath woodland, as follows:
1. Type. Pine-heath. 4
Facies a. Pine-heath with dominance of Juniperus com-
munis.
Facies b. Pine-heath with dominance of Rubus species.
*Facies c. Pine-heath with dominance of Arctostaphylos uva-
urst.
186 BROOKLYN BOTANIC GARDEN MEMOIRS
*Facies d. Pine-heath with dominance of grasses.
*Facies e. Pine-heath with dominance of Vaccinium myrtillus
and V. vitis-idaea.
*Facies f. Pine-heath with bog-moss substratum.
2. Type. Broad-leaved Tree-heath.
Facies a. Birch-heath.
*Facies b. Oak-heath.
Those facies of the forest-heaths in Germany which are similar
physiognomically with the ones in New Jersey are marked with an
asterisk. Facies e in Germany, with the prevalence of two species
of Vaccinium, is represented in New Jersey by a pine forest with an
undergrowth of Gaylussacia resinosa, Kalmia angustifolia, Vaccinium
pennsylvanicum and V. vacillans. The oak-heath we have described
and also the heathland, where the bearberry is common, as on Nan-
tucket.
We are able, therefore, by this comparative study to correlate
certain American plant formations with those of Europe. The species
of plants represented in each are in general different, but physiog-
nomically the contrast can be made with general correctness.
America p Europe
Heathland (with huckleberries, blue- Heathland (Heide, with heather, etc.).
berries and bearberries, etc.).
Oak-heath (with low oaks, etc.). Oak-heath (Eichen-heide).
Coremal (heathland with broom-crow- Low pinc-heath (Kiefern-heide).
berry in Nantucket with addition
of dwarf pine on plains of New
Jersey).
Pine-barrens (Long Island and New Pine-heath (Kiefern-heide).
Jersey, incipient on Nantucket
(Figs. 7 and 8)).
Much remains to be done in the study of the phytogeography of
America, but we have reached a stage in our investigations where it
is profitable to compare the American plant formations with those of
Europe and other parts of the world. This comparison leads to a
clarification of our concepts and also brings about a correlation of
our knowledge, so that it is possible to formulate certain principles
upon such comparative study. The three international phyto-
geographic excursions have done much to stimulate this kind of
comparative investigation, where the studies of botanists in other
climates and in other countries can be used for the extension of our
knowledge of the fundamental principles upon which phytogeography,
ecology and plant physiology depend.
SOME BOTANICAL PROBLEMS THAT PALEOBOTANY
HAS HELPED TO SOLVE
ARTHUR HOLLICK
Museum of the Staten Island Association of Arts and Sciences
Among the savants whose investigations in connection with the
vegetable kingdom have earned for them the distinction of having
their names graven on the walls of the laboratory building of the
Brooklyn Botanic Garden are four to whom paleobotany lays claim,
at least in part—Adolf Théodore Brongniart, Leo Lesquereux, Oswald
Heer, and Marquis Gaston de Saporta—and it is my privilege briefly
to outline some of the botanical problems which they and other paleo-
botanists have helped to solve.
Paleobotany, as a science, is much younger than botany. Living
plants were noticed, studied and classified long before fossil plants
had received any serious attention. In fact it was only a little more
than a century ago that fossil plants began to be recognized as the
probable actual remains of former living plants.
The science of paleobotany may be said to have been born in 1804,
when Ernst Friedrich, Baron von Schlotheim, issued his Beschreibung
Merkwiirdiger Kraiiter-Abdriicke und Pflanzen-Versteinerungen: ein
Beitrag zur Flora der Vorwelt, generally known and cited as Schlot-
heim’s Flora der Vorwelt, in which he discusses the prevailing theories
in regard to the nature and origin of fossils, or “ petrifactions’’ as they
were commonly called at that time, and uses the following epoch-
making words: ‘“‘. . . and more recent observations and investigations
have even led us to the very probable supposition that they may be
the remains of an earlier so-called pre-Adamic creation, the originals of
which are now no longer to be found. . . . In the continued investiga-
tion of this subject this opinion, with certain restrictions, has in fact
gained a high degree of probability with the author of the present
work, so that he ventures to announce his treatise as a contribution
to the flora of the ancient world.’’
In the light of what we know and take for granted today this
statement sounds strangely elemental in connection with a work of
that nature; but in reality it represented an expression of the most
advanced thought of the period when it was issued, and to Schlotheim
should be given the credit for having laid the foundation upon which
187
188 BROOKLYN BOTANIC GARDEN MEMOIRS
Brongniart and his contemporaries and successors erected the super-
structure which gradually developed into the science of paleobotany
as we now recognize it.
The first great basic fact, therefore, that paleobotany proclaimed
was that our living flora had an ancestry whose elements were different
from those now in existence. As facts accumulated, and the floras of
the successive periods in the earth’s history became better and better’
known, the phylogenetic development from low and simple types of
vegetation to successively higher and more complex ones was demon-
strated, and a rational, philosophical basis for systematic botany was
established. Before that time any system of taxonomic arrangement
of the vegetable kingdom was purely theoretical. Relationships
were recognized, but they were often lacking in explanation; and it
is significant that every real advance which has been made in tax-
onomy has been in accord with our constantly increasing knowledge
of phylogeny. Paleobotany is thus constantly helping to solve the
broad problem of the why and wherefore of our modern systematic
arrangement of the vegetable kingdom and rendering it more and more
truly scientific.
If certain of the apparent anomalies in modern taxonomy are
critically examined in the light of paleobotanical knowledge they
become anomalies no longer. As an example we may consider the
case of a monotypic genus such as Ginkgo. It is represented by a
single living species, G. biloba L., and its nearest affinities among living
coniferae are apparently with the Taxaceae. What is the meaning of
its isolation in our scheme of classification? Does it represent a recent
development of a new generic type in connection with which new
species are destined to be evolved in the future, or does it represent
an ancient type of vegetation of which it is now the sole survivor?
Paleobotany has supplied the answer to these questions by demon-
strating that the genus was formerly represented by many species and
that it was merely one of a number of allied genera all of which are
now extinct.
The genus Sequoia, with its two living species, is a similar, although
not quite so striking an example of generic isolation; and in the same
category may be mentioned Nelumbo, Liriodendron, Sassafras—each
represented by but two species—and Liquidambar by three. All of
these represent vanishing generic types as evidenced by the many
known fossil species in each, now entirely extinct. It was, of course,
reasonable to infer that such was the case in these and other similar
instances; but paleobotanical discoveries alone furnished the definite
proof.
It is, however, within the domain of what we broadly designate as
HOLLICK: BOTANICAL PROBLEMS AND PALEOBOTANY 189
the science of ecology, and especially in connection with the problems
of phytogeography, that the work of paleobotanists has been of un-
expected value, in furnishing explanations of many puzzling facts of
modern plant distribution. In this connection we may hark back to
certain of the genera already discussed taxonomically and consider
them in relation to their present geographic distribution.
The genus Liriodendron is represented by one species in eastern
and middle North America and one in eastern Asia. The two species
of Sassafras have the same distribution. Of the two species of
Nelumbo one has a range in America extending from New Jersey to
Colombia and the other is Asiatic. What is the meaning of the
occurrence of only two species representing each genus and each of
the species in such widely separated regions? We can not imagine
that a genus could originate two specific types independently, each one
in a different part of the world, even in a single fortuitous instance;
and it is almost as difficult to believe that a genus could originate in
Asia and develop a single species which somehow subsequently mi-
grated to America and there evolved into a different species, or vice
versa. Discussions of the possibilities of evolution, mutation, and
migration afforded theoretical but unsatisfactory explanations. The
discoveries of paleobotany, however, supplied actual facts, and these
showed that in all such instances the genera were formerly world-wide
in their distribution as well as prolific in species. A single example in
this connection is sufficient. Fossil remains of some twenty-five
species of Nelumbo have been brought to light, from the United
States, British America, Greenland, England, Holland, Germany,
Hungary, France, Portugal, Egypt and Japan. The problem of the
modern distribution of any such genus in two widely separated parts
of the world, therefore, has nothing to do with any phenomena of
evolution, or mutation, or migration in modern times. It is merely
a matter of elimination of species in past times, throughout the inter-
mediate regions where they formerly flourished. On the same basis
may also be explained the geographic isolation of Sequoia with its two
living species confined to a narrow belt on the western slope of the
Sierras in California, and Taxodium with its three living species con-
fined to the coast region of the eastern and southern United States and
the northern part of Mexico. The discoveries of paleobotany have
demonstrated that in past ages both of these genera included many
species and that they were widely distributed. They flourished not
only in similar latitudes to those in which they now occur, but also
northward beyond the Arctic circle as far as exploration has been
carried. The climatic conditions of the Ice Age exterminated them
everywhere in the North. The mountain systems of the Eurasian
190 BROOKLYN BOTANIC. GARDEN MEMOIRS
continent, extending in an east and west direction, formed barriers
which prevented their migration southward and there they became
extinct. In North America, however, with its mountain systems
extending in a north and south direction, migration to more congenial
regions was possible and here they continued to exist. Their present
isolated geographic distribution was, therefore, determined long ago,
by a combination of climatic changes and topographic features, and is
not a modern phenomenon that can be satisfactorily explained by
present conditions alone.
Incidentally it may also be pertinent to recall that the genus
Sequoia enjoys the unique distinction of having been found in the
fossil form previous to its discovery as an element in our existing
flora. Cones and leaf-bearing twigs, representing what we now know
as the genus Sequoia, were found in Europe and were described and
figured (but not, of course, under the modern generic name) before
the living trees on our western coast were discovered. This fact,
however, can hardly be cited as an instance in which paleobotany has
been of assistance to botany, inasmuch as it involves the question in
nomenclature whether or not the generic name first applied to the
fossil remains should have precedence over that subsequently given to
the living trees.
I shall not attempt, in this paper, to discuss the debt which botany
owes to those paleobotanical students who have made special studies
of.the internal structure of fossil plants, and thus determined exact
botanical relationships along lines of modern morphological investi-
gations. This is a relatively recent phase in the development of
paleobotany and the results attained are familiar to us all. The dis-
covery of the extinct class or order of plants, the Pteridosperms or
Cycadofilicales, and its taxonomic relations to the Pteridophytes and
Gymnosperms, is due to their labors, as is also the determination of
the exact affinities of many extinct families and genera with those now
living. They have filled in the details of the broad phylogenetic
sequence outlined by the earlier paleobotanists and they represent
the field of work in which botany and paleobotany are most closely
and intimately related today and in which it is impossible to dissociate
them.
FURTHER NOTES ON THE STRUCTURAL DI-
MORPHISM OF SEXUAL AND TETRASPORIC
PLANTS IN THE GENUS GALAXAURA
MARSHALL A. HOWE
The New York Botanical Garden
At the meeting of the Botanical Society of America, held at Colum-
bia University in December, 1916, the writer presented a short paper,
since published,! in which evidence was brought forward to show that
Galaxaura obtusata, a calcified red alga of the family Chaetangiaceae,
presents itself in two forms of the same general habit, but differing
markedly and constantly in the microscopic structure of the cortex.
In the one, the middle layer of the cortex consists chiefly of large
chambers, more or less filled with lime in the natural condition, while
in the other the chambers are comparatively small and the cortex is
pseudoparenchymatous throughout. There are also other differences
in the form and relations of the cortex cells, as pointed out in the
paper to which reference has been made. In a monograph of the
genus Galaxaura, published by Kjellman in 1900,? these characters
were made the basis of two groups of species, denominated by him the
“Cameratae’’ and the “Spissae.’’ In the recently published paper,
the present writer drew attention to the facts that plants showing
the ““Cameratae’’ and ‘‘Spissae’’ structure are commonly collected
together throughout the West Indian region, that they show the
same or parallel variations in external habit, that they can not, in
fact, be separated without a microscopic examination, and that, when-
ever reproductive organs can be found, the plants of the ‘‘Cameratae”’
structure are always tetrasporic, while those of the “‘Spissae”’ struc-
ture are always antheridial or cystocarpic. The writer therefore
expressed the conviction “that the ‘Spissae’ and ‘Cameratae’ char-
acters, first accurately pointed out by Kjellman, do not offer a proper
basis for subgeneric groupings of species as supposed by him,” but
merely distinguish the sexual and tetrasporic phases in the life-cycle
of a single species.
Since reading and publishing the short paper to which reference
1 Structural dimorphism or sexual and tetrasporic plants of Galaxaura obtusata.
Bull. Torrey Club 43: 621-624. 10 Ja 1917.
* Kjellman, F. R. Om Floridé-slagtet Galaxaura, dess organografi och sys-
tematik. Kongl. Sv. Vet.-Akad. Handl. 33!: 1-109. pr. I-26. 1900.
14 191
192 BROOKLYN BOTANIC GARDEN MEMOIRS
has been made, the writer has been investigating some of the other
sections of Galaxaura as proposed or recognized by Kjellman in his
monograph of the genus and as accepted without question by writers
on the red algae during the past seventeen years, and grounds have
been found for believing that a similar relation exists between several
other pairs of groups hitherto considered to be independent. The
Fic. 1. Cortex of a tetrasporic plant of Galaxaura marginata (Ell. & Soland.)
Lamour, in section, illustrating the cortex structure of the section ‘‘ Brachycladia”’;
enlarged about 210 diameters. (After Bgrgesen.)
Fic. 2. Cortex of antheridial plant of Galaxaura marginata (Ell. & Soland.)
Lamour., in section, illustrating the cortex structure of Kjellman’s section “‘ Vepre-
culae,’’ enlarged about 300 diameters. (After Bgrgesen, under name Galaxaura
occidentalis Borg.)
Fic. 3. Figures showing the essentially free filaments, short and long (bases
only of the long), constituting the cortex of Galaxaura flagelliformis Kjellm., and
illustrating the cortex structure of Kjellman’s section ‘‘Rhodura’’; enlarged about
154 diameters. (After Kjellman.)
Fic. 4. Cortex of Galaxaura squalida Kjellm., in section, illustrating the cortex
structure of the section ‘‘ Microthoé’’; enlarged 210 diameters. (After Bgrgesen.)
HOWE: DIMORPHISM IN GALAXAURA 193
evidence associating the groups ‘‘Vepreculae’’ and ‘‘ Brachycladia”’
seems particularly convincing. The name ‘‘Vepreculae’’ was given
by Kjellman to a “‘section’’ of the genus in 1900. ‘“‘ Brachycladia”’
was proposed by Sonder as a separate genus in 1853 and is recognized
as an independent genus by De-Toni in his ‘‘Sylloge Algarum,”’ though
by Kjellman it is properly considered to represent a section of Galax-
aura. In the “Brachycladia”’ group, the cortex is essentially fila-
mentous, as to its two outer layers at least (TEXT-FIGURE I), and the
cells separate easily after decalcification, though forming a more or
less coherent epidermis in the natural calcified condition. The outer-
most or superficial cells are usually oval or ellipsoid and obtuse or
apiculate. In plants of the section ‘‘Vepreculae,” the cortex (TEXT-
FIGURE 2) may be said to be parenchymatous or pseudoparenchyma-
tous rather than filamentous. The epidermis here consists of cells
that are firmly united both before and after decalcification and these
cells have their longest axis parallel to the general surface instead of
at right angles to it. In some parts of the thallus, especially at the
edges of the flattened branches, the surface shows few or numerous,
scattered or crowded, blunt or apiculate, papilla-like cells, which are
probably homologous with the outermost or epidermal cells in the
“Brachycladia”’ section, though they do not here form the epidermis,
the firmly united epidermal cells of the ‘‘Vepreculae”’ section being
probably homologous with theewidely spaced subepidermal stalk-cell$
of the ‘“‘Brachycladia”’ section. Now, an examination of a wide
series of plants of the ‘“‘ Brachycladia”’ structure from the West Indies,
as well as an examination of the type material of nearly all of the
species from various parts of the world referred to this section by
Kjellman, indicates that whenever reproductive organs are found,
the plants of this group are always tetrasporic, and, in the same way,
plants showing the ‘‘Vepreculae”’ structure are always antheridial or
cystocarpic. Moreover, in the West Indies, at least, the writer’s
personal experience in collecting shows that plants of these two types
of cortex-structure often occur together and that they show the same
or parallel variations in external habit. They resemble each other
very much in size and habit (PLATE III; PLATE IV, FIGURE 1), but may
usually be distinguished under a hand-lens, if not at sight, by differ-
ences in the texture of the epidermis, that of the ‘‘ Vepreculae”’ being
more compact and parenchymatous and often more smooth and
shiny. Of the occurrence of these two forms together, three cases.
may be cited: In one collection (no. 6515) of 40 plants, all believed
referable to Galaxaura marginata, found growing together just below
low-water mark near Guantanamo Bay, Cuba, 26 have been examined
microscopically and of these 26, 13 were of the ‘‘ Brachycladia”’ struc-
194 BROOKLYN BOTANIC GARDEN MEMOIRS
ture and tetrasporic, 7 were of the ‘‘ Vepreculae”’ structure and cysto-
carpic, 5 were of the ‘‘Vepreculae’’ structure and antheridial, and 1
was of the “ Vepreculae”’ structure and apparently sterile. Of 9 plants
(no. 6966) found growing together at the mouth of Guanica Harbor,
Porto Rico, 6 were of the “ Vepreculae”’ structure, 2 of them being
obviously antheridial, and 3 were of the ‘‘ Brachycladia’’ structure,
2 of them obviously tetrasporic. Of 5 specimens (no. 7468) found
near low-water mark on Muertos Island, Porto Rico, 2 were of the
‘“Brachycladia”’ structure and tetrasporic, and 3 were of the ‘‘ Vepre-
culae’’ structure, I being cystocarpic, I antheridial, and 1 apparently
sterile. In some cases, a considerable series of specimens, all of one
group, has been collected, but in collecting the red algae it often
happens, as is well known, that the plants found at one time and place
may be either all tetrasporic or all sexual. Without waiting for the
results of cultural experiments which might furnish absolutely complete
proofs of the suggested genetic continuity, it seems to the writer that
the evidence is overwhelming that the so-called species of the Kjell-
man’s “‘ Vepreculae”’ section are simply the sexual phases of the species
of the “‘Brachycladia”’ section. It is of interest to note that Bérgesen,
in a recent instalment of his admirable series of papers on “‘ The Marine
Algae of the West Indies,’’* relying upon the sectional distinctions
proposed by Kjellman, appears to have described and figured the
antheridial plant (sect. ‘‘Vepreculae’’) of Galaxaura marginata (Ell.
& Sol.) Lamour. as a new species under the name Galaxaura occi-
dentalis Bérg., taking the tetrasporic plant (sect. ‘‘ Brachycladia’’)
to be the true G. marginata.4
When we come to examine the alleged species of some of the
other sections of the genus Galaxaura, as monographed by Kjell-
man, we find strong evidences of other correlations similar to those
already described for the Cameratae-Spissae and Brachycladia-Vepre-.
culae groups. In Kjellman’s section ‘‘Rhodura,” the peripheral
elements of the thallus are so manifestly and predominantly fila-
mentous (TEXT-FIGURE 3) that there is little ground for using the term
‘“cortex’’ in connection with these plants, yet there is commonly a
* 2: 109-113. f. II8—I23. 1916.
4The original of the Corallina marginata of Ellis and Solander (Nat. Hist. Zooph.
115. pl. 22. f. 6. 1786) was from the Bahama Islands, and, like most of the Ellis
and Solander types, it is not certainly known to be now in existence. However,
there is, in the herbarium of the Royal Botanic Gardens at Kew, an old fragment,
inscribed in the hand of Lamouroux:
Galaxaura marginata
Worallina. .~ teas oes Sol. et Ell.
Bahame
which may or may not represent an authentic bit from the Ellis collection. This is
antheridial and has the ‘‘ Vepreculae”’ structure.
HOWE: DIMORPHISM IN GALAXAURA 195
dimorphism in these peripheral assimilatory filaments, one set being
long and another short, and the short ones, more or less even-topped,
may sometimes be said to form a loose cortex. Whenever repro-
ductive organs are found on plants of the ‘“‘Rhodura’”’ section, they
are always tetrasporangia, never antheridia or cystocarps. In the
section ‘‘Microthoé,”’ one finds a firm, compact, pseudoparenchym-
atous cortex—usually firm and coherent, even after decalcification
(TEXT-FIGURE 4). In some of the species or forms belonging in this
section, the smooth firm epidermis bears, in certain parts of the thallus,
numerous long assimilatory filaments, and, when these are particularly
abundant, plants of the section ‘‘Microthoé’’ may look much like
those of the section ‘‘Rhodura,’’ but, generally speaking, it may be
said that the firm smooth cortex of plants of the ‘‘ Microthoé”’ section
and the rough shaggy exterior of plants of the “Rhodura”’ section
give them a very different appearance and it is no wonder that they
have been considered not only as different species but also as members
of different sections of the genus. But members of the ‘‘ Microthoé”’
section, except when apparently sterile, are always either antheridial
or cystocarpic—never tetrasporic—just as members of the ‘‘ Rhodura”’
section are always tetrasporic and never sexual. And plants of the
“Microthoé”’ section and those of the ‘‘Rhodura”’ section grow often
so closely associated—often intertangled in the same tuft—that it
seems to be a fair inference that they represent phases in the develop-
ment of one and the same species. Two plants from a collection (no.
1859) of about 100 specimens made by the writer near Santurce,
Porto Rico, in 1903, are shown on PLATE IV. Not all of the material -
in this collection has been examined microscopically, but, roughly
speaking, about 80 of the 100 have the ‘“Rhodura”’ structure, some
of them being obviously tetrasporic and others apparently sterile;
and about 20 of the 100 are of the ‘‘Microthoé”’ structure, some of
them being obviously antheridial or cystocarpic and others apparently
sterile. These plants of the ‘‘Rhodura”’ section appear to represent a
condition of what .Kjellman described as a new species under the
name Galaxaura flagelliformis, though usually less “ flagelliform”’ than
Kjellman’s original; the plants of the ‘‘Microthoé”’ section represent
what Kjellman described as a new species under the name Galaxaura
squalida. The two forms, as shown in FIGURES 2 and 3 (PLATE IV),
differ much in habit, yet, if we consider only the lower part of the
Galaxaura squalida, where the cortex is more or less covered with
free assimilative filaments, it looks a good deal like the shaggy tetra-
.sporic plant, G. flagelliformis. These Porto Rican specimens lying
under the no. 1859 were collected by the writer in his less experienced
and less critical days and were put together under one field number as
196 BROOKLYN BOTANIC. GARDEN MEMOIRS
representing a single species, as in all probability they do, even though
the current system of classification would require us to put them not
only in different species-covers, but also in different sections of the
genus. Likewise, in Bermuda, these two forms, Galaxaura flagelli-
formis and G. squalida, occur and in one instance, at least, they have
been placed together under one field number by F. S. Collins (8486
in herb. N. Y. Bot. Gard.).
In a similar way, Galaxaura subverticillata Kjellm., a tetrasporic
plant representing the section “‘Rhodura,” and G. rugosa (Ell. & Sol.)
Lamour., a sexual plant representing the section “‘Microthoé,’’ are,
in all probability, phases in the life-cycle of one and the same species.
As instances of their occurrence together may be mentioned the
writer’s no. 2042 (Santurce, Porto Rico), in which the two, the G.
subverticillata with young tetrasporangia and the G. rugosa with cysto-
carps, were found intertangled in the same tuft; the writer’s nos.
7470 (G. subverticillata) and 7469 (G. rugosa), growing close together
and sometimes intermingled, near the low-water line on Muertos
Island (Caja de Muertos), Porto Rico; the writer’s no. 4909a, G.
subverticillata, tetrasporic, occurring with or near no. 4911, G. rugosa,
cystocarpic, and other forms of Galaxaura at Montego Bay, Jamaica.
It must be confessed, however, that G. subverticillata occurs also with
sexual plants that agree more closely with G. squalida than with G.
rugosa and that just as the lines of distinction between G. flagelliformis
and G. subverticillata often seem vague and uncertain, so also do G.
squalida and G. rugosa appear to intergrade.
The plants included by Kjellman in his section ‘Eugalaxaura”’
appear to be all sexual, never tetrasporic. The cortex is here smooth
and firm, much as in the section “ Microthoé,’’ but the epidermal cells
are commonly smaller, the cortex dissolves into its constituent fila-
ments more readily on decalcification, the thallus is more distinctly
jointed, and free superficial assimilatory filaments are of less frequent
occurrence. The tetrasporic phases ofthe ‘Eugalaxaura’’ forms are
apparently to be found in the section “‘Rhodura,” this section supply-
ing the tetrasporic conditions for both the section ‘‘Microthoé”’ and
the section ‘‘Eugalaxaura.’’ From size and association (at Santurce,
Porto Rico, and elsewhere) more than from any similarity in habit (for
the two are, as a rule, strikingly different in habit), the writer believes
that Galaxaura cylindrica (Ell. & Sol.) Lamour. of the section ‘‘ Euga-
laxaura”’ finds its tetrasporic phase in G. lapidescens (Ell. & Sol.)
Lamour., of the section ‘‘Rhodura,” as this species has been recently
limited and defined by B¢grgesen.® And, with less assurance, it may
5 Mar. Alg. Dan. W. I. 2: 95-99. f. 102-104. 1916.
BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME I, PLATE III.
Howe: GALAXAURA MARGINATA (ELL. & SOLAND.) LAMouR. (ANTHERIDIAL, SECTION
““VEPRECULAE.’’)
BROOKLYN BOTANIC GARDEN Memoirs VoLuME |, PLATE IV.
HOWE: 1. GALAXAURA MARGINATA (ELL. & SOLAND.) Lamour. (TETRASPORIC, SECTION
“ BRACHYCLADIA.’’)
2. GALAXAURA FLAGELLIFORMIS KJELLM. (SECTION ‘‘ RHODURA.’’)
3. GALAXAURA SQUALIDA KJELLM. (SECTION ‘‘ MICROTHO#.’’)
HOWE: DIMORPHISM IN GALAXAURA 197
be surmised that Galaxaura oblongata (Ell. & Sol.) Lamour.® has its
tetrasporic condition in G. comans Kjellm. And just as the line of
demarkation between Galaxaura oblongata and G. cylindrica seems a
little uncertain and arbitrary, so also is the line of separation between
G. comans and G. lapidescens. Where there is so much difference in
habit as there is between the “Rhodura’”’ forms on the one hand and
the ‘‘Microthoé”’ and ‘‘Eugalaxaura’’ forms on the other, there is
manifestly more need for a cultural demonstration of their correlation
as alternating generations than there is in the case of the Cameratae-
Spissae and Brachycladia-Vepreculae groups, where the two
phases have the same outward appearance. But while experimental
demonstration or further observations in the field may be desirable
for a precise correlation of the “‘ Rhodura’”’ forms, the existing evidence
that these ‘‘Rhodura”’ forms represent tetrasporic phases of “ Micro-
thoé”’ and ‘“‘Eugalaxaura’”’ forms seems convincing.
EXPLANATION OF PLATES III AND IV
PLATE III
Photograph of a formalin-preserved antheridial specimen of Galaxaura marginata
(Ell. & Soland.) Lamour., representing Kjellman’s section ‘‘ Vepreculae.’’* Speci-
men from San Juan, Porto Rico (Howe 2304); natural size.
PLATE IV
Fic. 1. Photograph of a formalin-preserved tetrasporic specimen of Galaxaura
marginata (Ell. & Soland.) Lamour., representing the section “ Brachycladia.”’
Specimen from San Juan, Porto Rico (Howe 2304); natural size.
Fic. 2. Photograph of a formalin-preserved specimen representing a form of
Galaxaura flagelliformis Kjellm. and belonging in Kjellman’s section “ Rhodura.”’
Specimen from Santurce, Porto Rico (Howe 1859a); natural size.
Fic. 3. Photograph of a formalin-preserved specimen of Galaxaura squalida
Kjellm. Specimen from Santurce, Porto Rico (Howe 1859b); natural size.
6 Galaxaura fragilis of Kjellman and of Boérgesen; not Dichotomaria fragilis
Lamarck, the type specimen of which in herb. Mus. Paris. appears to have the
structure of the ‘‘Spissae”’ group.
THE UREDINALES OF OREGON!
H. S, JACKSON
Purdue University
Since no account of the rusts of any of the states bordering on the
Pacific coast is available for reference by the students of the flora of
that region, it has seemed desirable to bring together in the form of
an annotated list the results of a study of the species occurring in
Oregon, which has extended, intermittently, over a period of eight
years.
The account is the result of a study begun by the author in 1909
at the Oregon Agricultural College and finally prepared in the form
here presented at the Purdue University Agricultural Experiment
Station. ,
On account of the great diversity of conditions the State of Oregon
presents a wonderful field for work in any phase of botanical study.
The area of the State is approximately 96,000 square miles, an area
considerably greater than that included in all the New England states
combined. The great range of climatic and topographical features
existing in the State offers favorable conditions for the development
of a flora not only large in number of species, but very diverse in char-
acter. Within the confines of the State is to be found a range in alti-
tude from sea level to perpetually snow-capped mountains. The
annual rainfall varies from over 80 inches in some localities to below
10 inches in others, resulting in the development of a flora:almost
tropical in its luxuriance on the one hand, and one having many of
the characteristics of a desert on the other.
The diversity of the Phanerogamic flora which has developed
under these conditions offers an especially attractive field for the
student of the parasitic fungi. Many species of all groups are to be
found especially in those portions of the State having a heavy annual
rainfall.
Rusts are found in great profusion in all sections of the State.
The first collections of this group made by the writer were accumu-
lated in connection with an effort to obtain general material for class
use. The greater part of the collections have been made in connection
with local excursions, the primary object of which was recreation.
‘Contribution from the Botanical Department of Purdue University Agri-
cultural Experiment Station.
198
JACKSON: UREDINALES OF OREGON 199
Many of the collections made by the writer at localities outside of
Benton County have been picked up in spare moments on trips taken
in connection with Experiment Station or Extension Service duties.
During 1914 and 1915, however, a number of special excursions were
made primarily for collecting this group of fungi.
In addition to those made by the writer, several hundred collections
made by his former associates, assistants and students at the Oregon
Agricultural College are included. The greater number of these were
collected by Prof. H. P. Barss, Mr. F. D. Bailey and Mr. G. B. Posey.
To these have been added a considerable number of records obtained
from miscellaneous sources. Several of these were obtained from the
herbarium of the New York Botanical Garden, and of the National
Museum. A considerable number are in the Arthur Herbarium at
the Purdue University Agricultural Experiment Station. The greater
number of these were obtained originally from phanerogamic speci-
mens mainly collected and distributed by pioneer botanists of the
region, particularly W. C. Cusick, Thomas Howell, J. B. Lieberg and
E. P. Sheldon, E. R. Lake and others.
A few collections were made in Oregon by Dr. David Griffiths and
associates, most of which were distributed in his “‘ West American
Fungi.’”’ Mr. E. Bartholomew collected at a few localities in Oregon
in 1915 and distributed the specimens in the exsiccati, ‘Fungi Colum-
biani’’ and ‘‘ North American Uredinales,’’ which he edits. A number
of specimens of rusts, the records of which were obtained mainly
from the Arthur Herbarium, were made by Moses Craig, at one time
botanist at the Oregon Agricultural College. It is evident that he
made quite an extensive collection of rusts in Oregon, but the location
of his collection at the present time is unknown to the writer.
Oner of the most interesting collections which it has been the
privilege of the writer to examine was made by Dr. J. R. Weir, mainly
in the northeastern and southwestern parts of the State. This col-
lection consists of about 130 numbers and was sent to this laboratory
for study in 1915 and 1916. Another interesting collection of about
30 numbers was made by Dr. E. P. Meinecke in southwestern Oregon
and forwarded to the writer for study.
Approximately thirteen hundred collections have been examined in
the preparation of this account and are listed in the following pages.
By far the greater number of these, about one thousand, were made in
western Oregon, including the Cascade Mountains. Of this number
about six hundred were made in the Willamette Valley, four hundred
having been collected in Benton County, mostly in the vicinity of
Corvallis. Two hundred and fifty are listed from the Cascade moun-
tain region, most of the collections having been made in the vicinity
200 BROOKLYN BOTANIC GARDEN MEMOIRS
of Mt. Jefferson or in Hood River County. About sixty collections
are recorded from the counties bordering on the coast, only a few of
which are from the southern coast counties. Less than three hundred
collections have been made in eastern Oregon, almost half of which
are from the mountainous region of the northeastern part.
It will be seen from the above summary of the distribution of the
collections recorded that only a very small portion of the State has
been explored for this group of fungi. Much remains to be done,
particularly in southwestern Oregon and in eastern Oregon. The
mountains of the Coast Range have been explored only in Benton
County, where several collections have been made on Mary’s Peak.
The coast counties also offer a rich field for the collector. The region
around Corvallis in Benton County is the only portion of the State that
can be said to have been carefully explored for rusts. Corvallis is
situated at the junction of the Willamette River and Mary’s River
and lies in the edge of the foothills of the Coast Range.
In spite of the fact that the exploration so far conducted is in-
adequate to furnish a very accurate idea of the rust flora of the State,
records for 220 species are brought together in the account which
follows. These occur on about 500 different host plants. In connec-
tion with the study of the collections 10 species have been found
which are believed to be new to science, 8 of which are described in
the following pages. A number of European species not previously
recorded for North America have also been collected in the State.
A large number of unrecorded host plants have been encountered.
The number of speciés of economic importance found in the State
is worthy of mention. All of the grain rusts recorded for North
America with the exception of the corn rust (Puccinia Sorghi) are
known to occur in the State, including the recently discovered Puccinia
glumarum. All of the rusts troublesome to florist’s crops in the
greenhouse, including Puccinia Antirrhini, are known to occur. The
Pacific coast rust of pears and quinces promises to become of con-
siderable economic importance. The large number of forest-tree
rusts found in the State offers an especially attractive field for investi-
gation. Many heteroecious species are known to occur whose life
history is not yet determined. The rusts occurring on Salix are
especially in need of investigation.
In the notes given in connection with the account of the species
which follows, an effort has been made to summarize the available
information on the life history as shown by any culture work which
may have been conducted either by American or European authors.
Notes on the distribution in North America are given whenever it
was considered of sufficient importance. The genera are listed alpha-
JACKSON: UREDINALES OF OREGON 201
betically under each family and the species similarly under the genus.
The host plants are also arranged alphabetically under each species.
No attempt has been made to give a full list of synonyms. In
general, however, sufficient synonymy is given to show the origin of
the specific name used as well as any names which have been in general
use. For convenience of reference the specific nomenclature used is
that in use in this laboratory, and in general conforms to that used in
the ‘“‘North American Flora.’’ For similar reasons the generic nomen-
clature follows that proposed by Dr. Arthur, for use in the “ North
American Flora,’”’ except that Melampsora, Puccinia and Uromyces,
are retained. In order to follow this system consistently it has been
found necessary to establish a number of new combinations.
Under each host is given a list of the specimens examined, with
locality, county, date and collector, followed by the collector’s number.
Numbers between 1000 and 3450 refer to collections in the Oregon
Agricultural College Herbarium, if made by members of the botanical
staff or by students. If no collector is given it may generally be
assumed that the collection was made by the writer. In order to
prevent unnecessary repetition, Benton County is not cited following
collections made at Corvallis and Philomath. Similarly Hood River
County is implied for all collections from Hood River or Mt. Hood
and all collections made at Portland are from Multnomah County.
The writer is under great obligations to all those who have contrib-
uted specimens for study and especially to those whose names have
been mentioned previously. He is also greatly indebted to those
botanists who have given so generously of their time in determining
host plants. Dr. A. S. Hitchcock and Mrs. Agnes Chase have named
most of the grasses. Dr. Theo. Holm and Dr. K. K. Mackenzie
have each determined a considerable number of species of Carex.
Dr. C. R. Ball has named most of the willows. Dr. F. V. Coville and
Dr. K. M. Wiegand have each determined several specimens of Juncus.
Mr. Paul Standley has determined a large number of specimens from
miscellaneous families.
Dr. J. C. Arthur and his former associates very kindly determined
a considerable number of specimens of the rusts occurring on grasses
and sedges, and verified the determinations of others, which the writer
sent from Oregon at various times during the period when the col-
lections were being made. The greater part of the collections recorded,
however, have been worked over since the writer took charge of the
work of this laboratory. During this period it has been his privilege
to be able to consult freely with Dr. Arthur and to have the unre-
stricted use of the collections, catalogues, and manuscript notes on
the rusts, which have been accumulated at the Purdue University
02 BROOKLYN BOTANIC GARDEN MEMOIRS
Le)
Agricultural Experiment Station during the many fruitful years of
Dr. Arthur’s administration of the department of botany. Without
this assistance the preparation of this account in the form presented
would not have been possible and the writer takes great pleasure in
acknowledging his indebtedness to Dr. Arthur and to the various
assistants in this laboratory for any help which they may have given.
COLEOSPORIACEAE
1. Coleosporium Adenocaulonis sp. nov.
O.and I. Pycnia and aecia unknown.
iI. Uredinia hypophyllous, few, scattered on conspicuous angular
yellowish spots, small, 0.I-o.2 mm. across, early naked, orange yellow
fading to whitish, becoming pulverulent, ruptured epidermis con-
spicuous; urediniospores globoid to ellipsoid, 18-24 by 23-26 yp, wall
light golden brown or colorless, 2-3 » thick, prominently and moder-
ately verrucose; pores indistinct.
III. Telia unknown.
ON CARDUACEAE:
Adenocaulon bicolor Hook.—Corvallis, Sept. 20, 1914, 1549.
This species is very inconspicuous, developing very small sori on
the under side of the leaves on yellowish spots.
2. COLEOSPORIUM MADIAE Cooke, Grevillea 7: 102. 1879.
On CarRDUACEAE: II, III.
Madta citriodora Greene—Mary’s Peak, Benton Co., Aug. 15, 1914,
1514.
Madia exigua (Sm.) Greene—Corvallis, July 29, 1914, 1475;
Philomath, Aug. 15, 1914, 15106.
Madia glomerata Hook.—Corvallis, Aug., 1889, E. R. Lake, July,
1910,:2759, July 209, 1915, 3247; Portland, Aug. 24, 1915, E. Bar
tholomew, 5964 (Barth. Fungi Columb. 4970).
Madia racemosa (Nutt.) T. & G.—Corvallis, July, 1910, rz60,
Sept. 12, 1910, 1928; Wren, Benton Co., June 26, 1914, 1316, 1317,
1322, 1328; Elk City, Lincoln Co., Aug. 20, 1914, 2538; Philomath,
May I0, 1914, 3246.
Madia ramosa Piper—Corvallis, July 29, 1914, 1470.
Madia sativa Molina—Corvallis, Aug. 12, I910, 1163, July 29,
1914, 1474.
The aecial connection of this very common species has not been
demonstrated by cultures and no field observations have been made
in Oregon. Judging from distributional data, however, it seems
probable that Peridermium californicum Arth. & Kern may be gen-
etically connected.
JACKSON: UREDINALES OF OREGON 203
From field observations made by the writer it is evident that in
western Oregon this species overwinters in the uredinial stage.
3. COLEOSPORIUM OCCIDENTALE Arth. North American Flora 7: 94.
1907.
On CarbDuaceseE: II.
Senecio triangularis Hook.—Mary’s Peak, Benton Co., Aug. 15,
1914, 1518.
This species is known otherwise only from the type collection
made in Falcon Valley, Washington, on S. hydrophiloides Rydb., by
W. N. Suksdorf in 1900.
The aecial connection is not known and no clues are available.
The aecia, in common with other species of Coleosporium whose life
history is known, should be looked for on the leaves of Pinus sp.
The above collection, however, was made in a region where no pines
exist in a radius of several miles. It is probable that this species, in
common with some other members of the genus, is capable of being
carried over the winter in the uredinial stage.
The only pine-leaf-inhabiting Peridermium known to the writer
in the present range of this species is P. montanum Arth. & Kern,
which has been shown to be genetically connected with a Coleosporium
on Aster and Solidago, referred to C. Solidaginis.
4. COLEOSPORIUM SOLIDAGINIS (Schw.) Thiim. Bull. Torrey Club 6:
eee -iS78.
Uredo Solidaginis Schw. Schr. Nat. Ges. Leipzig 1: 70. 1822.
Peridermium acicolum Und. & Earle, Bull. Torrey Club 23: 400.
1896.
Peridermium montanum Arth. & Kern, Bull. Torrey Club 33: 413.
1906.
On PinaceseE: I.
Pinus contorta Dougl.—North slope Mt. Hood, Aug. 7, 1914, 16z0.
On CarpDuAcEAE: II, III.
Aster conspicuus Lind|l.—Hilgard, Union Co., July 10, 1914, 1532;
Austin, Grant Co., Aug. 1915, J. R. Weir, 1509.
Aster Cusecku Gray ?—Corvallis, Sept. 21, 1914, 1548.
Aster Douglas Lindl.—Hood River, Aug. 26, 1915, E. Bartholo-
mew, 5972 (Barth. Fungi Columb. 4977); Corvallis, June 29, 1914,
G. B. Posey, 1370.
Aster foliaceus frondeus Gray—Hood River, July 22, 1915, 3137;
Clatskanie, Columbia Co., May 20, 1914, F. D. Bailey, 2564, Oct.
29, 1914, 2531; Corvallis, June 29, 1914, G. H. Godfrey, 1307.
Aster Hallit Gray—Corvallis, July 29, 1914, 1471; Wren, Benton
Co. July 26, 10L4, 737.
204 BROOKLYN BOTANIC GARDEN MEMOIRS
Solidago caurina Piper—North slope Mt. Hood, Aug. 7, 1914,
1605.
Solidago elongata Nutt.—Corvallis, July 29, 1915, 3244; Scotts,
7 miles N. of Fort Klamath, Klamath Co., Sept. 20, 1913, E. P.
Meinecke, Cr D 7.
Solidago missouriensis Gray?—Sumpter, Baker Co., Aug. 21, 1915,
J. R. Weir, 267.
Solidago tolmieana Gray ?—Hood River, July 23, 1915, 3254.
The life history of this species was first demonstrated by Clinton
(Science N.S: 25: 289. 1907; Ann. Rep. Conn. Exp. Sta. 29@G:
320. 1907; 1907: 375. 1908). He successfully infected Solidago
rugosa with aeciospores of Peridermium acicolum on Pinus rigida.
The single collection of aecia listed above (1610) agrees with
the description of P. montanum Arth. & Kern and was collected in the
immediate vicinity of Solidago caurina (1605). The possibility of
genetic relationship was made note of at that time. Hedgcock
(Mycologia 4: 144. 1912; Phytopath. 3: 16. 1913) has also made
similar observations and more recently (Phytopath. 6: 65. - 1916)
has cultured this Peridermium successfully on Aster conspicuus, using
aecial material on Pinus contorta collected in Montana. Weir and
Hubert (Phytopath. 6:68. 1916) working independently from Hedg-
cock, with similar aecial material, have also demonstrated by cultures
that this Peridermium has its uredinia on both Aster and Solidago,
having obtained infection on A. laevis geyeri, S. canadensis and S.
M1SSOULIENSIS.
Sydow (Monographia Ured. 3: 621. 1915) suggests that the form
on Aster in North America is different from C. Solidaginis on Solidago
and should either be united with the Asiatic C. Asterum (Diet.) Syd.
or that it represents an unrecognized species having a different Pert-
dermium as its aecial form. The culture work of Weir and Hubert
(1. c.), however, shows that P. montanum is genetically connected with
uredinia on both Aster and Solidago and does not lend support to
Sydow’s view.
While the two species of Peridermium included here are widely
separated as to range and are morphologically distinguishable, it
seems best until further culture work is conducted to recognize but
one American species.
UREDINACEAE
5. CALYPTOSPORA COLUMNARIS (Alb. & Schw.) Kiihn; Rab.-Wint.
Fungi Eur. 3521. 1886. (Hedwigia 26: 28. 1887.)
Aecidium columnare Alb. & Schw. Consp. Fung. 121. 1805.
Calyptospora Geoppertiana Kiihn, Hedwigia 8: 81. 1869.
JACKSON: UREDINALES OF OREGON 205
ON PINACEAE: I.
Abies grandis Lindl.—Scottsburg, Lane Co., Sept. 1, 1914, G. G.
Hedgcock, 20210.
Abies magnifica A. Murr.—Road to Crater Lake, Union Creek,
Camp Grant, Klamath Co., Sept. 23, 1913, E. P. Meinecke, Cr D 20.
On VaAccINIACEAE: III.
Vaccinium macrophyllum (Hook.) Piper—Austin, Grant Co., June,
1913, J. R. Weir, 25; Sumpter, Baker Co., June, 1913, J. R. Weir, 24;
Silver Creek, Josephine Co., July 28, 1913, E. P. Meinecke, Sz (D6) Dr.
Vaccinium myrtilloides S. Wats.—Road to Crater Lake, Union
Creek to Camp Grant, Klamath Co., Sept. 23, 1913, E. P. Meinecke,
er D ro.
Vaccinium ovalifolium Smith—Larch Mt., Multnomah Co., Aug.
1910, 1756; North slope Mt. Hood, Aug. 7, 1914, 1608.
Vaccinium ovatum Pursh—Dothan, Douglass Co., Sept. 8, 1914,
G. B. Posey, 1932; Waldo, Josephine Co., Sept. 5, 1916, J. R. Weir,
280; Oregon, April 19-31, 1911, H. D. House.
Vaccinium parviflorum Smith—Whitewater Ranger Station, near
Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 1750.
Vaccinium scoparium Lieb.?—Mary’s Peak, Benton Co., Aug. 15,
1914, 1284. ,
Specimens of aecia collected in various parts of North America on
Abies balsamea, A. concolor and A. lasiocarpa are now referred to this
species in the Arthur herbarium.
The life history was first demonstrated by Hartig (Allg. Forst.- u.
Jagdzeitg. 289. 1880), who conducted culture investigations using
aecia on Abies pectinata and telia on Vaccinium Vitis-idaea. He
obtained successful infection in both directions. Other European
investigators, notably Dr. G. Winter, have amply confirmed these
results. (Klebahn, Die Wirtsw. Rostpilze 391. 1904.)
In America, Arthur (Mycologia 2: 231. 1910) was the first to
culture this species and succeeded in obtaining aecia on Abies Fraseri
following exposure to infection from telia on Vaccinium pennsyl-
vanicum sent by W. P. Fraser from Nova Scotia. Later in the same
year Fraser made the first field collection of aecia on Abies balsamea
(Science 30: 814. 1909) and later (Mycol. 4: 177. 1912; 6: 27.
1914) confirmed Arthur’s work by obtaining infection on Abies bal-
samea from telia on Vaccinium pennsylvanicum.
6. CurysoMyxA WEIRII Jackson, Phytopath. 7: 353. 1917.
On PINACEAE:
~ Picea Engelmanii Parry—Whitman Nat. Forest, Oregon, July 17,
1913, J. R. Weir, 2717.
206 BROOKLYN BOTANIC GARDEN MEMOIRS
This species differs from C. Abietis in the narrower, somewhat
smaller spores which do not long remain in chains but soon break
apart. No evidence of germination has been seen in any of the col-
lections. This is the only American representative of the genus as
restricted by Arthur. (Result Sci. Congr. Bot. Vienne 338. 1906.)
It is known to the writer otherwise only from single collections from
British Columbia and Idaho. It is doubtless not uncommon in the
northwest.
7. CRONARTIUM FILAMENTOSUM (PK.) Hedgcock, Phytopath. 2: 177.
EO12.
Peridermium filamentosum Pk. Bot. Gaz. 7: 56. 1882.
Uredo coleosporioides Dietel & Holway, Erythea 1: 247. 1893.
Peridermium stalactiforme Arth. & Kern, Bull. Torrey Club 33:
419. 1906.
Cronartium coleosporioides Arth. N. Am. Flora 7: 123. 1907.
On PINACEAE: I.
Pinus contorta Doug|.—Scotts, Anna Creek, Klamath Co., May 23,
1912, E. P. Meinecke, used for inoculation on Castilleja miniata; Gold
Center, June 20, 1914, H. F. Wilson, 1856; North slope Mt. Hood,
elev. 3,000-4,000 ft.; Aug. 7, 1914, 3332; Sumpter, Baker Co.,
May, 1916, J. R. Weir.
On SCROPHULARIACEAE: II, III.
Castilleja sp.—North slope Mt. Hood, 3,000-4,000 ft., Aug. 7,
1914, 2612, 1615 (collected near 3332); Ashland Toll House, Jackson
Co., Sept. 27, 1913, E. P. Meinecke, Cr D 22.
Hedgcock (I. c.) was the first to publish a record of connection
of Peridermium filamentosum with a Cronartium on Castilleja by cul-
tures. He considered this distinct, however, from Cronartium coleo-
sporioides, which Meinecke had cultured in 1911 (Phytopath. 3: 167—
168. 1913) and shown to have for its aecial form P. stalactiforme.
Meinecke’s culture material was collected in Klamath Co., Oregon.
Further culture work has been carried on by Weir and Hubert
(Jour. Agr. Research 5: 781-785. 1916) in which it is shown that
the gall-forming Peridermium on Pinus contorta which has previously
been commonly referred to P. Harknessti Moore is but a form of P.
filamentosum.
All the records of the aecial stage given above are of the gall-
forming type. The Hood River material was collected in the im-
mediate vicinity of the telial form on Castilleja.
8. CRONARTIUM PYRIFORME (Pk.) Hedgc. & Long, Alt. Stage Peri-
dermium pyriforme 3, 1914.
Peridermium pyriforme Peck, Bull. Torrey Club 6: 13. 1875.
JACKSON: UREDINALES OF OREGON 207
Cronartitum Comandrae Peck, Bull. Torrey Club 11: 50. 1884.
Peridermium Betheli Hedgc. & Long, Phytopath. 3: 251. 1913.
ON PINACEAE: I.
Pinus ponderosa Dougl.—Hood River Co., May I0, 1910, 3333;
Sumpter, Baker Co., May, 1916, J. R. Weir.
On SANTALACEAE: II, III.
Comandra umbellata (L.) Nutt.—Corvallis, June 20, 1909, E. R.
Lake, 3068, July 24, 1914, 2570, Road to Ashland toll house, Jackson
Co., Sept. 27, 1913, E. P. Meinecke, Cr D 23; Hood River Co., June
20, 1914, 1995, July 22, 1915, 3143; Dufur, Wasco Co., June 30, 1914,
1337; Indian Creek, Malheur Co., Sept. 16, 1897, E. P. Sheldon, 8934.
The collection of aecia on Pinus ponderosa made at Hood River
consisted of a large fusiform gall at the base of the trunk of a young
tree about 2 inches in diameter. The gall entirely encircled the tree
which was noticeably stunted from the effects of the parasite. The
foliage also showed a distinct yellow cast.
The life history of this common and widespread species was first
demonstrated by Hedgcock and Long (Il. c.). They succeeded in
obtaining the development of uredinia on Comandra umbellata by
exposing them to infection from aecia on Pinus ponderosa collected in
Washington and California and on Pinus pungens from Pennsylvania.
In a later publication the authors (Bull. U. S. Dept. Agr. 247: 1-20.
1915) discuss the economic importance of this fungus as a disease of
pines and record in detail the results of extensive culture work.
g. HyALopsora Aspiptotus (Peck) Magn. Ber. Deuts. Bot. Ges. 19:
Bo2. TOOT: ;
Uredo Aspidiotus Pk. Ann. Rept. N. Y. State Mus. 24: 88. 1872.
ON POLYPODIACEAE:
Phegopteris Dryopteris (L.) Fée—Austin, Grant Co., Aug. I915,
J. R. Weir, 764.
10. HYALOPSORA LAEVIUSCULA (D. & H.) Arth. North Am. Flora 7:
LIZ. 1907.
Uredo laeviuscula Dietel & Holway, Erythea 2: 127. 18094.
ON POLYPODIACEAE:
Polypodium occidentale (Hook.) Maxon—Vicinity of Mt. Jefferson,
July 27, 1907, E. R. Lake, 2508; Corvallis, March 25, 1915, G. B.
Posey, 2626; Hood River Co., May 16, 1915, 3042; Bridal Veil,
Multnomah Co., May 18, 1915, 3025.
Polystichum munitum (Kaulf.) Presl., Mary’s Peak, Benton Co.,
Apr. 23, 1915, G. B. Posey, 3041.
The urediniospores in this species are smooth in all collections as
shown by very careful examination with the oil immersion objective.
15
208 BROOKLYN BOTANIC GARDEN MEMOIRS
This species has not previously been recorded on the latter host
so far as the writer is aware.
11. HyALopsora PoLypopit (DC.) Magn. Ber. Deuts. Bot. Ges. 19:
Boz. .190L.
Uredo Polypodiu DC. Fl. Fr. 6: 81. 1815.
ON POLYPODIACEAE:
Filix fragilis (L.) Underw.—Road to Lost Lake, Hood River Co.,
July 24, 1915, 3024.
12. MELAMPSORA sp.
II. Uredinia amphigenous, chiefly epiphyllous, scattered or occa-
sionally gregarious, round, 0.5-I1 mm., early naked, somewhat
pulverulent, orange fading to yellowish, ruptured epidermis not con-
spicuous; uredospores ellipsoid or obovoid, 15-19 by 21-24 yu, wall
colorless, uniformly 2.5—3 uw in thickness, moderately to closely verru-
cose-echinulate; paraphyses numerous, chiefly peripheral, clavate or
occasionally capitate, 18-26 by 45-70 u, wall colorless, usually uni-
formly 1-2 pu thick, occasionally thickened at apex to 4 um.
ON SALICACEAE:
Populus alba V..—Sheridan, Yamhill Co., July 7, 19014; ie
Barss, 1935; Cottage Grove, Lane Co., July 17, 1914, 1933; Philo-
math, July 20, 1915, 3309.
The only other American collection on this host known to the
writer is one in the Arthur herbarium, collected by E. Bethel, Aug. 7,
1913, at San Jose, Cal. These specimens differ from all other North
American collections on Populus. It seems most probable that this
is an introduced European species. Only uredinia are present in
American collections and it is quite impossible to assign it to any
known species without telial material. A description of the uredinial
stage drawn up from the Oregon collections is given for the benefit
of those who may have occasion to study this form.
The Oregon collections were all made from low, rapidly growing
water sprouts.
13. MELAMPSORA ALBERTENSIS Arth. Bull. Torrey Club 33: 517.
1906.
Caeoma occidentalis Arth. Bull. Torrey Club 34: 591. 1907.
On PINACEAE: I.
Pseudotsuga mucronata (Raf.) Suds.—Southeast Mt. Jefferson,
Linn Co., July 3, 1914, F. D. Bailey, 1841; Sumpter, Baker Co.,
July 20, 1913, J. R. Weir, 275; Corvallis, June 1910.
The life history of this species has been studied by Arthur (Myco-
logia 4: 29 and 59, 1912), who obtained infection resulting in pycnia
JACKSON: UREDINALES OF OREGON 209
and aecia on Pseudotsuga by exposing the foliage to infection from
germinating telia on Populus tremuloides collected in Colorado. Out
of four trials, three were successful. No infection was obtained on
Larix.
It is noteworthy in this connection that all of the northwestern
collections have larger spores than specimens from Colorado. The
former show spores 20-28 by 24-32 u while the average of the latter
are 16-20 by 19-26 uw. The culture work was conducted with Colorado
material, nearly if not all of which was collected in immediate associa-
tion with M. albertensis on P. tremuloides. The type of Caeoma
occidentale, on the other hand, was collected in British Columbia and
has larger spores. It seems entirely possible that the northwestern
collections may represent a different species and have genetic relation-
ship with some form on Populus other than M. albertensis.
14. MELAMPSORA ARCTICA Rostr. Medd. Grénland 3: 535. 1888.
ON SALICACEAE:
Salix Bebbiana Sarg.—Sumpter, Baker Co., Aug. 1915, J. R.
Weir, 167. >
Salix fendleriana And.—Sumpter, Baker Co., June, 1913, J. R.
Weir, 8.
Salix lutea Nutt.—Sumpter, Baker Co., June, 1913, J. R. Weir, 4.
Salix sitchensis Sanson—Dothan, Douglass Co., Sept. 8, 1914, G.
B. Posey, 3342.
Salix sp.—Scott’s, 7 miles from Fort Klamath, Klamath Co.,
Sept. 20, 1913, E. P. Meinecke, Cr D 6.
It is with considerable hesitation that the above collections have
been referred to this species. Only those collections which have small,
rather thin-walled uredospores, accompanied by an abundance of
thin-walled, clavate paraphyses, are included.
Fraser, working with material collected in Nova Scotia (Mycol.
4: 187. 1912; 5: 238. 1913), has made a cultural study of this
species. He succeeded in obtaining infection on Abies balsamea with
production of pycnia and aecia following exposure to germinating
telia from Salix discolor.
15. MeLAMPsoRA BIGELOW! Thiim. Mitth. Forstl. Vers. Oest. 2: 37.
1879.
On PINACEAE: I.
Larix occidentalis Nutt.—Hood River Co., elevation 4000°, July
23, 1915, 3305, 3305.
On SavicacEaE: II, III.
Salix Bebbiana Sarg.—Austin, Grant Co., Aug. 1915, J. R. Weir,
162, 163; Sumpter, Baker Co., July 19, 1913, J. R. Weir, 272.
210 BROOKLYN BOTANIC GARDEN MEMOIRS
Salix cordata Muhl.—Sumpter, Baker Co., Aug. 1915, J. R. Weir,
167.
Salix Piperi Bebb.—Philomath, Oct. 29, 1911, 3346; The Dalles,
Wasco Co., Aug. 26, 1915, E. Bartholomew (Barth. Fungi Columb.
4730).
Salix pseudocordata Anders.—Hilgard, Union Co., July 10, 1914,
1530.
Salix scouleriana Barr.—Corvallis, Sept. 19, 1910, 1165; St.
Johns, Multnomah Co., June 23, 1915, W. E. Lawrence, 3347; Austin,
Grant Co:, Aug: 1915, J. R. Weir, 765; Portland, Aug. 24, 19n5:ae@
Bartholomew (Barth., N. Am. Ured. 1417).
Salix sp.—Calamity, Aug. 1901, Griffiths & Morris (Griffiths, W.
Amer. Fungi 341); Crater Lake, Klamath Co., Sept. 22, 1913, E. P.
Meinecke, Cr Pk D (2) 13; Hood River Co., May 14, 1914, 1500,
Aug. 5, 1914, 1483, 1484; Beaverton, Washington Co., July 15, 1914,
F. D. Bailey, 1507; Austin, Grant Co., Aug. 25, 1915, J. R. Weir, 262.
The above specimens are tentatively assigned to this species.
There are quite certainly not less than four species of Melampsora on
Salix in North America. The characteristics by which they may be
separated in the uredinial stages are not well worked out at the present
time. The larger spored forms have been included here under M.
Bigelow.
Arthur (Jour. Myc. 11: 60. 1905) first established the fact that
this rust has its aecia on Larix. He succeeded in infecting Larix
decidua in two trials, by inoculating with basidiospores from germi-
nating telia on Salix amygdaloides collected in Wisconsin. This
result was later confirmed (Jour. Myc. 13: 194. 1907) with telial
material collected in Indiana.
Weir and Hubert (Phytopath. 6: 372. 1916) have succeeded in
obtaining infection of this species from Salix bebbiana Sarg. collected
in Montana on Larix occidentalis, and from S. cordata mackenzieana
collected in Idaho on Larix europea. The same authors (Phytopath.
7: 109. 1917) have recently repeated the work with the last-named
species of Salix and obtained infection with development of pycnia
and aecia on both L. occidentalis and L. europea.
16. Melampsora confluens (Pers.) comb. nov.
Uredo confluens Pers. Obs. Myc. 1: 98. 1796.
On GROSSULARIACEAE: I.
Ribes lacustre (Pers.) Poir—Philomath, May 3, 1913, F. D.
Bailey, 1107.
On SALICACEAE: II, III.
Salix argophylla Nutt.—Freewater, Umatilla Co., June 17, 1913,
F. D. Bailey, 7764, Aug. 12, 1915, F. D. Bailey, 3344.
JACKSON: UREDINALES OF OREGON 211
Salix scouleriana Barr.—Cascade Locks, Hood River Co., Aug.
Rie toro, 7775, Myrtle Creek, Douglass-Co.,.June 9, 1914, F: D:
Bailey, 3345; Hood River Co., July 23, 1915, 3343, 3366; Ashland,
Jackson Co., Sept. 10, 1914, 3340, 3341.
Salix sp.—Scott’s, 7 miles north of Fort Klamath, Klamath Co.,
Sept. 20, 1913, E. P. Meinecke, Cr D 4, Cr D 9; Austin, Grant Co.,
Aug. 25, 1915, J. R. Weir, 263; Grant’s Pass, Josephine Co., Sept. 3,
1916, J. R. Weir, 266; White Pine, Baker Co., July 20, 1913, J. R.
Weir, 270; Unity, Baker Co., Aug. 1915, J. R. Weir, 278.
It is impossible to assign with any degree of certainty the collec-
tions which should be referred to this combination. Only those col-
lections having small, rather thick-walled spores, accompanied by an
abundance of capitate, thick-walled paraphyses, are included. It is
possible that some of the collections included under M. Bigelowii
should be referred here.
No culture work has been conducted in America. A summary of
European work has been made by Klebahn (Die Wirtsw. Rostpilze
424. 1904).
In addition to the above, aecia have been collected in America
on Ribes saxosum from Utah, R. vallicola, Colorado, and R. lacustre,
British Columbia.
17. MELAMPSORA LINI (Pers.) Desmaz. Pl. Crypt. (Fasc. 41) 2049.
1850.
Uredo miniata Lint Pers. Syn. Fung. 216. Igol.
On LINACEAE:
Linum Lewis Pursh—Blue Mts., 7,000—8,000 ft., eastern Oregon,
1897 (from phanerogamic specimen in Gray Herb. Harvard Univ.);
Hermiston, Umatilla Co., May 12, 1915, 2664.
Arthur (Jour. Myc. 13: 207. 1907) has shown this species to be
autoecious. He sowed basidiospores from Linum usitatissimum on
the same host and on L. Lewisit and obtained the development of
pycnia and aecia.
18. MELAMPSORA OCCIDENTALIS Jackson, Phytopath. 7: 354. 1917.
On SALIcCACEAE: II, III.
Populus trichocarpa Nutt.—Corvallis, Sept. 1909, 1069, Oct. 15,
1912, 1024 (type), March 12, 1916, G. B. Posey; Trail to Sulphur
Springs, Benton Co., Nov. 2, 1914, 3369; Scott’s, N. of Fort Klamath,
Klamath Co., Sept. 20, 1913, E. P. Meinecke, CrD2; Clatskanie,
Solumbia'Co., Oct. 6; 1914, F. D. Bailey, 3358, Oct. 29, 1914, EF: D:
Bailey, 3306; Sumpter, Baker Co., Aug. 21, 1915, J. R. Weir, 265;
Medical Springs, Union Co., Aug. 1913, J. R. Weir, 777.
This species differs from all other species of Melampsora on Populus
JAD BROOKLYN BOTANIC GARDEN MEMOIRS
in the large size of the urediniospores which are only slightly flattened
and are evenly verrucose-echinulate. The teliospores are much longer
than those of M. Medusae and are thickened at the apex. The charac-
ter of the telial sori suggests that this species may be closely allied
to M. albertensis. The sori are much larger as are also both uredinio-
and teliospores.
This species may be the same as that recently cultured by Weir
and Hubert (Phytopath. 7: 108. 1917), who used telial material from
P. trichocarpa referred to M. Medusae and obtained successful infec-
tion on Larix europea and L. occidentalis. The actual material used
for infection and the aecia resulting have not been seen by the writer
but telial material sent by Dr. Weir from Montana agrees with the
form described above. Aecia from the same locality on L. occidentalis
agree in general with aecia of Melampsora Medusae and M. Bigelowit.
The walls of the aeciospores are however somewhat thinner, I-2 y,
and considerably thickened on opposite sides to 3-5 uw. They measure
17-19 by 19-26 w.. Additional culture work, and a careful comparison
of the resulting aecia with those of M/. Medusae would be desirable.
In any case, the morphological characters of the uredinial and telial
stages are considered sufficient to warrant separation.
19. Melampsora Piscariae sp. nov.
O.and I. Pycnia and aecia unknown.
II. Uredinia hypophyllous, scattered, rounded, 0.3-0.5 mm.
across, early naked, somewhat pulverulent, orange yellow fading to
whitish, ruptured epidermis conspicuous; urediniospores globoid to
ellipsoid, 14-16 by 16-19 yu; wall colorless, 1.5—2 uw in thickness, finely
and closely verrucose-echinulate; paraphyses numerous, intermixed
with the spores, capitate, smooth or with an occasional conical echinu-
late marking, 32-644 long; heads 12-18 y broad, wall uniformly
thick, 2.5-4 um.
III. Telia not seen.
ON EUPHORBIACEAE:
Piscaria setigera (Hook.) Piper (Eremocarpus setigerus Benth.)—
Corvallis, Sept. 20, 1914, 3308, type.
Known only from the type locality.
This species is referred to the genus Melampsora with considerable
confidence in spite of the absence of telia, on account of the structure
of the sorus, the character of the spores, and the presence of scattered
capitate paraphyses.
20. MELAMPSORELLA ELATINA (A. & S.) Arthur, N. Amer. Flora 7:
Tits, ViOO7.
Aecidium elatinum Alb. & Schw. Consp. Fung. 121. 1805.
JACKSON: UREDINALES OF OREGON 213
Melampsorella Cerastii (Pers.) Schroet. Krypt. Flor. Schles. 3:
366. 1887.
On PINACEAE: I.
Abies grandis Lindl.—Mary’s River, west of Wren, Benton Co.,
Aug. 2, 1914, 1297; Sumpter, Baker Co., July 20, 1913, J. R. Weir,
270.
Abies lasiocarpa (Hook.) Nutt.—Crater Lake, Klamath Co.,
Sept. 9, 1916, J. R. Weir, 209; Sumpter, Baker Co., July, 1913, J. R.
Weir, 274.
On CARYOPHYLLACEAE: II, III.
Cerastium vulgatum L.—Corvallis, May I, 1915, 2667.
Cerastium viscosum L.—Corvallis, June 28, 1915, 3070.
Stellaria borealis Bigel.—Corvallis, April 5, 1914, 1287.
This rust, which is doubtless common throughout the state, is
remarkable in that both stages develop from a perennial mycelium.
The aecial stage forms large or small witches’ brooms on the branches
of various species of Abies, each leaf of which bears the conspicuous
aecia in two rows on the under surface of the leaves.
The life history was first worked out by Fischer (Zeitschr. fiir
Pflanzenkr. 11: 321. 1901) and has been amply confirmed by other
European investigators. A summary of this work has been made by
Klebahn (Die Wirtsw. Rostpilze 397. 1904).
In America Arthur (Mycol. 4:58. 1912), using aecial material on
Abies lasiocarpa collected in Colorado, has succeeded in obtaining
infection resulting in uredinia on Cerastium oreophilum.
21. MELAMPSOROPSIS PIPERIANA Arthur, N. Amer. Flora 7: 120.
1907.
ON ERICACEAE:
Rhododendron Californicum Hook.—Newport, Lincoln Co., June
2, 1892, A. Isabel Mulford (Specimen in Herb. N. Y. Bot. Gard. and
in Herb. J. C. Arthur), May 16, 1914, G. H. Godfrey, 1280; Larch
Mt., Multnomah Co., Aug. I910, rzz8; Parmelia Lake, near Mt.
Jefferson, July 3, 1914, F. D. Bailey, 1939; Trail to Hanging Valley,
near Mt. Jefferson, Aug. 11, 1914, H. P. Barss & G. B. Posey, 1623.
22. MELAMpPSOROPSIS PyROLAE (DC.) Arth. Résult Sci. Congr. Bot.
Vienne 338. 1906.
Aecidium (?) Pirolae DC. Fl. Fr. 6: 99. 1815.
Aecidium conorum Piceae Reess, Abh. Nat. Ges. Halle 11: 102.
1869.
- Chrysomyxa Pirolae Rostr. Bot. Centr. 5: 127. 1881.
Peridermium conorum Piceae Arth. & Kern, Bull. Torrey Club 33:
431. 1906.
214 BROOKLYN BOTANIC GARDEN MEMOIRS
On PINACEAE: O, I.
Picea Engelmanit Parry—Sumpter, Baker Co., Sept. 25, 1909,
G. G. Hedgcock, 1916.
On PyrovaceEaE: II, III.
Pyrola secunda L.—North slope Mt. Hood, Aug. 7, 1914, 1607;
Trail to Elk Meadows, Hood River Co., July 23, 1915, 3061; Columbia
Highway, Multnomah Co., Aug. 19, 1916, J. R. Weir, 269.
Pyrola sp.—Grant’s pas Josephine Co:, Sept. 3, 1or6;))ae
Weir, 268.
The genetic relation of this species with Peridermium conorum
Piceae was first suggested by Rostrup (I. c.). So far as the writer is
aware the first culture work confirming this observation was made by
Fraser (Mycol. 4: 183. 1912), who succeeded in obtaining infection
resulting in pycnia and aecia on the cones of Picea mariana and P.
canadensis, following exposure to germinating telia on Pyrola americana
and P. elliptica.
23. Milesia Polystichii Wineland n. sp.
O. and I. Unknown.
II. Uredinia hypophyllous, scattered, roundish, 0.2-0.3 mm. across,
bullate, brownish yellow, tradily dehiscent by a central pore, peridium
well developed, cells above polygonal, approximately isodiametric,
diameter about 7 yw, cells at the sides elongated to 21 uw, outer walls
2-2.5, inner walls 2.5-3 4; urediniospores obovoid, ellipsoid, or
oblong, 18-23 by 26-35, wall colorless, 1.5-2.5 4 in thickness,
strongly and sparsely echinulate, pores obscure.
III. Telia unknown.
ON POLYPODIACEAE:
Polystichum munitum (Kaulf.) Presl.—Grant’s Pass, Josephine
Co., Sept. 5, 1916, J. R. Weir, 260 (type).
This species was separated from material referred to Hyalopsora
laeviuscula in the writer’s herbarium by Miss Grace O. Wineland who
has been studying the fern rusts of North America in this laboratory.
24. PUCCINIASTRUM ABIETI-CHAMAENERII Kleb. Prings. Jahrb. f.
Wiss. Bot. 34: 387. 1900.
On PINACEAE: I.
Abies grandis Lind|.—Dee, Hood River Co., July 23, 1915, 3355.
Abies lasiocarpa Nutt.—North slope Mt. Hood, 4,500 ft., Aug. 9,
1914, 3295.
On ONAGRACEAE: II, III.
Chamaenerion angustifolium (L.) Scop.—Bonneville, Multnomah
Co., Aug. II, 1910, 7075; Garden Home, Multnomah Co., Aug. 1911,
1990; Southwest slope Mt. Jefferson, July 3, 1914, F. D. Bailey,
JACKSON: UREDINALES OF OREGON 215
3247; Odell, Hood River Co., Aug. 5, 1914, 1678; Crater Lake,
Klamath Co., Sept. 21, 1913, E. P. Meinecke, Cr Pk D (2) 2; Portland,
Aug. 24, 1915, E. Bartholomew (Barth., N. Am. Ured. 1482).
This species is separated from P. pustulatum largely on the basis
of culture investigations. All of the culture work has been conducted
with the above host species or other members of the same genus or
section of Epilobium. European investigators have amply demon-
strated the connection of this form with aecia on Abies (Klebahn,
Die Wirtsw. Rostpilze 393. 1904). In America, Fraser, working in
Nova Scotia (Mycol. 4: 176. 1912), was the first to conduct culture
experiments. He obtained, in three trials, the development of aecia
on Abies balsamea, from sowings with teliosporic material from C.
angustifolium. With the aecia thus obtained he sowed back to
Chamaenerion and obtained uredinia. Weir & Hubert (Phytopath.
6: 373. 1916) conducted similar work with Idaho material and suc-
ceeded in obtaining the development of aecia on Abies lasiocarpa.
The aecia thus obtained were sown back on the telial host (Phytopath.
7: 109. 1917), with the result that uredinia were developed in
abundance.
25. PUCCINIASTRUM GaALII (Link) Fischer, Ured. d. Schweiz 471.
1904.
Caeoma Galu Link, in Willd. Sp. Pl. 62: 21. 1825.
On RusBiAcEseE: II.
Galium triflorum Michx.—Corvallis, April 29, 1914, F. D. Bailey,
1992, May 9, 1914, r991, July 5, 1914, H. P. Barss, 1996, June 29,
1914, G. B. Posey, 1373, July 10, 1915, 3104; Oregon City, Clackamus
Co., Aug. 20, 1915, E. Bartholomew, 5934 (Barth. N. Am. Ured.
1679); Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 187.
This species is known to the writer from North America only from
the above collections and a specimen collected by J. W. Macoun in
British Columbia in 1915, one from Palmer Lake, Colorado, Sept. 6,
1913, by E. Bethel, both in the Arthur Herbarium, and a collection
made by Dr. H. Fitzpatrick and the writer at Michigan Hollow Swamp
near Ithaca, N. Y., July, 1916. All are on G. triflorum. The species
is evidently common in western Oregon and if aecia are developed,
doubtless occur on Abies grandis. Field observations made by the
writer, however, would indicate that this rust winters over in the
uredinial stage.
26. PUCCINIASTRUM GOODYERAE (Tranz.) Arth. N. Am. FI. 7: 105.
1907.
Uredo Goodyerae Tranz. Trudi S. Peterb. Obshch. Est. Otd. Bot.
2312380 1853;
216 BROOKLYN BOTANIC GARDEN MEMOIRS
ON ORCHIDACEAE: II,
Goodyera Menziesi1 (Lindl.) Morong.—Parmelia Lake, West slope
Mt. Jefferson, July 3, 1914, F. D. Bailey, 7627; North slope Mt.
Hood, Aug. 9, 1914, 1620.
27. PUCCINIASTRUM MyrrtILii (Schum.) Arth. Résult. Sci. Congr.
Bot. Vienne 337. 1906.
Aecidium ? Myrtilla Schum. Enum, PI. Saell. 2: 227. 1803.
Pucciniastrum Vacciniorum (DC.) Dietel, in E. & P. Nat. Pfl.
Be en LOO.
ON VACCINIACEAE: II.
Oxycoccus macrocarpus (Ait.) Pursh—Astoria, Clatsop Co., Aug.
1916, G. M. Darrow, comm. C. L. Shear 2905.
Vaccinium caespitosum Michx.—Mary’s Peak, Benton Co., Aug.
15, LO14;,.7577, 1520:
Vaccinium macrophyllum (Hook.) Piper—Whitewater Ranger
Station, West slope Mt. Jefferson, Aug. 12, 1914, H. P. Barss &
G. B. Posey, 3314; Ashland, Jackson Co., Sept. 10, 1914, 33106.
Vaccinium ovatifolium Sm.—Whitewater Ranger Station, West
slope Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 3315.
Vaccinium sp.—North slope Mt. Hood, 4,000 ft., Aug. 7, 1914,
1606, 1609; Sucker Creek, Josephine Co., July 27, 1913, E. P. Mein-
ecke, Sz (D6) D3.
Clinton (Rep. Conn. Agr. ‘Exp. Sta. 1909-1910: 719. 1911)
was the first to show that the aecial stage of this species occurred on
Tsuga canadensis. He successfully infected Gaylussacia baccata by
sowing with aeciospores from Tsuga, resulting in the development
of the typical uredinia of this species.
Fraser in 1912 (Mycol. 5: 237. 1913) confirmed Clinton’s work
by obtaining the development of aecia on the leaves of Tsuga canadensis
following sowings from teliosporic material on Vaccinium canadense.
The same author in 1913 (Mycol. 6: 27. 1914) obtained aecia on
Tsuga canadensis following sowing of teliosporic material from Gay-
lussacia resinosa. The aecia developed in these experiments are
similar to those of Peridermium Peckii, but may represent an unde-
scribed form.
No aecia collected in the west have been referred to this species
though they doubtless occur on Abies or Tsuga heterophylla.
28. PUCCINIASTRUM PUSTULATUM (Pers.) Dietel, in E. & P. Nat. Pfl.
Tete Aa 1807.
Uredo pustulata Pers. Syn. Fung. 219. 1801.
ON ONAGRACEAE:
Epilobium adenocaulon Haussk.
Corvallis, Oct. 29, 1911, F. DB:
JACKSON: UREDINALES: OF OREGON Zb7
Bailey, 1773, Nov24, 1911, F..D. Bailey, rz7z, June 18, 1914,°F. D.
Bailey, 3218, July 29, 1914, 1450; Hilgard, Union Co., July 10, 1914,
1533, 1535, Glendale, Douglass Co., July 17, 1914, 1505, North slope
Mt. Hood, Aug. 7, 1914, 1488; Whitewater Ranger Station, West
slope Mt. Jefferson, H. P. Barss & G. B. Posey, 3219; Ashland,
Jackson Co., Sept. 10, 1914, 3221.
Epilobium brevistylum Barbey—Corvallis, July 14, 1914, G. B.
Posey, 3220.
No successful culture work has been conducted with this form, as
here interpreted, either in Europe or America. Aecia doubtless occur
on species of Adzes.
From field observations it is quite evident that in western Oregon
at least this species overwinters in the uredinial stage.
29. PUCCINIASTRUM PYROLAE (Pers.) Dietel, in E. & P. Nat. Pfl.
eer Ar. ESOT.
Aecidium Pyrolae Pers. Gmel. Syst. Nat. 2: 1473. 1791.
Uredo Chimaphilae Peck, Ann. Rep. N. Y. State Mus. 46: 33.
1893.
ON PYROLACEAE:
Chimaphila umbellata (L.) Nutt. (C. occidentalis Rydb.)—Spencer
Creek, Klamath Co., 5,000 ft., July 10, 1903, E. B. Copeland, 3774
(Sydow, Ured. 7795); Whitewater Creek along trail to Hanging Valley,
Mt. Jefferson, Aug. 11, 1914, H. P. Barss & G. B. Posey, 1908; North
slope Mt. Hood, Aug. 7, 1914, 1674.
Pyrola secunda L., Klamath Co., July 10, 1903, E. B. Copeland
(Sydow, Ured. 1791).
30. PUCCINIASTRUM SPARSUM (Wint.) E. Fischer, Beitr. Krypt.
Schweiz 27: 469. 1904.
Melampsora sparsa Wint. in Rab. Krypt. Fl. 1!: 245. 1881.
ON ERICACEAE:
Arbutus Menziesit Pursh—Myrtle Creek, Douglass Co., June 8,
1914, F. D. Bailey, 1837; Glendale, Douglass Co., July 17, 1914,
1298; Ashland, Jackson Co., Sept. 10, 1914, 1838; Corvallis, April,
O11, 3374; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir,
244.
Arctostaphylos Manzanita Parry—Grant’s Pass, Josephine Co.,
Sept. 3, 1916, J. R. Weir, 245, 247.
Arctostaphylos nevadensis A. Gray—Northwest slope Mt. Jefferson,
Aug. 14, 1914, H. P. Barss & G. B. Posey, 3290.
No aecial collections have been referred to this species in America.
Fischer (Cent. fiir Bakt. 46: 333. 1916) has cultured this species.
He used germinating telial material on Arctostaphylos alpina and sowed
218 BROOKLYN BOTANIC GARDEN MEMOIRS
on species of Abies and Picea, obtaining the development of pycnia
and aecia on Picea excelsa. This is the only case in which culture
investigations have shown the aecia of Pucciniastrum to occur on
Picea.
31. UREDINOPSIS COPELANDII Sydow, Ann. Myc. 2: 34. Feb. 1904.
Uredinopsis Atkinsoniu Magn. Hedwigia 43: 123. Mar. 1904.
Peridermium balsameum Peck, Rept. N. Y. State Mus. 27: 104.
1975... Dp:
On PINACEAE: I.
Abies grandis Lindl.—Trail to Sulphur Springs, Corvallis, Benton
Co., Nov. 7, 1914, 3339; Corvallis, Feb. 2, 1914, 3300.
Abies nobilis Lind|l.—Mary’s Peak, Benton Co., Feb. 7, 1914,
F. D. Bailey, 3337, Aug. 15, 1914, 3334, 333513339 3338.
On PoLypopiaAcEAE: II, III.
Athyrium cyclosorum Rupr.—Hoover, Linn Co., Aug. 19, 1914,
H. P. Barss & G. B. Posey, 3032; Mary’s Peak, Benton Co., Aug.
15, 1914, 3033, 3034; Elk City, Lincoln Co., Aug..20, 1914 4f2a75e
Hood River Co., July 23, 1915, 3031; Grant’s Pass, Josephine Co.,
Sept. 5, 1916, J. R. Weir, 258; Yaquina, Lincoln Co.} July 177m
3035.
There seems to be no good reason for separating U. Copelandit
Sydow from U. Atkinsoni. All gradations in the length of the beak
of the urediniospores are found on the above collections. Most of the
material has urediniospores with long beaks and some of the collec-
tions show spores with both long and short beaks.
Field observations as well as a study of morphological characters
would support the view that the aecia commonly referred to Peri-
dermium balsameum occurring on Abies grandis and A. nobilis in
western Oregon are genetically connected.
Fraser (Mycol. 5: 236. 1913) has cultured U. Atkinsonit by
sowing aeciospores of Peridermium balsameum on Aspidium Thelepteris
followed sparingly by the development of uredinia.
32. UREDINOPsIS PTERIDIS Dietel & Holway, Ber. Deuts. Bot. Ges.
1355330; etoose
Aecidium pseudo-balsameum Diet. & Holw. Erythea 7: 98. 1899.
Peridermium pseudo-balsameum Arth. & Kern, Bull. Torrey Club
33: 430: *T906.
On PINACEAE: I.
Abies amabilis (Loud.) Forb.—Whitewater Creek, near Mt. Jeffer-
son, Aug. 11, 1914, H. P. Barss & G. B. Posey, 3294.
Abies grandis Lindl.—Corvallis, Aug. 1910, 3299, May 8, 1909,
comm. Clarence D. Learn, April 29, 1914, F. D. Bailey, 3303; Ump-
JACKSON: UREDINALES OF OREGON 219
qua Nat. Forest, near Diston, Lane Co., Oct. 27, 1909, Geo. G. Hedg-
cock; Philomath, Jan. 6, 1914, 3298; Wren, Benton Co., Aug. 3,
1914, 3296; Ashland, Jackson Co., Sept. I0, 1914, 3297; Mary’s
Peak, Benton Co., Feb. 7, 1914, G. H. Godfrey, 3307, Aug. 15, 1914,
3302; N. slope Mt. Hood, 4,000 ft., Aug. 9, 1914, 1616.
On PotypopiaAcEsE: I], III.
Pteridium aquilinum pubescens Underw.—Corvallis, Sept. 1909,
1142, July, 1910, 1082, Aug. 1910, 1058, Oct. 6, 1914, 3109; Bonneville,
Multnomah Co., Aug. II, 1910, 1076; Scappose, Columbia Co.,
July 25, 1911, 1067; North slope Mt. Hood, Aug. 9, 1914, 1617;
Ashland, Jackson Co., Sept. 10, 1914, 1993; Portland, Aug. 24, 1915,
E. Bartholomew (Barth. N. Am. Ured. 1485); Grant’s Pass, Josephine
Gay Sept. 5, 1916, J. R. Weir, 250.
This species in all its stages is very common in western Oregon
and the association of the infected aecial and telial hosts is everywhere
apparent.
From field observations made by the writer and others it has
been assumed that Peridermium pseudo-balsameum was the aecial
stage of this species. Recently Weir and Hubert (Am. Jour. Bot.
4: 328-332. 1917) have conducted cultures showing the genetic con-
nection of this species with aecia on Abies grandis. The authors
evidently do not consider the aecia identical with P. pseudo-balsameum.
The description which they give, however, agrees very well with the
type of that species.
According to the writer’s present interpretation, there are two
closely related species of Peridermium on Abies grandis in western
Oregon. One is to be referred to P. balsameum and is presumably
genetically connected with Uredinopsis Copelandii (cf. 31). The
other is P. pseudo-balsameum and is genetically connected with the
species under discussion.
The walls of the aeciospores in P. balsameum are considerably
thinner than those of P. pseudo-balsameum. In the former they are
I-1.5 uw while in the latter they are 2—2.5 yw in thickness.
PUCCINIACEAE
33. EARLEA SPECIOSA (Fr.) Arth. Résult Sci. Congr. Bot. Vienne 341.
19006.
Aregma speciosa Fr. Syst. Myc. 3: 496. 1832.
Phragnuidium speciosum Cooke, Grevillea 3: 171. 1875.
On. ROSACEAE:
Rosa gymnocarpa Nutt.?—I, Austin, Grant Co., Aug. 1915, J. R.
Weir, 188.
220 BROOKLYN BOTANIC GARDEN MEMOIRS
34. GYMNOSPORANGIUM BETHELI Kern, Bull. Torrey Club 34: 459.
1907.
Roestelia Betheli Kern, Bull. Torrey Club 34: 461. 1907.
On MAtaceaeE: I.
Crataegus Douglas Lind|.—Joseph, Wallowa Co., Aug. 19, 1899,
C. L. Shear (Ellis & Ev. Fungi Columb. 1480).
ON JUNIPERACEAE: III.
Juniperus occidentalis Hook.—Whitney, Baker Co., Aug. 1915,
JoR: Weir, 770.
Another specimen on Crataegus sp. indet. from eastern Oregon
(ex herb. Ellis) bearing no date is in the Arthur herbarium and has
been examined by the writer.
The life history of this species was first demonstrated by Arthur
(Jour. Myc. 14: 23. 1908) and later repeatedly confirmed. Telia
are otherwise known only on Juniperus scopulorum from Colorado,
Idaho and Montana. Aecia are known to occur only on Crataegus
sp. in the Rocky Mt. region and in eastern Oregon and Washington.
35. GYMNOSPORANGIUM BLASDALEANUM (Dietel & Holway) Kern,
Bull NV] Boe. Gard."7 3.437. DOr:
Aecidium Blasdaleanum Dietel & Holway, Erythea 3: 77. 1895.
Gymnosporangium Libocedri Kern, Bull. Torrey Club 35: 509.
1908.
On Mataceae: I.
Amelanchier florida Lind|l.—Eugene, Lane Co., July I1, 1914,
G. B. Posey, 3276; Albany, Linn Co., June 11, 1913,.D. W. Kune
baugh, 3171; Cottage Grove, Lane Co., June 13, 1913, 37066, [ume
20, 1913, C. E. Stewart, 3777; Lebanon, Linn Co., Aug. 2) iaae
F. D. Bailey, 3774; Crater Lake, Klamath Co., 7,000 ft., Sepia
1913, E. P. Meinecke, Cr Pk D 11; Jackson Co.,. July, tome
Reimer, 7791; Between Albany, and Lebanon, Linn Co., June 13,
1913, C. E. Roberts, 1788; Lost Prairie, Sept. 1891, M. Craig; Halsey,
Linn Co., June 9, 1913, 3170; Corvallis, July 29, 1915, 3750; Ashland,
Jackson Co., Sept. 10, 1914, 3047; N. W. Mt. Jefferson, Whitewater
station, Aug. 17, 1914, H. P. ‘Barss & G. B.-Posey, 3043; Aueageee
1916, H. P. Barss.
Crataegus Douglasii Lindl.—Halsey, Linn Co., June 9, 1913, 3214;
Albany, Linn Co., D. W. Brumbaugh, 32172; Cottage Grove, Lane
Co., May 21, 1913, 3169, June 14, 1913, 32090; Eugene, LaneiCos
May 8, 1913, 3173.
Cydonia japonica Pers.—Eugene, Lane Co., June, 1914, G. H.
Godfrey.
Cydonia vulgaris L., Halsey—Linn Co., June 9, 1913, 3166; Irving,
Lane Co., 1913, Comm. S. J. Quigley, 287z; Creswell; Lane Co.,
JACKSON: UREDINALES OF OREGON 221
May 5, 1913, Comm. K. V. Miller, 7873; Eugene, Lane Co., Aug.
1912, 1054; Talent, Jackson Co., May 18, 1916, F. C. Reimer.
Pyrus baccata Linn.—Lorane Valley, Lane Co., May, 1915, C. E.
Stewart, 3367.
Pyrus communis L.—Kerby, Josephine Co., June 1, 1899, Comm.
E. F. Meissner, 7845; Brownsville, Linn Co., May 24, 1913, D. W.
Brumbaugh, rgzz; Eugene, Lane Co., May 8, 1913, 3172, April 22,
1915, 2620.
Pyrus diwersifolia Bong. (P. rwularis Dougl.)—Cottage Grove,
Pane Co.,))une 13, 1913, 3775; 3211:
Pyrus toensis (Wood) Bailey—Cottage Grove, Lane Co., June 13,
1913, 1554; Eugene, Lane Co., May 21, 1913, 3270.
Pyrus malus L.—Eugene, Lane Co., July 10, 1913, J. O. Holt,
moe Cottage Grove, Lane Co:, May 23, 1913, C. E. Stewart, ror3,
June 20, 1915, C. E. Stewart, 887.
Sorbus aucuparia Linn.—Cottage Grove, Lane Co., June 13, 1913,
3176; Eugene, Lane Co., June 1, 1914, G. H. Godfrey, 3222.
Sorbus hybrida Linn.—Cottage Grove, Lane Co., May 21, 1913,
fmge. June 13, 1913, 3207.
ON JUNIPERACEAE:
Libocedrus decurrens Torr.—Eugene, Lane Co., Feb. 28, 1913,
3213, Feb. 21, 1914, F. D. Bailey, 1675, Mar. 20, 1914, 3070; Breiten-
bush Hot Springs, Marion Co., Mar. 27, 1915, E. A. Hartley, 2621;
Cottage Grove, Lane Co., Mar. 8, 1914, C. E. Stewart, 7888; Ashland,
Jackson Co., Sept. 10, 1914, 17839; Corvallis, Mar. 30, 1915, J. G.
Corsaut, gor; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir,
185.
This species is very common in its aecial stage on all the native
members of the Malaceae as well as most of the cultivated fruits and
ornamental plants belonging to this family which may occur in the
range of the incense cedar. There is considerable evidence also that
the disease is gradually spreading beyond the natural range of the
telial host. Observations made at Corvallis support this view.
Previous to 1915 no specimens of this species had been collected in
Benton County though careful search had been made many times.
The incense cedar does not occur naturally in that locality but is
frequently planted for ornament. There are several fine examples on
the campus at the Oregon Agr. College. In 1915 a very sparing in-
fection of the aecial stage was found in the vicinity on native hosts
only, and the cedar trees on the campus were found to be sparingly
infected. Reports of the occurrence on quince have come from Salem,
far north of the natural range of the incense cedar. The writer
believes that the disease will gradually spread throughout the Wil-
222 BROOKLYN BOTANIC GARDEN MEMOIRS
lamette Valley on the incense cedars planted for ornament and, in
certain cases, will become a serious menace to cultivated pears and
quinces.
The life history of the species was first worked out by Arthur
(Mycol. 1: 252. 1909; 4: 57. 1912). He succeeded in showing
that aecia occurred on Crataegus and Amelanchier. The telial material
used for the cultures was collected at Eugene, Oregon, by Prof. A. R.
Sweetzer. The writer has also studied this species in some detail
(Phytopath. 4: 261-269. 1914; Ore. Expt. Sta. Biennial Crop Pest
Rep. II: 204-212. 1915) and has reported the results of culture
work and field observations. Similar work is also briefly reported by
O’Gara (Science N.S. 39: 620-621. 1914). The previous records of
the occurrence of this species on Malus floribundus Siebold and Sorbus
sambucifolia Roem. made by thé writer (I. c.) should be corrected to
read Pyrus toensis and S. aucuparia respectively. For the correct
determination of these hosts the writer is indebted to Prof. W. W.
Eggleston.
This species has since been successfully cultured in the greenhouse
on Pyrus sinensis by Prof. H. P. Barss. The writer, using aecial
material on quince, the result of infection experiments made in 1914,
has obtained sparing infection on Libocedrus resulting in telia which
matured in February 1915. The trees were kept in the greenhouse
at the Oregon Agr. College.
36. GYMNOSPORANGIUM HARKNESSIANUM (EIl. & Ev.) Kern, Bull.
N.Y. Bot. Gardiiy-14aa> ort.
Roestelia Harknessiana Ell. & Ev. Kern, Bull. Torrey Club 34:
462. M1907.
On MALaAcEaE: I.
Amelanchier alnifolia Nutt.—Redmond, Crook Co., July 2, 1914,
1393; Fort Rock, Lake Co., Oct. 10, 1915, Wendover, 3375.
This very interesting species has otherwise been reported only
from northern California. The telia form is unknown. In the col-
lection made by the writer listed above, the aecia occurred only on
fruits and twigs. There was every evidence that this fungus is
perennial. Some of the specimens show fresh aecia on branches having
four annual rings, surrounding or extending from cankered areas
bearing evidence of old aecial cups. There is slight hypertrophy.
The branches are frequently girdled and killed.
37. GYMNOSPORANGIUM JUNIPERINUM (L.) Mart. Fl. Crypt. Erlang.
393i 1St7.
Tremella juniperina L. Sp. Pl. 1157. 1753.
JACKSON: UREDINALES OF OREGON 223
On MALAceseE: I.
Sorbus occidentalis (S. Wats.) Greene—North slope Mt. Jefferson,
along trail to Hanging Valley, Aug. 15, 1914, H. P. Barss & G. B.
Posey, 7395; Columbia Highway, Multnomah Co., Aug. 19, I916,
J. R. Weir, 2709.
ON JUNIPERACEAE: III. ;
Juniperus sibirica Burg.—North slope Mt. Jefferson, Aug. 26,
1916, H. P. Barss, 3399.
The genetic connection of the forms of this alpine species has
been abundantly demonstrated by European investigators, first by
Hartig (Lerb. Baum-Kr. 133. 1882), and later by many others.
Arthur in 1911 (Mycol. 4: 57. 1912), using telial material from J.
sibirica collected in Colorado, succeeded in obtaining infection result-
ing in pycnia only on Sorbus americana. The species is known in
America only from the Rocky and Cascade Mountains of the United
States and Canada.
38. GYMNOSPORANGIUM JUVENESCENS Kern, Bull. N. Y. Bot. Gard.
72 449. - LOL. ‘
On Matacece_: I.
Amelanchier sp.—Hurricane Creek, Wallowa Co., July 24, 1897,
E. P. Sheldon, 8622.
On JUNIPERACEAE: III.
Juniperus scopulorum Sarg.—White Pine, Baker Co., Aug. 1915,
J. R. Weir, 769.
This species causes witches’ brooms on the telial host somewhat
similar to the eastern G. nidus-avis Thax.
Arthur has repeatedly cultured it, showing that the aecia occur
on Amelanchier and Sorbus (Jour. Myc. 13: 203. 1907; 14: 18.
foeo; Mycol. 1:'239. 1909; 4: 195. I912).
39. GYMNOSPORANGIUM KERNIANUM Bethel, Mycologia 3:157. IgI1.
ON JUNIPERACEAE: III.
Juniperus occidentalis Hook.—Redmond, Crook Co., July 2, 1914,
1392, May 15, 1915, 3390.
The above specimens are somewhat doubtfully referred to this
species. The witches’ brooms are large and open, sometimes reaching
2-3 feet in diameter. The teliospores are somewhat more tapering
at the apex than is typical for the species and average shorter and
somewhat narrower, 19-22 by 45-65 u. The only aecia collected in
the vicinity are properly referred to G. Harknessianum. There was
no very direct field evidence, and unless the above collections repre-
sent an undescribed form there is little possibility that the two can
be genetically connected. Arthur (Mycol. 4:62. 1912) has cultured
16
224 BROOKLYN BOTANIC GARDEN MEMOIRS
G. Kernianum on Amelanchier but obtained the development of
pycnia only. Field observations and collections by Bethel in Colorado
indicate strongly that the aecial stage occurs on Amelanchier. The
Roestelia, however, is quite different from R. Harknessiana.
40. GYMNOSPORANGIUM KOREAENSE (P. Henn.) Jackson, Jour. Agr.
Research 5: 1006. 1916.
Roestelia koreaensis P. Henn. in Warburg, Monsunia 1: 5. 1899.
Gymnosporangium asiaticum Miyabe, Bot. Mag. Tokyo 17: 34.
1903. (Hyponym.)
Gymnosporangium Haraeanum Syd. Ann. Myc. 10: 405. 1912.
Gymnosporangium chinense Long, Jour. Agr. Research 1: 353.
1914.
ON MALaAcEsE: I.
Pyrus sinensis Lind|.—Portland (Orient), June 11, 1914, 2666.
On JUNIPERACEAE: III.
Juniperus chinensis L.—Portland (Orient), March 29, 1915, 2668.
This species has been shown by the writer (I. c.) to have been
established at Portland (Orient), Oregon, on trees imported from
Japan. It has been cultured on Pyrus sinensis and Cydonia vulgaris.
41. GYMNOSPORANGIUM NELSONI Arth. Bull. Torrey Club 28: 665.
I9Ol.
ON JUNIPERACEAE: III.
Juniperus occidentalis Nutt.—Austin, Grant Co., April, 1916, J. R.
Weir, 257.
Juniperus scopulorum Sarg.—Whitman Nat. Forest, Aug. 1915,
J. R. Weir, 766.
This species causes conspicuous galls on the branches of Juniperus.
The aecial stage has been collected on Amelanchier, Cydonia, Pera-
phyllum, Pyrus and Sorbus.
Arthur (Mycol. 4: 61. 1912; 7: 78. 1915) has conducted cul-
tures, using telial material from Colorado. Weir & Hubert (Phyto-
path. 7: 109. 1917) have recently confirmed these results, using
material collected in Montana, on J. communis and J. scopulorum.
42. GYMNOSPORANGIUM NOOTKATENSIS (Trel.) Arth. Am. Jour. Bot.
3: 44. TOr6.
Uredo nootkatensis Trelease, Alaska Harr. Exped. 5: 36. 1904.
Uredo Chamaecyparidis-nutkaensis Tubeuf, Nat. Zeits. Forst.-
Landw..2:-9ly T0n4:
ON JUNIPERACEAE:
Chamaecyparis nootkatensis (Lamb.) Spach—North slope Mt.
Jefferson, trail to Hanging Valley, Aug. 15, 1914, H. P. Barss & G. B.
JACKSON: UREDINALES OF OREGON 225
Posey, 1394; Whitewater Ranger station, Aug. 28, 1916, H. P. Barss;
Foot of Mt. Jefferson, Aug. 28, 1916, H. P. Barss.
The material collected by Barss and Posey in 1914 contained
teliospores in the uredinia and forms the basis of the transfer of the
very interesting and much discussed Uredo nootkatensis to Gymno-
sporangium. A full account of the history of this species has been
given by Arthur (I. c.). In the collections of 1916 made in the same
locality by Prof. Barss, teliospores were found in great abundance
with the uredinia, and in many sori predominated. The uredinio-
spores were germinated in this laboratory and the germ tubes found to
develop in the usual way for urediniospores.
43. GYMNOSPORANGIUM SorsI (Arth.) Kern, Bull. N. Y. Bot. Gard. 7:
age. I91L.
Aecidium Sorbi Arth. Bull. Torrey Club 33: 521. 1906.
ON MALAcEAE: I.
Sorbus occidentalis (S. Wats.) Greene—Whitewater Ranger station,
Mt. Jefferson, Aug. 28, 1916, H. P. Barss.
There is little doubt that the suggestion of the genetic relationship
of this species with Gymnosporangium nootkatensis (cf. 42) originally
made by Kern (Science 31: 833. I910) and later re-affirmed by
Arthur (Am. Jour. Bot. 3: 43-44. 1916) will prove to be correct.
The above collection extends the range of the aecia to correspond
exactly with the range of the known collections of uredinia and is the
most southern record.
It seems best, however, for the purpose of this list to retain the
above name till actual cultures confirming the prediction have been
made.
44. GYMNOSPORANGIUM TUBULATUM Kern, Bull. N. Y. Bot. Gard. 7:
Nets “TORT.
Roestelia tubulata Kern; in M. E. Jones, Bull: Univ. Mont. 61: 64.
I9IO.
On MALACEAE: I.
Crataegus Douglasu Lind|l.—Minam River, Wallowa Co., Oct. 5,
1897, E. P. Sheldon, 9061; Wallowa Nat. Forest, Sept. 28, 1910,
G. G. Hedgecock, 1944.
The above collections were found in the Arthur herbarium at the
Purdue University Experiment Station. The specimens show chiefly
foliage infection, though the first-mentioned collection also includes
infected fruit.
‘Weir (Phytopath. 5: 218. 1915) has recently demonstrated by
cultures that the telia, which were previously unknown, occur on the
twigs of Juniperus scopulorum forming irregularly lobed galls. Telia
226 BROOKLYN BOTANIC GARDEN MEMOIRS
have been collected only in Idaho and western Montana. Weir and
Hubert in 1916 (Phytopath. 7: 109. 1917) have confirmed the above
results.
45. KUNKELIA NITENS (Schw.) Arth. Bot. Gaz. 63: 504. 1917.
Aecidium nitens Schw. Schrift. Nat. Ges. Leipzig 1: 69. 1822.
ON ROSACEAE:
Rubus nigrobaccus Bailey—Freewater, Umatilla Co., June 27,
1913, F. D. Bailey, 7743.
Rubus vitifolius Cham. & Schlecht. (cult. loganberry)—LaGrand,
Union Co. July 209 1914 "C2 ©. (Cate; ra5r:
Kunkel’s results (Bull. Torrey Club 40: 361.. 1913; 43: 55a:
1916; Amer. Jour. Bot. I: 37. 1914) indicate that two rusts on
Rubus, both commonly referred to Gymnoconia interstitialis or Caeoma
nitens, occur in North America, one a short-cycled form having the
morphology of a Caeoma, the other a brachy-form with caeomoid
aecia and telia of the type of Puccinia (P. Peckiana Howe). Arthur
(1. c.) has recently based the genus Kunkelia on the short-cycled form.
The inclusion of the Oregon collections under Kunkelia follows
the disposition made of them by Arthur.
46. NYSSOPSORA ECHINATA (Lev.) Arth. Result. Sci. Congr. Bot.
Vienne 342. 1906.
Triphragmium echinatum Lev. Ann. Sci. Nat. III. 9: 247. 1848.
ON UMBELLIFERAE:
Ligusticum Cusicki Coult. & Rose—Steins Mts., Harney Co.,
Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 340).
Ligusticum purpureum Coult. & Rose—North slope Mt. Jefferson,
Aug. 13, 1914, H. P. Barss & G. B. Posey, 2540.
47. PHRAGMIDIUM DISCIFLORUM (Tode) J. F. James, Contr. U. S.
Nat. Herb. 3: 276. A895.
Ascophora disciflora Tode, Fungi Meckl. 1: 16. 1790.
ON ROSACEAE:
Rosa sp. cult.—Empise, Coos Co., Oct. 2, 1911, comm. J77R:
Brown, 3154; Portland, May 10, 1914, comm. W. C. Dietz, 3156;
Eugene, Lane Co., June 1, 1914, G. H. Godfrey, 3147; Sutherlin,
Douglass Co., March 9, 1915, comm. Gladys Franz, 2571.
48. PHRAGMIDIUM IMITANS Arth. N. Am. Flora 7: 165. 1912.
On ROSACEAE:
Rubus leucodermis Dougl., Philomath, May 10, 1914, 7830.
Rubus neglectus Pk., Ore. Agr. Coll. Pathologium, Corvallis, July
30, 1915; 3027:
Rubus strigosus Michx.—Stream banks, Eastern Oregon, 4,000-
5,000 ft. elev., July, 1897, W. C. Cusick, 1729.
JACKSON: UREDINALES OF OREGON 227
49. PHRAGMIDIUM IVESIAE Sydow, Ann. Myc. 1: 329. 1903.
Phragmidium affine Sydow, Ann. Myc. 2: 29. 1904.
ON ROSACEAE:
Potentilla blaschkeana Turcz.—Philomath, June 20, IgI10, 1503;
Austin, Grant Co., June, 1913, J. R. Weir, 147; Baker Co., June, 1913,
J. R. Weir,. 77; Sumpter, Baker Co., June, 1913, J. R. Weir, 3; Hil-
gard, Union Co., July 10, 1914, 1534.
Potentilla glomerata A. Nels.—Andrews, Harney Co., Aug. 1901,
Griffiths & Morris (Griffiths, West Am. Fungi 577d).
Potentilla gracilis Dougl.—Corvallis, June 20, 1909, E. R. Lake,
1499, July 29, 1914, 1477; Wren, Benton Co., June 26, 1914, 1323.
Potentilla sp.—Corvallis, June, 1910, rzz0, 3149, Aug. 1911, F. D.
Bailey, 7071.
50. PHRAGMIDIUM JONEsII Dietel, Hedwigia 44: 128. Yrg905.
On ROSACEAE:
Ivesia Baileyi S. Wats.—Steins Mts., Harney Co.,. July 27, 1898,
W. C. Cusick, Phan. Herb. 1967. (From specimen in herb. Field
Museum 108727.)
51. PHRAGMIDIUM MONTIVAGUM Arth. Torreya 9: 24. 1909.
On ROSACEAE:
Rosa gymnocarpa Nutt.—North slope Mt. Hood, Aug. 9, 1914,
1478; Bank of Minam River, Union Co., alt. 5,100 ft., Oct. 4, 1897,
E. P. Sheldon, 9053.
Rosa pisocarpa Gray?—Hilgard, Union Co., July 10, 1914, 1537.
Rosa sp.—Trail Creek Cafion, Wallowa Co., May 18, 1897, E. P.
Sheldon, 8073; Corvallis, May 1, 1914, 1466, April 25, 1915, G. B.
Posey & C. M. Schearer, 3753, April 28, 1915, 3151; North slope Mt.
Hood, Aug. 7, 1914, 1619; Mouth of Salmonberry River, Tillamook
Co., July 17, 1915, G. VanGundia, 3089.
52. PHRAGMIDIUM OCCIDENTALE Arth.; Earle, in Greene, Pl. Baker.
2° 4.) 1001.
On ROSACEAE:
Rubus parviflorus Nutt.—Wallowa Lake, Wallowa Co., Aug. 1899,
C. L. Shear, 952 (Griffiths, W. Am. Fungi 329); Jackson Co., July 9,
1903, E. B. Copeland (Sydow, Ured. 1788); Mt. Hood, Aug. 31, 1901,
E. W. D. Holway, Aug. 7, 1914, 1636; Glen Brook, Benton Co.,
Aug. 1909, 1119; Trail to Hanging Valley, Mt. Jefferson, H. P. Barss
& G. B. Posey, 1785; Mary’s Peak, Benton Co., Aug. 15, 1914, 1285;
Elk City, Lincoln Co., Aug. 20, 1914, 1626; Dothan, Douglass Co.,
Sept. 8, 1914, 1930; Corvallis, May 4, 1915, 3059; Unity, Baker Co.,
Aug. 1915, J. R. Weir, 242; Austin, Grant Co., Aug. 1916, J. R.
Weir, 238.
228 BROOKLYN BOTANIC GARDEN MEMOIRS
53. PHRAGMIDIUM POTENTILLAE (Pers.) P. Karst. Bidr. Finl. Nat.
Folk 35: 49, "1879.
Puccinia Potentillae Pers. Syn. Fung. 229. 1801.
ON ROSACEAE:
Potentilla aracnoides Lehm.—Austin, Grant Co., Aug. 1915, J. R.
Weir, r6r.
Potentilla Hippiana Lehm.—Austin, Grant Co., Aug. 1915, J. R.
Weir, 158.
54. PHRAGMIDIUM ROSAE-ACICULARIS Liro, Bidr. Finl. Nat. Folk. 65:
428. 1908.
ON ROSACEAE:
Rosa nutkana Pres|.—Bridal Veil, Multnomah Co., May 18, 1915,
3348; Edge of woods on Minam River, Union Co., Aug. 11, 1897,
E. P. Sheldon, 8667.
Rosa sp.—Corvallis, July 28, 1914, 3746.
55. PHRAGMIDIUM ROSAE-CALIFORNICAE Dietel, Hedwigia 44: 125. 1905.
ON ROSACEAE:
Rosa gymnocarpa Nutt.—Corvallis, July 29, 1914, H. P. Barss,
1469; Mary's Peak, Benton Co., Aug. 15, 19m, 1512; 1570)) 752
Ashland, Jackson Co., Sept. 10, 1914, 3084.
Rosa nutkana Presl.—Corvallis, July 29, 1914, 1473; Portland,
Aug. 23, 1915, E. Bartholomew, 5950 (Barth. N. Amer. Ured. 1626);
Hood River, Aug. 26, 1915, E. Bartholomew, 50973 (Barth. Fungi
Columb. 4834); Austin, Grant Co., Aug. 1915, J. R. Weir, 157;
Bend, Crook Co., Sept. 11, 1916, J. R. Weir, 202.
Rosa pisocarpa Gray—Corvallis, April 5, 1914, 1523.
Rosa sp.—Bonneville, Multnomah Co., Aug. II, I910, 1072,
Corvallis, Aug. 10, I91I, 31752, Spring -1914, H. C.. Gilbert, 3755,
Eugene, Lane Co., July 11, 1914, G. B. Posey, 1467; ‘Ashland, Jackson
Co., Sept. 30, 1914, 3350; Whitewater Creek, near Mt. Jefferson, Aug.
11, 1914, H. P. Barss & G. B. Posey, 3362.
56. PILEOLARIA TOXICODENDRI (Berk. & Rav.) Arth. N. Am. Flora 7;
147.- 1907:
Uromyces Toxicodendri Berk. & Rav. Grevillea 3: 56. 1874.
Pileolaria brevipes Berk. & Rav. Grevillea 3: 58. 1874.
On ANACARDIACEAE:
Rhus diversiloba T. & G.—Corvallis, April 29, 1914, F. D. Bailey,
1831; Grant’s Pass, Josephine Co., Sept. 5, 1916, J. R. Weir, 256;
Jim Creek, Wallowa Co., June 14, 1897, E. P. Sheldon, 8279.
57. PoLYTHELIS FUSCA (Pers.) Arth. Résult. Sci. Cong. Bot. Vienne
B40.) TODO:
Aecidium fuscum Pers., in Gmel. Syst. Nat. 2: 1473. 1791.
JACKSON: UREDINALES OF OREGON 229
ON RANUNCULACEAE:
Anemone oregana A. Gray—Mary’s Peak, Benton Co., May 23,
IQI5, 3030.
Anemone quinquefolia L.2—North slope Mt. Hood, Aug. 9, 1914,
go2T.
58. PUCCINIA Apsintui (Hedw. f.) DC. Fl. Fr. 6: 56.. eT8rs,
Uredo (Puccinia) Artemis Hedw. f.; DC. in Lam. Encycl. Meth.
Bot. 63-245. ° 180s.
Puccimia similis E. & E. Bull. Torrey Club 25: 508. 18908.
ON CARDUACEAE:
Artemisia dracunculoides Pursh—Sherman, Sherman Co., July 1,
1914, 2671, May 16, 1915, 2672.
Artemisia frigida Willd.—Bend, Crook Co., Sept. 11, 1916, J. R.
Weir, 212.
Artemisia ludoviciana Nutt.—Eastern Oregon, Aug. 1914, H. F.
Wilson, 3321; Eugene, Lane Co., July 20, 1914, F. D. Bailey, 1504;
Portland, Aug. 21, 1915, E. Bartholomew, 5939 (Barth. Fungi Columb.
5048); Sumpter, Baker Co., June, 1913, J. R. Weir, 9z; Grant’s Pass,
Josephine Co., Sept. 3, 1916, J. R. Weir, 249; Hood River Co., July
29° 1915, 3136.
Artemisia rigida (Nutt.) A. Gray—Eastern Oregon, 3,500 ft.
altitude, Sept. 1900, W. C. Cusick, 2504; Lost Valley, Wheeler Co.,
Sept. 9, 1894, J. B. Lieburg, 888.
Artemisia tridentata Nutt.—Redmond, Crook Co., Sept. 15, 1913,
Kirk Whited, 3787, July 2, 1914, 2555; Sherman, Sherman Co.,
July 1, 1914, 1938; Umatilla, Umatilla Co., July 11, 1914, 1293, May
mm o05, 3036, Park, Union Co:, Oct. 9; 1897, E:-P. Sheldon, o773.
This species, presumably a brachy-form, though no pycnia have
yet been observed, is not to be confused with any other species on this
host genus. The only other species recognized in North America is
P. conferta (cf. 90) which is a micro-form.
59. Puccinia abundans (Pk.) comb. nov.
Aecidium abundans Pk. Bot. Gaz. 3: 34. 1878.
Puccinia Crandall Pam. & Hume, Proc. Dav. Acad. Sci. 7: 250.
1899.
Puccinia Kreagert Ricker, Jour. Myc. 11: 114. 1905.
ON CAPRIFOLIACEAE: I.
Symphoricarpos albus (l.) Blake—Head of Applegate Creek,
Jackson Co., July 29, 1913, fs. P. Meinecke, Cr D (1) 5; Bridal Veil,
Multnomah Co., May 18, 1915, 3054, Mary’s Peak, Benton Co.,
May 21, 1915, 3036; Hilgard, Union Co., July 9, 1914, 2546; Hood
River, May 14, 1914, 2566, July 21, 1915, 3063; Springbrook, Yamhill
230 BROOKLYN BOTANIC GARDEN MEMOIRS
Co., May 14, 1914, F. D. Bailey, 2567; Philomath, April 26, 1914,
2572; Corvallis, April 28, 1915, 2612; Grant’s Pass, Josephine Co.,
Sept. 3; 1916,.J) Ro Weir, 767;
On Poaceae: II and III.
Festuca confinis Vasey (Poa Kingi S. Wats.)—Steins Mts., Harney
Co., July 2, 1896, J. B. Leiberg, 2945.
Festuca idahoensis Elmer—Hilgard, Union Co., July 10, I9g14,
1358, 1362; Redmond, Crook Co., July 2, 1914, 1424, 1430.
Festuca rubra L.—Hilgard, Union Co., July 10, 1914, 1366; Mary’s
Peak, Benton Co., Aug. 15, 1914, 1571, 1573; Newport, Lincoln Co.,
July 18, 1915, 3207:
Festuca subulata Trin.—Ashland, Jackson Co., Sept. 10, 1914, 1563.
The connection between this common western form on Festuca
with Aecidium abundans was shown by Arthur in 1910 (Mycologia 4:
27. 1912). In three trials, using telial material on F. confinis, col-
lected in Colorado and Utah, infection resulting in pycnia and aecia
on Symphoricarpos racemosus was obtained.
60. PuccInIA ACETOSAE (Schum.) Koern. Hedwigia 15: 184. 1876.
Uredo Acetosae Schum. Enum. PI. Saell. 2: 231. 1803.
ON POLYGONACEAE:
Rumex acetosella L.—Maples Station, Tillamook Co., Sept. 15,
1915, F. D. Bailey, 3702; Corvallis, Oct. 19, 1915, G. B. Posey, 3000.
This species has been recorded previously from North America
only from Florida on R. hastatulus (Holway, North Am. Ured. 1: 35.
1906). Specimens on that host referred to this species are in the
Arthur herbarium also from S. Carolina and Massachusetts and on
R. acetosella from Massachusetts, Florida, New York and Indiana.
All the specimens bear uredinia only. It is possible that some or all
of the material should be referred to Uromyces Acetosae Schroet., as
the two species are indistinguishable in the uredinial stage.
61. PUCCINIA AMBIGUA (Alb. & Schw.) Lagerh., in Bubak, Sitz. Ver.
Bohm. Ges. Wiss. 1898, 28: 14. 1808.
Aecidium Gali ambiguum Alb. & Schw. Consp. Fung. 116. 1805.
Puccinia difformis Kunze, Myc. Hefte 1: 71. 1817.
Allodus ambigua Arth. Résult Sci. Congr. Bot. Vienne 345. 1906.
ON RUBIACEAE:
Galium aparine L.—Wren, Benton Co., June 26, I914, 1330;
Ashland, Jackson Co., Sept. 10, 1914, 30906.
This species possesses aecia and telia only in the life cycle. It
has been studied by Bubak (1. c.) who found that primary aecia were
followed by secondary aecia. Later Trebaux (Flora 81: 394-404.
1895) repeated this observation and conducted culture work con-
JACKSON: UREDINALES OF OREGON 231
firming Bubak’s contention. This species should not be confused
with P. punctata Lk. (cf. 156) which occurs on the same host from
this region.
62. PucctinIA ANGELICAE (Schum.) Fckl. Symb. Myc. 52. 1869.
(Not P. Angelicae E.& E. 1884.)
Uredo Angelicae Schum. Enum. PI. Saell. 2: 233. 1803.
Puccinia Archangelicae Blytt, Christiania Vid. Selsk. Forhandl.
No. 6:51. 1896.
Bullaria Angelicae Arth. Résult Sci. Congr. Bot. Vienne 346.
1906.
ON UMBELLIFERAE:
Angelica genuflexca Nutt.—Woodburn, Clackamas Co., Sept. 1885,
Thomas Howell.
Angelica Lyallit Wats.?—Larch Mt., Multnomah Co., Aug. 10,
1910, 2613.
This species is evidently rare in North America having been
reported otherwise only from a single collection from Washington on
A. genuflexa and one from New York on A. atropurpurea. It is a
brachy-form though pycnia have not been seen in American collec-
tions. This species has smooth teliospores and is easily separable
from Puccinia Ellisii (cf. 98) on the same hosts from our region, which
has verrucose spores.
63. PUCCINIA ANOMALA Rost. Thiimen, Flora 1877: 92. 1877.
Puccinia straminis simplex Koern. Land. u. Forstw. Zeit. no. 50.
1865. —
Puccinia Hordes Otth. Mitt. Nat. Ges. Bern. 1870: 114. 1871.
(Not P. Hordet Fckl. 1860.)
Puccinia simplex Erikss. & Henn. Getreideroste 238. 1896. (Not
P. simplex Peck. 1881.)
Aecidium Ornithogalum Bubak, Ann. Myc. 3: 223. 1905.
On Poaceae: II, III.
Hordeum montanense Schribn.—Corvallis, July 26, 1914, 1474.
Hordeum murinum L.—Corvallis, July 8, 1914, G. B. Posey, 1354.
Hordeum nodosum L.—Corvallis, July 26, 1914, 3257.
Hordeum vulgare L.—Corvallis, July 6, 1914, 7683, Aug. 13, 1914,
1691, 1708.
This, the leaf rust of barley, is evidently very common in Oregon,
much more so than the collections listed above would indicate. It is
evidently not abundant in America except on the Pacific coast. In
the Arthur herbarium, specimens on wild barleys are represented only
from Oregon, California and Utah. On the cultivated barley speci-
mens are at hand only from California, Iowa and Wisconsin. It is
232 BROOKLYN BOTANIC GARDEN MEMOIRS
evidently spreading into the eastern United States as the writer col-
lected it in August, 1916, at Ithaca and Savanna, New York.
Tranzschel has shown that this rust in Russia has its aecia on
Ornithogalum umbellatum and O. narbonense (Mycol. Cent. 4: 70.
I9I4).
64. PuCCINIA ANTIRRHINI Diet. & Holw: Hedwigia 36: 298. 1897.
ON SCROPHULARIACEAE:
Antirrhinum majus L.—Portland, Aug. 1909, comm. Charles
Ladd, roSo0, Aug. 28, 1914, comm. P. C. Schmeir, 1974; Salem,
Marion Co., July, 1911, comm. Mrs. Lord, 1727; Corvallis, June 26,
1912, 1085, Aug. 1912, 1025.
The snapdragon rust is very common in Oregon both in gardens
and in the greenhouse. For a long time it was known to occur only
in California. As snapdragons came to be used more commonly in
greenhouse culture the rust has gradually spread through the distri-
bution of cuttings, till at the present time it is known to occur in most
of the central and eastern states.
65. PUCCINIA ARNICALIS Pk. Bot. Gaz. 6: 227. 1881.
On CARDUACEAE:
Arnica cordifolia Hook.—Near Aneroid Lake, July 1, 1899, II,
BR. Lake; 7407.
A very distinct species having minutely verrucose teliospores,
not thickened at the apex, and is known only from the Rocky moun-
tain and Pacific coast regions.
66. PUCCINIA ASARINA Kunze, in Kunze & Schmidt, Myk. 1: 7o.
1817.
Puccinia Asari Link in Willd. Sp. Pl. 67: 68. 1825.
Dicaeoma asarinum Kuntze, Rev. Gen. Pl. 3: 467. 1808.
ON ARISTOLOCHIACEAE:
Asarum caudatum Lind|.—Portland, Aug. 30, 1915, E. Bartholo-
mew, 50977 (Barth. Fungi Columb. 4840).
This micro-form is known from North America on the above host,
otherwise only from California, Idaho and Washington.
67. PUCCINIA ASPERIFOLII (Pers.) Wettst. Verh. Zool.-Bot. Ges. Wien
35: 541. 1885.
Aecidium asperifolii Pers. Obs. Myc. 1: 97. 1796.
Puccinia dispersa Erikss. Zeitsch. f. Pflanzenkr. 4: 257. 1894.
ON POACEAE:
Secale cereale L.—Hood River, June 19, 1914, 1402; Corvallis,
July 28, 1914, 2682; Bend, Crook Co., Sept. 11, 1916, J. R. Weir, 243.
The leaf rust of rye is evidently common throughout the state.
JACKSON: UREDINALES OF OREGON 255
This species has its aecia on species of Anchusa and Lycopsis in Europe
‘as was first shown by De Bary (Monatsber. k. Akad. d. Wiss. Berlin
211. 1866). No aecia referable to this species have been found in
America, but Arthur (Mycologia 1: 236. 1909) obtained the develop-
ment of pycnia on Lycopsis arvensis secured from Europe, following
exposure to germinating telia on rye collected in Indiana. This
culture indicates that the European and American rusts are identical.
68. PUCCINIA ASPERIOR E. & E. Bull. Washb. Lab. 1: 3. 1884.
Puccinia oregonensis Earle, Bull. N. Y. Bot. Gard. 2: 349. I902.
Allodus oregonensis Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
Allodus asperior Orton, Mem. N. Y. Bot. Gard. 6: 193. Ig16.
ON UMBELLIFERAE:
Leptotaenia dissecta Nutt.—Corvallis, June and July, 1898, M.
Craig, April 14, 1899, M. Craig, June, 1910, 2674, March 24, 1914, G. B.
Posey,2665; Mary’s River near Wren, Benton Co., June 5, 1915, 2673.
This is one of the most common and conspicuous of rusts, prob-
ably widely distributed throughout western Oregon. The type of
P. oregonensts, the second collection listed, was described as on Sanicula
bipinnata, which is clearly an error for the above host as was first
pointed out by Holway (N. Am. Ured. 14: 93. 1913).
69. PucciniA ASTERUM (Schw.) Kern, Mycologia g: 224. I917.
Aecidium asterum Schw. Schrift. Nat. Ges. Leipzig 1: 67. 1822.
Aecidium Solidaginis Schw. Schrift. Nat. Ges. Leipzig 1: 68.
1822.
Caeoma asteratum Link in Willd. Sp. Pl. 6: 51. 1825.
Caeoma (Aecidium) erigeronatum Schw. Trans. Am. Phil. Soc. II.
a 2O2. 1632.
Puccinia extensicola Plowr. Brit. Ured. Ustil. 181. 1889.
Puccinia Caricis-Erigerontis Arth. Jour. Myc. 8: 53. 1902.
Puccinia Caricts-A steris Arth. Jour. Myc. 8: 54. 1902.
Puccinia Caricis-Solidaginis Arth. Bot. Gaz. 35: 21. 1903.
ON CARDUACEAE: I.
Aster sp.—Philomath, May 10, 1914, 7309, 3066; Hilgard, Union
Co., July 10, 1914, 3056; Corvallis, May 9, 1914, 3367, 3368, April 31,
1915, W. E. Lawrence, 3048; Hood River, May 14, 1914, 3022;
Sumpter, Baker Co., June 1913, J. R. Weir, 85.
Erigeron speciosus DC.—Near Whitewater ranger station, Mt.
Jefferson, Aug. 16, 1914, H. P. Barss & G. B. Posey, 3292.
Euthamia occidentalis Nutt.—Mary’s River, Benton Co., June
1898, M. Craig.
234 BROOKLYN BOTANIC GARDEN MEMOIRS
On CypERACEAE: II and III.
Carex athrostachya Olney—Philomath, May Io, 1914, 3286, Cor-
vallis, June 29, 1914, G. B. Posey, 1333.
Carex canescens L.—Hood River, Aug. 5, 1914, 3005.
Carex Deweyana Schw.—Glendale, Douglass Co., Aug. 17, 1914,
1409; Elk City, Lincoln Co., Aug. 20, 1914, 1382, 1383; Philomath,
May 10, 1914, 3284; Trail to Sulphur Springs, Benton Co., Nov. 3,
1912, 3288; Corvallis, Apr. 29, 1914, F. D. Bailey, 3283, May 19, 1913,
TOR".
Carex festiva Dewey—Newport, Lincoln Co., July 18, 1915, 3279.
Carex Goodenowi J. Gay (C. vulgaris E. Fr.)—Hood River, June
20, 1914, 1405.
Carex phyllomanica W. Boot ?—Grant’s Pass, Josephine Co., Sept.
2, 1916, |x ie Wel, 220:
Carex praegracilis Boot (C. marcida Boot)—Corvallis, May 9,
1914, 3287.
Carex scoparia Schk.—Corvallis, June 24, 1914, F. D. Bailey,
1386.
Carex stipata Muhl.—Sherwood, Washington Co., July 10, 1914,
F. D. Bailey, 7355; Hood River, May 14, 1914, 30142 Hulbbara:
Clackamas Co., May 27, 1914, 3011; Portland, May 19, 1914, F. D.
Bailey, 3003; Corvallis, Aug. 10, 1910, 1189, May 9, 1914, 3285,
July 29, 1915, 3281; Eddyville, Lincoln Co., Aug. 8, 1915, Hoerner,
3350.
Carex straminea Willd.—Hood River, July 24, 1915, 3280.
Carex subfusca W. Boot.—Corvallis, July 29, 1914, 1444.
Carex sp.—Ashland, Jackson Co., Sept. 10, 1914, 3008; Corvallis,
July 29, 1914, 1442; Cottage Grove, Lane Co., July 14, 1914, 1350;
Philomath, Jan. 6, 1914, z78o.
In 1901 Arthur (Jour. Myc. 8: 54. 1902) first began culture
work showing that aecia which occur commonly on Aster, Solidago and
related hosts are genetically connected with uredinia and telia on
various species of Carex. The culture work conducted by Arthur is
extensive and extends over a period of years from 1901-1914. In this
series of culture work aecia have been produced on various species of
Aster, Solidago, Erigeron, Leptilon and Euthamia, using telia from
Dulichium and from many species of Carex from various parts of
North America (Jour. Myc. 8:54. 1902; 11: 58. 1905; I2seme
1906; 14: 13. 1908; Bot. Gaz! 35:15, 21. - 1903; Mycol.-mitaee
190Q;'2:224. 1910; 4:15, 16:' IOT2; 7: 70; ST: ~ TOLS).- eeeoueee
1911 (Mycol. 4: 181. 1912) confirms Arthur’s results in part by
successfully infecting Aster acuminatus using telial material from
Carex trisperma L.
JACKSON: UREDINALES OF OREGON 235
This study has also shown that the species as here considered
is a composite form made up of several distinct physiological races.
The species is separable from all other American species of Puccinia
on Carex by the presence of two pores in the upper part of the rather
small (12-19 by 16—-23 uw) urediniospores and the medium-sized (12-20
by 35-50 yp) teliospores.
' The aecia of this species should not be confused with those of
P. stipae (cf. 166), which occur on the same generic hosts and other
Carduaceae in the west. In P. Asterum the aecia are cupulate, the
peridium conspicuous, the spores globoid, nearly colorless and small
(11-15 by 13-18 w). In P. stipae the aecia occur on hypertrophied
areas, the individual cups are gall-like and open by a central aperture,
the peridium is evanescent, the spores are cinnamon brown, globoid,
large (21-26 by 22-29 uw). The aeciaof the latter species have not been
found in Oregon, but doubtless occur abundantly in the eastern part
of the state.
70. PUCCINIA ASTERIS Duby, Bot. Gall. 2: 888. 1830.
On CARDUACEAE:
Aster conspicuus Lind|.—Hilgard, Union Co., July 10, 1914, 1538;
Crater Lake, Klamath Co., Sept. 3, 1916, J. R. Weir, 182.
Aster sp.—Corvallis, Aug. 10, 1911, F. D. Bailey, 7775, May 1,
1915, 3050; Austin, Grant Co., June 1913, J. R. Weir, zo2.
71. PUCCINIA ATRO-FUSCA (Dudley & Thompson) Holway, Jour. Myc.
FOs1228.. O04.
Uromyces atro-fuscus Dudley & Thompson, Jour. Myc. 10: 55.°
1904.
ON CYPERACEAE:
Carex Douglasit Boot.—Enterprise, Wallowa Co., July 24, 1897,
E. P. Sheldon (from Phan. spec. 8634).
This species possesses amphispores which were first mistaken for
the teliospores of a Uromyces. It may be distinguished from other
Carex rusts by the presence of the amphispores together with the
normal urediniospores, the latter are 20-26 w long and thin walled,
1.5-2 u thick. The aecial connection is unknown. It is known only
from the Rocky mountain and Pacific coast regions.
72. PuccINIA BALSAMORRHIZAE Pk. Bull. Torrey Club 11: 49. 1884.
Trichobasis Balsamorrhizae Pk. Bot. Gaz. 6: 276. 1881.
On. CARDUACEAE:
_ Balsamorrhiza deltoidea Nutt.—Corvallis, July 29, 1914, 1472;
Hermiston, Umatilla Co., May 12, 1915, 2663.
Balsamorrhiza sagittata (Pursh) Nutt.—Hood River, Aug. 11, 1909,
3186; Durfur, Wasco Co., June 19, 1914, 1836.
236 BROOKLYN BOTANIC GARDEN MEMOIRS
73. PUCCINIA BICOLOR Ell. & Fv. Bull. Torrey Club 27: 572. 1900.
ON CICHORIACEAE:
HMieracium cinereum Howell—Hood River, July 22, 1915, 3325.
Hieracium Scoulert Hook.—White Pine, Baker Co., June 1913,
Jick WWem, a7
Hieracium sp.—Durfur, Wasco Co., June 30, 1914, 1338.
This very distinct micro-form is known otherwise only from the
type collection made at Waitsburg, Wash., May 7, 1900, on H. Scouleri,
by R. M. Horner and distributed in E. & E. Fungi Col. 1570.
74. PUCCINIA BISTORTAE (Strauss) DC. Fl. Fr. 6: 61. 1815.
Uredo Polygoni Bistortae Strauss, Ann. Wett. Ges. 2: 103. 1870.
ON POLYGONACEAE:
Polygonum imbricatum Nutt.—Oregon?, Aug. 1880, Thos. Howell.
Polygonum Newberryt Small—Crater Lake, Klamath Co., Sept. 22,
1913, E. P. Meinecke, Cr Pk D (2) 14; N. slope Mt. Jefferson, 2,600
ft., Marion Co., Aug. 16, 1914, H. P. Barss & G.. B.. Posey aazea,
Aug. 27, 1916, He JP. Barss, 3305.
This species may be distinguished from other North American
Polygonumrusts by the medium-sized teliospores (16-21 by 26-35 pu)
with wall of uniform thickness, without hyaline umbo. It is not
known elsewhere in North America on the first mentioned host and
otherwise only from Washington on P. Newberryi. Aecia are un-
known.
75. PucCINIA BLASDALEI Diet. & Holw. Erythea 1: 248. 1893.
On ALLIACEAE:
Allium attenuifolium Kellog—Corvallis, June 2, 1915, C. E. Owens,
268T. .
Allium acuminatum Hook.—Hood River Co., June 10, 1917, Leroy
Childs. ;
This rust may be distinguished from other Allium rusts by the
strongly developed stroma in the telial sori, and the tendency to form
confluent telia covering large areas on the stems and leaves. The
teliospores are large (16-26 by 40-61 yw), thickened to 4-10 uw at the
apex.
76. PUCCINIA CALOCHORTI Pk. Bot. Gaz. 6: 228. 1881.
Allodus Calochorti Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
On LILIACEAE:
Calochortus macrocarpus Dougl.—Redmond, Crook Co., July 21,
1912, Kirk Whited, 3782; Hills near Malheur River, Harney Co.,
June 6, 1901, W. C. Cusick, 2544; Powder River Mts., Baker Co.,
Aug. 1896, C. V. Piper, 2460.
JACKSON: UREDINALES OF OREGON 237
All of the above specimens were secured from phanerogamic
specimens, the first from the herbarium of the Oregon Agr. College,
the others from the herbarium of the N. Y. Botanical Garden. The
species is an opsis-form.
77. PUCCINIA CAMPANULAE Carm. Smith’s English Flora 5: 365.
1826.
Puccinia Campanulae Fckl. Sym. Myc. 53. 1869.
On CAMPANULACEAE:
Campanula Scoulert Hook.—Mary’s Peak, Benton Co., Aug. 15,
1914, 2559; Hood River, July 24, 1914, 3023.
A comparison of the above collections with European material
shows that the rust is identical and should be referred as above. This
is a micro-form unrecorded in America so far as the writer is aware,
and known otherwise from North America only from collections made
by the writer and others, on C. rotundifolia at Fall Creek, Ithaca, New
York.
78. PucciniA CHELONIsS Diet. & Holw. Hedwigia 36: 297. 1897.
ON SCROPHULARIACEAE:
Chelone nemorosa Dougl.—Mt. Hood, Sept. 1, 1901, E. W. D.
Holway.
A micro-form known otherwise only from Washington on the same
host.
79. PUCCINIA CHRYSANTHEMI Roze. Bull. Soc. Myc. Fr. 17:92. 1900.
On CARDUACEAE:
Chrysanthemum sinense Sabine—Portland, Nov. 1914, W. H.
Dunham, 1986.
The above collection from a greenhouse is the only collection we
have seen from Oregon. It is doubtless not infrequent in greenhouses
throughout the state. The life history is unknown. This rust is
evidently a native of Japan, having been introduced into America
and Europe where it has become widespread on cultivated chrys-
anthemums.
80. PucciniA CicHori (DC.) Bell, in Kickx. Fl. Fland. 2:65. 1867.
Uredo Cichoru DC. Fl. Fr. 6: 74. 1815.
ON CICHORIACEAE:
Cichorium intybus L.—Corvallis, Sept. 21, 1914, G. B. Posey, 1931.
81. Puccinia CicuTAE Lasch, Klotsch. Herb. viv. myc. No. 787.
1845.
Puccinia Cicutae Thiim. Bull. Soc. Imp. des Nat. Moscow 52: 136.
1877.
238 BROOKLYN BOTANIC GARDEN MEMOIRS
On UMBELLIFERAE:
Cicuta occidentalis Greene?—Klamath Falls, Klamath Co., Sept. 8,
1916, J. R. Weir, 223.
Cicuta sp.—Eastern Oregon, June, 1885, T. Howell.
The last-mentioned specimen is from the herbarium of W. G.
Farlow. It is marked on Peucedanum. ‘The host is clearly Cicuta sp.
82. PuCCINIA CIRCAEAE Pers. Roemer’s Neues Mag. 1: 119. 1794.
ON ONAGRACEAE:
Circaea pacifica Asch. & Magn.—West of Noon station, Benton
Co., Aug. 8, 1914, H. P. Barss, 7296; Hood River, July: 24, tens,
3062; Near Mary’s Peak, Benton Co., Aug. 15, 1914, 3263; Sumpter,
Baker Co., July 16, 1913, J. R. Weir, r99.
83. Puccinta Cirsii Lasch, in Rabh. Fungi Eur. No. 89. 1859.
Puccinia tnclusa Syd. Monog. Ured. I: 56. 1902.
ON CARDUACEAE:
Cirsium americanum (Gray) Robinson—Wren, Benton Co., June
26, 1914, 1332; Corvallis, May 20, 1915, 3242.
Cirsium edule Nutt.—Elk City, Lincoln Co., Aug. 20, 1914, 2520.
Cirsium undulatum (Nutt.) Spreng.—Sherman, Sherman Co.,
July 1, 1914, 1906.
84. PUCCINIA CLAYTONIATA (Schw.) Pk. Bull. N. Y. State Mus. 6:
226. 1899.
Caeoma (Aecidium) claytoniata Schw. Trans. Am. Phil. Soc. II.
4: 204. 1832.
Puccinia Mariae-Wilsoni G. W. Clinton; Peck, Bull. Buff. Soc.
INdty ci: 22 66. 1873:
Allodus claytoniata Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
ON PORTULACACEAE:
Claytonia lanceolata Pursh?—Austin, Grant Co., May, 1916, J. R.
Weir, 206.
85. PucctiniA CLEMATIDIS (DC.) Lagerh. Tromsé Mus. Aarsh. 17:
47. 1895.
Aecidium Clematidis DC. FI. Fr. 2: 243. 1805.
Aecidium Aquilegiae Pers. Icon. pict. IV. 58. 1806.
Puccinia tomipara Trel. Trans. Wis. Acad. Sci. 6: 127. 1885.
Puccinia Agropyri E. & E. Jour. Myc. 7: 131. 1892.
Puccinia cinerea Arth. Bull. Torrey Club 34: 583. 1907.
Puccinia alternans Arth. Mycol. 1: 248. 1909.
Puccinia obliterata Arth. Mycol. 1: 250. 1909.
JACKSON: UREDINALES OF OREGON 239
ON RANUNCULACEAE: I.
Aquilegia formosa Fish.—Myrtle Creek, Douglass Co., June 9,
1914, F. D. Bailey, 2573; Hood River, May 14, 1914, 2565, May 9,
1915, 3040; Bridal Veil, Multnomah Co., May 18, 1915, 3051.
Aquilegia truncata Fisch. & Mey.—Pokegama, Jackson Co., July
9, 1903, E. B. Copeland (Sydow, Ured. 1767), E. B. Copeland, 3717
(Rocky Mt. Herb. 45896). .
Clematis Drummondiu T. & G.—Freewater, Umatilla Co., July
10, 1914, 2562.
Clematis hirsutissima Pursh (C. Douglasit Hook.)—Austin, Grant
Go; July, 1913, J. R. Weir, 783.
Clematis ligusticifolia Nutt.—Corvallis, Linn Co., Sept. 2, 1914,
F. D. Bailey, 2563, Benton Co., May 4, 1915, 3307.
Thalictrum occidentale A. Gray—Corvallis, May 4, 1912, 1147,
July 4, 1914, G. B. Posey, 3067, May 4, 1915, 3270; Wren, Benton
Co., June 26, 1914, 7331a; White Pine, July, 1913, J. R. Weir, 153;
Austin, Grant Co., Aug. 1915, J. R. Weir, 205.
On Poaceae: II, III.
Agropyron dasystachyum (Hook.) Vasey—Redmond, Crook Co.,
July 2, 1914, 1432.
Agropyron lanceolatum Scribn. & Sm.—Redmond, Crook Co.,
lye2, 1914, 1427.
Agropyron spicatum (Pursh) Rydb.—Wren, Benton Co., June 26,
1914, 1320.
Bromus carinatus Hook. & Arn.—Newberg, Yamhill Co., June 8,
1913, F. D. Bailey, 7197; Portland, July 10, 1905, J. J. Davis, Aug.
23, 1915, E. Bartholomew, (Barth. Fungi Columb. 4846); Corvallis,
Sept. 10, 1914, 71577; Philomath, May Io, 1914, 3193; Hood River,
May 14, 1914, 1587, 1588, 1593; Grant’s Pass, Josephine Co., Sept. 3,
1916, J. R. Weir, 207.
Bromus carinatus californicus Shear—Philomath, Jan. 6, 1914,
1148.
Bromus grandis (Shear) Hitchc.—Corvallis, June 4, 1914, 1384.
Bromus hordeaceus L.—Portland, May 21, 1914, 1582.
Bromus hordeaceus leptostachys Beck.—Hood River, May 20, 1914,
1585; Springbrook, Yamhill Co., May 14, 1914, F. D. Bailey, 1504,
1595.
Bromus marginatus Nees.—Hood River, Aug. 6, 1914, 1550, July
27, 1915, 3192; Corvallis, June 24, 1914, G. B. Posey, 1389, June 29,
1914, G. H. Godfrey, 1312; Rose City Park, Portland, Jan. 9, 1914,
1198; Redmond, Crook Co., July 2, 1914, 1422; Hilgard, Union Co.,
July 10, 1914, 7365.
Bromus secalinus L.—Cottage Grove, Lane Co., July 14, 1914,
1352.
Lz
240 BROOKLYN BOTANIC GARDEN MEMOIRS
Bromus tector'um L.—Clatskanie, Columbia Co., May 20, 1914,
Ff; De-Bailey, 1567;
Bromus villosus Forsk.—Myrtle Creek, Douglass Co., June 9,
1914, F. D. Bailey, 24006. |
Bromus vulgaris Shear—Ashland, Jackson Co., Sept. 10, I914,
1569; Corvallis, Feb. 14, 1914, 3261, July 4, 1914, G. B. Posey, 1478,
July 29, 1914, 1443; trail to Sulphur Springs, Benton Co., Nov. 3,
1914, 3195; Mary’s Peak, Benton Co., Sept. 15, 1914, 1574.
Elymus condensatus Presl.—Albany, Linn Co., Aug. 1907, D.
Griffiths.
Elymus glaucus Buckl.—Wren, Benton Co., June 26, 1914, 7321,
1331; Ashland, Jackson Co., Sept. 10, 1914, 1562, 1564; N.slope Mt.
Hood, Aug. 7, 1914, 1556; Mary’s Peak, Benton Co., Aug. 15, 1914,
1575; The Dalles, Wasco Co., July 1, 1914, 7341; Garden Home, |
Multnomah Co., July 20, 1915, 3202; Hood River, June 20, 1914,
1403, Aug. 5, 1914, 3204; Corvallis, Feb. 14, 1914, 3262; June 29,
1914, G. B. Posey, 1304, G. H. Godfrey, 1305, July 29, 1914, 1439,
1440.
Elymus triticoides Buckl.—Columbia River, near mouth of De-
schuttes River, Sherman Co., July 29, 1914, M. E. Peck.
Poa ampla Merrill—Hood River, July 22, 1915, 3250.
Puccinella Nuttalliana (Schult.) Hitche.—Grand Ronde Valley,
Union Co., July, 1914.
Sitanion californicum J. G. Smith—N. slope Mt. Jefferson, Aug.
12, 1or4, dP: Barss; 7500:
Sitanion glabrum J. G. Smith—Umatilla, Umatilla Co., July 11,
1914, 1370.
Sitanion jubatum J. G. Smith—Redmond, Crook Co., July 2, 1914,
1428. .
Sitanion velutinum Piper—Hood River, July 22, 1915, 3255;
Hermiston, Umatilla Co., May 12, 1915, 3179.
This common subepidermal species, as here considered, includes
nearly if not all the forms having aecia on Ranunculaceous hosts.
Dietel (Oesterr. bot. Zeitschr. 42: 261. 1892) was apparently
the first to culture this species. Klebahn (Die Wirtsw. Rostpilze 292.
1904) has presented a summary of Dietel’s work together with that of
other European investigators.
In America, Arthur has conducted extensive culture work beginning
in 1904, using telial material from various parts of the country, on five
different genera of grasses representing ten species, and has success-
fully cultured them on five genera of Ranunculaceae. His work indi-
cates the presence of a number of well-marked races. (Jour. Myc. 11:
62. 1905,13:197. 1907,14:15. 1908; Mycologia 1: 246, 248, 249.
1909, 2: 225. I9Q10, 4:54. O12, 73°73) 62... 1015, 6 sen eons
JACKSON: UREDINALES OF OREGON 241
This species may be distinguished from other grass rusts having
the telia long covered by the epidermis primarily on the urediniospore
characters. They are not accompanied by paraphyses, the wall is
moderately thin, 1.5—2 yu, pale yellow to cinnamon brown, and the pores
are scattered. The telia may or may not be surrounded by stromal
hyphae and are rather narrow, 13-24 wu.
86. Puccin1ia CLINTONII Peck, Rept. N. Y. State Mus. 28:61. 1876.
ON SCROPHULARIACEAE:
Pedicularis bracteosa Benth.2—N. slope Mt. Jefferson, Aug. 16,
1914, H. P. Barss & G. B. Posey, 2545.
Pedicularis racemosa Dougl.—N. slope Mt. Jefferson, 7,000 ft.,
ue. 13, 1914, H. P. Barss, 2544.
87. Pucctnta Cnict Mart. Fl. Mosq. 226. 1817.
Puccinia Cirsit-lanceolati Schroet., Cohn, Krypt. Fl. Schl. 3}:
Sh. USS?
On CARDUACEAE:
Cirsium lanceolatum (L.) Scop.—Corvallis, Oct. 21, 1911, F. D.
Bailey, 1963, Mar. 6, 1914, G. H. Godfrey & F. D. Bailey, 1965; Elk
City, Lincoln Co., Aug. 20, 1914, 1964; The Dalles, Wasco Co.,
July 1, 1914, 7334; Portland, Aug. 21, 1915, E. Bartholomew (Barth.
Fungi Columb. 5053).
88. PUCCINIA COMANDRAE Pk. Bull. Torrey Club 11: 49. 1884.
ON SANTALACEAE:
Comandra umbellata (L.) Nutt.—Dufur, Wasco Co., July 30, 1914,
2504; Hood River, May 18, 1915, 2660, July 22, 1915, 3141.
This micro-form, found commonly in the Rocky Mt. and Pacific
coast states, possesses teliospores having similar morphological char-
acters to those of the heteroecious rust P. pustulata (Curt.) Arth.,
which has aecia on Comandra and uredinia and telia on Andropogon.
A number of such correlations between micro-forms and the telia of
heteroecious forms whose aecia occur on the same host have been
pointed out by Travelbee (Proc. Ind. Acad. Sci. 1914: 233. 1915)
among species occurring in North America. Dietel (in Engler &
Prantl, Die Nat. Pflanzenf. 1!**: 69. 1897) was apparently the first
to point out this sort of correlation between P. mesneriana Thiim.
and P. coronata (P. Rhamnit).
89. PUCCINIA COMMUTATA Sydow, Monog. Ured. 1: 201. 1902.
Allodus commutata Arth. Résult. Sci. Congr. Bot. Vienne 345. 1906.
ON VALERIANACEAE:
Valeriana occidentalis Heller—Hilgard, Union Co., July 10, 1914,
1541.
242 BROOKLYN BOTANIC GARDEN MEMOIRS
go. PUCCINIA CONFERTA Diet. & Holw. Erythea 1: 250. 1893.
Puccinia recondita Diet. & Holw. Erythea 2: 128. 1894.
ON CARDUACEAE:
Artemisia ludoviciana Nutt.—Corvallis, Sept. 2, 1914, F. D. Bailey,
2h 32; Sept: 4, 19842500:
A micro-form in which the teliospores resemble quite closely those
of P. Absinthit (cf. 58), a brachy-form also common in the west. It
is probable that this should be considered a correlated form.
g1. PuccInIA CONVOLVULI (Pers.) Cast. Obs. 1: 16. 1842.
Uredo Betae Convolvuli Pers. Syn. Fung. 221. 1801.
ON CONVOLVULACEAE:
Convolvulus atriplicifolius (Hallier f.) House—Central Point, Jack-
son Co., Oct. 6, 1914, M. P. Henderson, 7949; Grant’s Pass, Josephine
Go., Sept..3; ro16, J. Rk, Weir, 227.
92. PUCCINIA CREPIDIS-ACUMINATAE Sydow, Oestr. Zeitschr. 51: 27.
19Ol.
On CICHORIACEAE:
Crepis gracilis (D. C. Eaton) Rydberg—Baker City, Baker Co.,
July,-1913;.) Re Weir 57.
93. PucciIntia CyAnt (Schleich.) Pass. Rabh. Fungi Eur. No. 1767.
1874.
Uredo Cyani Schleich. Pl. Helv. 95.
ON CARDUACEAE:
Centaurea Cyanus L.—Corvallis, June, 1913, 7745, April 8, 1914,
2551, July 29, 1914, 2552, July 4, 1914, G. B. Posey, 3106; Orenco,
Washington Co., April 2, 1915, 3060.
94. Puccini DEBARYANA Thiim. Flora 58: 364. 1875.
Puccinia compacta DeBary, Bot. Zeit. 16: 83. 1858. (Not P.
compacta Berk. 1855.)
On RANUNCULACEAE:
Anemone Drummondi Wats.—Mt. Hood, 7,000 ft., foot of Eliott
Glacier, Sept. 1, 1901, E. W. D. Holway.
g5. PuccINIA DENTARIAE (Alb. & Schw.) Fuckel, Symb. Mycol.
Nachtihur: 7. (87a,
Uredo Dentariae Alb. & Schw. Consp. Fung. 129. 1805.
ON CRUCIFERAE:
Dentaria tenella Pursh—Corvallis, April 5, 1914, 1288.
A micro-form occurring on the petioles and blades of the basal
leaves causing considerable distortion. So far as the writer is aware
this species is known from North America only from the above col-
lection.
JACKSON: UREDINALES OF OREGON 243
96. PuUCCINIA DICHELOSTEMMAE D. & H. Erythea 3: 78. 1895.
Allodus Dichelostemmae Orton, Mem. N. Y. Bot. Gard. 6: 183.
1916.
ON ALLIACEAE:
Hookera pulchella Salisb. (Brodiaea congesta Smith)—Dallas, Polk
Co., March 20, 1900, W. N. Suksdorf (Barth. N. Am. Ured. 1541);
Corvallis, April 28, 1915, 2611, May 1, 1915, 2069; E. of Wren Station,
Benton Co., April 17, 1915, 2678.
The first-mentioned collection bears aecia and a few telia. It
is probable that they belong together. The last specimen mentioned
consists of aecia only. The others bear telia only. The two stages
rarely occur together. This species may be separated from all other
species of Puccinia occurring on Alliaceae by the very large, broad
teliospores (38-45 by 43-58 uw) having smooth walls 5-7 p thick.
97. Puccinta Douctasi Ell. & Ev. Proc. Phil. Acad. 1893: 152.
1893.
Puccinia Richardsontui Sydow, Monog. Ured. 1: 317. 1902.
Allodus Douglas Orton, Mem. N. Y. Bot. Gard. 6: 198. 1916.
ON POLEMONIACEAE:
Phlox condensata (A. Gray) E. Nels.—N. slope Mt. Hood, 7,000 ft.,
Aue..7, 1914, III, 74094, I, 2624.
Phlox diffusa Hook.—N. slope Mt. Hood, 7,000 ft., Aug. 7, 1914,
1602, 1603; Sept. 1, 1901, E. W. D. Holway.
98. PucciniA ELiist DeToni, in Sacc. Syll. Fung. 7: 651. 1888.
Puccinia Angelicae E. & E. Bull. Wash. Lab. 1: 3. 1884. (Not
P. Angelicae Fckl. 1869.)
Puccinia Bakeriana Arth. Bull. Torrey Club 31: 3. 1904.
ON UMBELLIFERAE:
Angelica genufleca Nutt.—Corvallis, Sept. 7, 1901, E. R. Lake,
1490.
Otherwise known only from Idaho and Washington on the above
host, and from California on A. tomentosa. It is doubtless a brachy-
form, though no pycnia have been found. The teliospores are closely
and finely verrucose, a character which enables one to distinguish this
species easily from P. Angelicae (cf. 62).
99. PuCCINIA EPILOBII-TETRAGONI (DC.) Wint. in Rabenh. Krypt.
Mi Ss 254. Ser.
Uredo vagans a Epilobi-tetragont DC. Fl. Fr. 2: 238. 1805.
Puccinia Gayophyti Billings, in King, Geol. Expl. goth Par. 5: 414.
1871.
Puccinia Oenotherae Vize, Grevillea 5: 109. 1877.
Puccinia Boisduvaliae Pk. Bot. Gaz. 7: 45. 1882.
244 BROOKLYN BOTANIC GARDEN MEMOIRS
Puccima Clarkiae Pk. Bull. Torrey Club 11: 49. 1884.
Puccinia glabella Holw. N. Am. Ured. 1: 76. 1907.
ON ONAGRACEAE:
Boisduvalia densifolia (Lindl.) Wats.—Minam River, Wallowa
Co., Oct. 2, 1897, E. P. Sheldon, 9049; Corvallis, July, 1910, 1116,
Aug. 10, I9I1, 1723, Sept..20, 1914, 1546; Calapooya Valley, Douglas
Co., July 24, 1899, M. A. Barber (Rocky Mt. Herb. 40989); Grant’s
Pass, Josephine Co., Sept. 2, 3, 1916, J. R. Weir, 222, 251.
Boisduvalia glabella (Nutt.) Walp.—Burns, Harney Co., Aug. 1901,
Griffiths & Morris (Griffiths, W. Am. Fungi 385).
Boisduvalia stricta (A. Gray) Greene—Corvallis, Aug. 13, 1914,
1492; Medford, Jackson Co., June 26, 1915, G. B. Posey, 3275;
Wimer, Jackson Co., July 22, 1892, E. W. Hammond, 149 (Rocky Mt.
Herb. 48696).
Clarkia pulchella Pursh—Hilgard, Union Co., July 10, 1914, 1520.
Epilobium minutum Lindl.—Corvallis, Aug. 15, 1909, 1170.
Eptilobium paniculatum Nutt.—Cole’s Creek, Wallowa Co., June
10, 1897, E. P. Sheldon, 8263 (Rocky Mt. Herb. 70411); Hood River,
May 14, 1914, 1510, May 16, 1915, 3271; Hilgard, Union Co., July
10, 1914, 1530, 1544; Ontario, Mahheur Co., Aug. 1901, Griffiths &
Morris (Griffiths, W. Am. Fungi 383); Corvallis, Aug. 1910, 3065,
Aug. 10, 1911, F. D. Bailey, 1774; The Dalles, Wasco Co., June 19,
1914, 3107; N. slope Mt. Hood, Aug. 7, 1914, 1491; Near Cascade
Locks, Hood River Co., Aug. 11, 1910, 1073; Philomath, May 26,
1914, 3351, April 21, 1899, Moses Craig; Klamath Falls, Klamath
Cos Septi-8, 1916, |. Rs Weirn225:
Gayophytum ramossissimum T. & G.—Redmond, Crook Co., July
I, 1914, 2536; Hood River, July 23, 1915, 3272; Farewell Bend,
Crook Co., July 15, 1894, J. B. Lieberg, 435 (Rocky Mt. Herb. 66228).
Godetia amoena (Lihm.) Lilja.—Corvallis, July, 1910, 7775, Wren,
Benton-Go.,. June 26, 1914,°7327.
Sphaerostigma Boothit (Dougl.) Walp.—Muddy Station, John Day
Valley, May 12, 1885, Thomas Howell.
Sphaerostigma dentatum (Cav.) Walp.—Pleasant Creek, near
Wimer, Jackson Co., April 23, 1889, E. W. Hammond, 143.
As here considered, this species includes all the long-cycled autoe-
cious forms occurring on Onagraceae. The treatment follows the
disposition made of them by Bisby in his recent admirable discussion
of the Onagraceous rusts (Amer. Jour. Bot. 3: 538. 1916).
100. PUCCINIA EPIPHYLLA (L.) Wettst. Verhl. Zool.-Bot. Ges. Wein.
35: 541. 1885.
Lycoperdon epiphyllum L. Sp. Pl. 1653. 1753.
Aecidium Tussilaginis Pers. in Gmel. Syst. Nat. 2: 1473. 1791.
JACKSON: UREDINALES OF OREGON 245
Puccinia Poarum Niels. Bot. Tidsskr. II. 3:26. 1877.
On POACEAE:
Poa ampla Merrill—Hood River, May 14, 1914, 1591.
Poa annua L.—Hood River, July 22, 1915, 3190.
Poa macrantha Vasey—Newport, Lincoln Co., June 20, 1915, 3123.
Poa pratensis L.—Corvallis, May 19, 1913, F. D. Bailey, rz04,
March 29, 1914, G. B. Posey, 3126, April 29, 1914, F. D. Bailey,
3125, June 29, 1914, G. B. Posey, 1311; Philomath, May Io, 1914,
3124; Hood River, May 14, 1914, 1586, 1592; The Dalles, Wasco
wo July 1, 1914, 7302; N. slope Mt: Hood, Aug. 7, 1914, 1557;
Ashland, Jackson Co., Sept. 10, 1914, 1565; Kamela, Union Co.,
July 22, 1915, M. E. Peck; Klamath Falls, Klamath Co., Sept. 8,
1916; J. R. Weir, 224, 239.
Poa triflora Gilib.—Klamath Falls, Klamath Co., Sept. 8, 1916,
J. R. Weir, 239a.
Poa sp.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir,
228, 229; Klamath Falls, Klamath Co., Sept. 8, 1916, J. R. Weir,
218; Austin, Grant Co., Aug. 1915, J. R. Weir, 194.
This rust is especially common in western Oregon particularly on
blue grass. Only uredinia are known in the above collections as is
the common condition except in those made in the far north or at
high elevations.
Nielsen (Bot. Tidsskr. 2: 26. 1877) was the first to show the
relation between this rust and Aecidium Tussilaginis Gmel. He suc-
ceeded in infecting P. annua, P. trivialis, P. nemoralis, P. fertilis and
P. pratensis by sowing aeciospores from Tussilago farfara. He in-
fected the aecial host by sowing with teliospores from P. annua.
Additional observations and culture work have been recorded by
various European authors, which have been summarized by Klebahn
(Die Wirtsw. Rostp. 290. 1904).
101. PucciInIA ERtopHoRI Thiim. Bull. Soc. Imp. Nat. Moscow 55:
208. 1880.
Aecidium Ligulariae Thiim. Nov. Giorn. Bot. Ital. 12: 196. 1880.
Aecidium Cinerariae Rostr. Overs. Kong. Dansk. Vid. Selsk. Forh.
Koph. 1884-5: 17. 1884.
On CARDUACEAE: I.
Senecio ductaris Piper—Alpine meadow, E. Mt. Hood, 5,000 ft.,
July 23, 1915, 3320.
ON CYPERACEAE: III.
Eriophorum polystachyon L.—Alpine meadow, E. Mt. Hood, 5,000
oi Muly23, 1915, 33391.
In the alpine meadow where the above collections were made the
Aecidium was very abundant and in fine condition. A search was
246 BROOKLYN BOTANIC GARDEN MEMOIRS
made for overwintered telia on Cyperaceous hosts associated with the
Senecio and the only rust found was on very much weathered leaves
which have been determined by comparison of the microscopic struc-
ture as above indicated.
Tranzschel (Beitr. Biol. Ured. III: 4. 1907), working in Russia,
was the first to culture this species. He used telial material on
Eriophorum angustifolium to successfully infect Ligularia sibirica and
Senecio paluster.
In America, Arthur (Mycol. 8: 131. 1916), using aecial material
from New York on Senecio aureus, obtained successful infection result-
ing in uredinia and telia on EL. viridi-carinatum.
102. Puccinia Eriophyllii sp. nov.
O.and I. Pycnia and aecia unknown.
Il. Uredinia amphigenous and caulicolous, scattered, small,
roundish, 0.3-0.6 mm. across, early naked, pulverulent, pulvinate,
chestnut brown, surrounding epidermis not conspicuous; uredinio-
spores globoid, obovoid or oblong, 21-27 by 26-29 yu, wall chestnut
brown, 2-3 pv thick, moderately and finely echinulate, pores 2, approxi-
mately equatorial.
III. Telia amphigenous, scattered, small, round, 0.3-0.6 mm.
across, early naked, compact, pulvinate, blackish brown, ruptured
epidermis not conspicuous; teliospores ellipsoid to oblong, 18-21 by
26-30 pw, apex and base rounded, not constricted, wall chestnut brown,
2-2.5 m thick, uniform, minutely and obscurely verrucose; pedicel
colorless, deciduous.
ON CARDUACEAE:
Eriophyllum lanatum (Pursh) Forbes—Wren, Benton Co., June
26, 1914, 1319 (type).
Eriophyllum leucophyllum (DC.) Rydberg—Redmond, Crook Co.,
July 1, 1914, 3083.
Distinguished from other species of Puccinia on related hosts by
the very small teliospores.
103. Puccinia Fendleri (Tracy & Earle) comb. nov.
Aecidium Fendleri Tracy & Earle, in Green, Pl. Baker 1:17. 1901.
Puccinia Koeleriae Arth. Mycologia 1: 247. 1909.
ON BERBERIDACEAE: I.
Berberis aquifolium Pursh—Corvallis, May 12, 1914, 1276, June 7,
1908, J. C. Bridwell, 3380.
Berberis nervosa Pursh—Hilgard, Union Co., July 10, 1914, 2568.
ON POACEAE:
Koeleria cristata (L.) Pers.—Hilgard, Union Co., July 10, 1914,
1363.
JACKSON: UREDINALES OF OREGON 247
The aecia of this species have often been confused with those of
~ P. graminis (cf. 151). Arthur in 1908 (Mycol. 1: 246. 1907), using
telial material from Koeleria cristata from Colorado, obtained, as a
result of infection experiments, the development of aecia on Berberis
aquifolium.
104. PUCCINIA GEMELLA Diet. & Holway, in Sydow’s Monog. Ured. 1:
541. 1903.
On RANUNCULACEAE:
Caltha biflora DC.—N. slope Mt. Jefferson, 6,000 ft., Aug. 13,
1914, H. P. Barss & G. B. Posey, 1624; foot of Mt. Jefferson, 5,000 ft.,
Aug. 28, 1916, H. P. Barss, 3401.
A micro-form, differing from P. Treleasiana Pazsch., which occurs
in the Rocky Mt. region on Caltha sp., in the smooth spores.
105. PuccINIA GENTIANAE (Strauss) Link, in Willd. Sp. Pl. 67: 73.
1825.
Uredo Gentianae Strauss, Ann. Wett. Ges. 2: 102. I81I0.
ON GENTIANACEAE:
Gentiana oregana Engelm.—Sumpter, Baker Co., July 16, 1913,
J. R. Weir, zor.
106. PucciniA GILIAE Hark. Bull. Cal. Acad. 1: 34. 1884.
ON POLEMONIACEAE:
Navarettia intertexta (Benth.) Hook.—Corvallis, Aug. 1898, Moses
Craig, July, t910, 1721; Umpqua Valley, Douglass Co., June, 1887,
Thomas Howell, 7835; Hood River, Aug. 17, 1888, L. F. Henderson
(673), 1120.
This is a hemi-form distinct from P. plumbaria (cf. 150) which
is an opsis-form. In P. Giliae the telia are early naked and the spores
smooth. In P. plumbaria the telia are long covered by the cinereous
epidermis and the spores are finely and closely verrucose.
107. PUCCINIA GLUMARUM (Schmidt.) Erikss. & Henn. Zeits. Pflan-
zenkr. 4: 197. 1894.
Uredo glumarum Schmidt. Allg. Oekon. Fl. 1: 27. 1827.
Puccima neglecta West. Bull. Soc. Bot. Belg. 2: 248. 1863.
Trichobasis glumarum Ley.; Cooke, Myc. Fung. 208. 1865.
ON POACEAE:
Elymus glaucus Buckl.—Hood River Co., May 14, 1914, 1590,
1590, 1597, July 23, 1915, 3199.
Hordeum Gussoneanum Parl.—Corvallis, June 4, 1914, F. D.
Bailey, 1385.
Hordeum vulgare L.—Moro, Sherman Co., June 11, 1915, F. K
Ravn and A. G. Johnson.
248 BROOKLYN BOTANIC GARDEN MEMOIRS
Sitanion hystrix (Nutt.) J. S. Smith—Redmond, Crook Co., July
I, 1914, 1423, 1429.
Sitanion jubatum J. G. Smith—Ashland, Jackson Co., June 7,
1916, H. B. Humphrey; Klamath Falls, Klamath Co., Sept. 11, 1916,
JaR Were 3e:
Triticum aestivum L.—Moro, Sherman Co., June 25, 1915, D. E.
Stephens, 3372, 3379;. Medford, Jackson Co., June 8, 1915, F.-K:
Ravn, A. G. Johnson, 3370.
Triticum compactum Host.—Moro, Sherman Co., June II, 1915,
F. K. Ravn & A. G. Johnson (Barth. Fungi Columb. 4756); June 25,
1915, D. E. Stephens, 337333376; 6350; 3365:
Triticum diococcum L.—Moro, Sherman Co., June 11, 1915, F. K.
Ravn & A. G. Johnson.
Triticum vulgare L. (Collective)—Corvallis, June 10, 1915, 2676,
2679, 2080, June 12, 1915, 3134; Hood River, July ‘22, 1015;,3 742.
Moro, Sherman Co., June I1, 1915, F. K. Ravn, A. G. Johnson, M.
A. Carleton.
This very important wheat rust was first found in the United
States, May 21, 1915, at Sacaton, Arizona, by Dr. F: K. Ravn, the
eminent Danish cereal pathologist, who at that time was making a
tour of investigation of cereal diseases in company with pathologists
of the Department of Agriculture (Carleton, Science N. S. 42: 58.
1916). A few weeks later the rust was found by Dr. Ravn and party
at Medford and Corvallis and later was detected at Moro, Oregon.
There is every evidence that this rust has been present in the western
states for some years. Several collections reported above were made
in 1914 but were confused with P. Clematidis. The writer is indebted
to Dr. H. B. Humphrey for examining most of the collections and for
the detection of several specimens belonging here, previously referred
to other species.
108. PUCCINIA GRANULISPORA Ell. & Gall.; Ellis & Ever. Bull. Torrey
Club 22:65. 1865;
On ALLIACEAE:
Allium nevit Wats.?—Austin, Grant Co., Aug. 1915, J. R. Weir,
204.
109. PUCCINIA GROSSULARIAE (Schum.) Lagerh. Tromsd. Mus.
Aarsh. 17: 60. 1895.
Aecidium Grossulariae Schum. Pl. Enum. Saell. 2: 223. 1803.
Puccinia Pringsheimiana Kleb. Zeits. fiir Pflanzenkr. 4: 194.
1894.
Puccinia Magnusii Kleb. Zeits. fiir Pflanzenkr. 5: 79. 1895.
Puccinia albiperidium Arth. Jour. Myc. 8: 53. 1902.
Puccimia uniporula Orton, Mycol. 4: 201. I9gI2.
JACKSON: UREDINALES OF OREGON 249
ON GROSSULARIACEAE: I.
Ribes divaricatum Dougl.—Bridal Veil, Multnomah Co., May 18,
1915, 3252; Hilgard, Union Co., July 10, 1914, 3001.
Ribes lacustre (Pers.) Poir.—N. slope Mt. Hood, Aug. 7, 1914, 2561.
Ribes sanguineum Pursh—Bridal Veil, Multnomah Co., May 18,
1915, 3253:
Ribes sp.—Philomath, April 26, 1914, 2571; Corvallis, April 11,
1915, 3045.
On CypERAcEAE: II, III.
Carex festiva Dewey—Hilgard, Union Co., July 10, 1914, 1360.
Carex Goodenowi J. Gay—Hilgard, Union Co., July 10, 1914, 1359.
Carex Kelloggii W. Boot—Portland, Aug. 21, 1915, E. Bartholomew
5041 (Barth. Fungi Columb. 4962).
Carex magnifica Dewey—Clatsop Co., Nov. 7, 1913, 1195.
Carex mertensit Prescott—Mt. Hood, Aug. 7, 1914, 3004.
Carex monile Tuckerm.—Clatskanie, Columbia Co., May 20, 1914,
F. D. Bailey, 3073.
Carex nebraskensis Dewey—Hilgard, Union Co., July 10, 1914,
1361.
Carex phyllomanica W. Boot?—Klamath Falls, Klamath Co.,
Sept. 8, 1916, J. R. Weir, 254.
Carex spectabilis Dewey—W. slope Mt. Jefferson, July 3, 1914, F. D.
Bailey, 1417; Vicinity Mt. Jefferson, Aug. 12, 1914, H. P. Barss &
G. B. Posey, 3007.
Carex sp.—In open meadow along Minum River, Wallowa Co.,
Aug. 20, 1897, E. P. Sheldon, 87571.
This common form having aecia on Ribes sp. was first cultured
by Klebahn in 1892. The species has since been extensively studied
by the culture method in both Europe and America (Klebahn, Die
Wirtsw. Rostp. 295-302. 1904) under various names.
In America, Arthur began culture work in rgo1 and has reported
the results of numerous cultures (Jour. Myc. 8: 53. 1902; Io: II.
Bega. Ths 59. 1905; 12:65. 1906; 13: 196... T9075. T4213. 1908;
Mycol. 4: 13. 1912; 7: 67. 1915; 7: 78. 1915). The species is
doubtless represented by several biological strains and further culture
work will need to be conducted in order to determine their limits.
Considerable confusion has resulted on account of the variable number
and position of the germ pores in the urediniospores.
110. PUCCINIA GRUMOSA Syd. & Holw. in Sydow, Monog. Ured. 1:
641. 1903.
On LILIACEAE:
Stenanthium occidentale A. Gray—Bridal Veil, Multnomah Co.,
May 18, 1915, 2670; Hood River, July 24, 1915, 3082.
250 BROOKLYN BOTANIC GARDEN MEMOIRS
This species, described from a collection on Zygadenus elegans
made by Professor Holway at Banff, Alberta, has been previously
known only from the original collection. The above collections
clearly belong here and add a new host. The only other collection of
Puccinia on Stenanthium known to the writer is one obtained by him
in January, 1917, on a phanerogamic specimen of S. gramineum col-
lected in Georgia by A. W. Curtis, now in the herbarium of the New
York Botanical Garden. This has been referred to P. atropuncta,
a species known only from east of the Rocky mountains on related
hosts.
111. PuccInIA HARKNEsSI Vize, Grevillea 7: 11. 1878.
Puccinia cladophila Pk. Bot. Gaz. 4: 127. 1879.
ON CICHORIACEAE: ;
Lygodesmia juncea (Pursh) D. Don—Denio, Harney Co., Aug.
1901, Griffiths & Morris (Griffiths, W. Am. Fungi 396c).
Ptiloria paniculata (Nutt.) Green—Sherman, Sherman Co., July 1,
1914, 2535.
112. Puccinia Helianthi-mollis (Schw.) comb. nov.
Aecidium Helianthi-mollis Schw. Schrift. d. Nat. Ges. Leipzig 1:
68. 1822.
Puccinia Helaanitht Schw. Schrift. d. Nat. Ges. Leipzig 1: 73.
1822.
ON CARDUACEAE:
Helianthus annuus L.—Sherman, Sherman Co., July 1, 1914, 2525;
Corvallis, Aug. 1910, F. D. Bailey, 71729; Umatilla, Umatilla Co.,
July 11, 1914, 1468.
113. PuCCINIA HEMIZONIAE Ell. & Tracy, Jour. Myc. 7: 43. 1891.
ON CARDUACEAE:
Hemizonia truncata (DC.) Gray—Grant’s Pass, Josephine Co.,
July 12, 1887, Thos. Howell.
Lagophylla ramossissima Nutt.—Grant’s Pass, Josephine Co.,
Septi.2; 1016, J. KR. Weir, 220.
114. PucctniA HEUCHERAE (Schw.) Diet. Ber. der Deutsch. Bot. Ges.
O42. °1So1;
Uredo Heucherae Schw. Schrift. Nat. Ges. Leipzig 1: 71. 1822.
Puccimia Tiarellae B. & C. Grevillea 3: 53. 1874.
Puccinia spreta Pk. Rep. N. Y. State Mus. 29: 67. 1878.
Puccinia congregata E. & H. Bull. Calif. Acad. Sci. 1: 26. 1884.
ON SAXIFRAGACEAE:
Heuchera micrantha Dougl.—Hood River, Feb. I, 1915, 3266;
Mary’s River, Corvallis, June 5, 1915, 2674; Ashland, Jackson Co.,
Sept. 10, 1914, 2533.
JACKSON: UREDINALES OF OREGON 251
Leptaxis Menziesit (Pursh) Raf.—Hood River, July 24, 1915,
3318, 3359.
Mitella Brewert Gray?—N. slope Mt. Jefferson, Aug. 27, 1916,
FP. Barss, 3307.
Mitella ovalis Greene—Mary’s Peak, Benton Co., May 23, 1915,
3037.
Mitella sp.—N. slope Mt. Jefferson, 8,000 ft., Aug. 8, 1914, H. P.
Barss & G. B. Posey, 2529.
Tellima grandiflora (Pursh) Dougl.—Corvallis, July 15, 1910,
wee, Apr. 8, 1914, 3075; Austin, Grant Co:, Aug..1915, J. R. Weir,
214.
Tiarella unifoliata Hook.—Bridal Veil, Multnomah Co., Aug. 11,
1910, 1070; Ashland, Jackson Co., Sept. 10, 1914, 3028.
115. Puccinia hieraciata (Schw.) comb. nov.
Caeoma (Aecidium) hieraciatum Schw. Trans. Am. Phil. Soc. II.
A202. 1832"
Puccinia patruelis Arth. Mycol. 1: 245. 1909.
On CyYpPERACEAE: I], III.
Carex praegracilis Boott (C. marcida Boott)—Ontario, Malheur
Co., Aug. 1901, Griffiths & Morris; Andrews, Harney Co., Aug. 1901,
Griffiths & Morris (Griffiths, W. Am. Fungi 339a).
This species shows a distribution from coast to coast and has
aecia on Cichoriaceous hosts. Arthur (Il. c.) has conducted one
successful culture and obtained the development of pycnia and aecia
on Agoseris glauca following sowings of teliosporic material on Carex
pratensis from Colorado. Other aecia having a similar morphology,
including a rare form on Hieracium collected by Schweinitz, on which
the present name is based, are properly referred here. No aecial
collections have been made in Oregon.
116. PucctniA HreERAciI (Schum.) Mart. Fl. Mosq. 226. 1812.
Uredo Hieracu Schum. Enum. PI. Saell. 2: 232. 1803.
Puccinia sejuncta Syd. Ann. Myc. 1: 236. 1903.
ON CICHORIACEAE:
Meracium albifiorum Hook.—Hood River, May 16, 1915, 3312,
Wily 23, 1915, 3317.
Hieracium cinereum Howell—Hood River, July 22, 1915, 3324.
Hieracium gracile Hook.—N. slope Mt. Jefferson, Aug. 6, 1914,
ff. Barss; 2547,
Meracium scoulert Hook.—Austin, Grant Co., Aug. 1915, J. R.
Weir, 156.
Meracium sp.—Hilgard, Union Co., July 9, 1914, 33129; Austin,
Grant Co., Aug. 1915, J. R. Weir, 273; Klamath Falls, Klamath Co.,
Sept. 3, 1906, J. R. Weir, 240.
252 BROOKLYN BOTANIC GARDEN MEMOIRS
This species may occur on the same plants with Aecidium colum-
biense (cf. 215) which is doubtless the aecial stage of some heteroecious
rust not yet determined. Sydow (Il. c.) has described P. sejuncta
based on such a mixture.
117. PucctntiA HoLBoELLU (Hornem.) Rostr. Middelser om Groen-
land 3: 534. 1888.
Aecidium Holboellu Hornem. Fl. Dan. 37: 11. 1840.
Puccinia Barbareae Cooke, Grevillea 8: 34. 1879.
The type of P. Barbareae was described as on a “Cruciferous
plant’’ from Oregon, Dr. Lyall 67. The data on the type collection at
the Kew Herbarium reads ‘‘Oregon Boundary Commission, Ft. Coville
to Rocky Mts. 1861, Dr. Lyall 67.”" Since Ft. Coville is in north-
eastern Washington there would seem to be little chance of this col-
lection having been made within the state of Oregon. However,
since it has been recorded from our limits both in the original descrip-
tion and by Holway (N. Am. Ured. 1: 45. 1906) it is included here
with the above explanation. The species undoubtedly does occur in
eastern Oregon as the range includes all the surrounding states.
118. PUCCINIA HOLCINA Erikss. Ann. Sci. Nat. 9: 274. 1899.
ON POACEAE:
Holcus lanatus L.—Corvallis, June 10, 1915, 2678, June 12, 1915,
3113; Toledo, Lincoln Co., July 19, 1915, 3776; Yaquina, Lincola
Co., July 17, 1915, 37177; Salem, Marion Co., May 1,.1914,)Geeme
Godfrey, 3718; Portland, Aug. 19, 1915, E. Bartholomew (Barth.
Fungi Columb. 4852).
Evidently a common rust in western Oregon, otherwise known
from North America from a few other collections made on the Pacific
coast from California to Vancouver Island, B. C., and from a single
locality along the eastern coast in Massachusetts. This species may
be easily separated from P. Rhamni (cf. 159) in the telial stage by the
evenly thickened apices of the teliospores. The urediniospores of the
two rusts are similar. Those of the present species are somewhat
larger and globoid, 19-24 by 23-27 uw, while in P. Rhamni they are
globoid or broadly ellipsoid, 16-20 by 18-24 wu.
The aecial connection is unknown. The rust has evidently been
introduced from Europe and is able to maintain itself by over-wintering
urediniospores.
119. PuccintiA HypocHoErtpis Oud. Nederl. Kruidk. Arch. II, 1: 175.
1872.
ON CARDUACEAE:
Hypochaeris radicata L.—Myrtle Creek, Douglass Co., June 9,
1914, F..D. Bailey, 2543.
JACKSON: UREDINALES OF OREGON 253
120. Puccinia insperata sp. nov.
O. Pycnia not seen.
I. Aecia chiefly hypophyllous and petiolicolous; in crowded
groups on yellowish spots 2-3 mm. across; cupulate, 0.2-.25 mm.
broad; peridium whitish, margin recurved, lacerate; peridial cells
rhombic, 19-27 by 35-45 uy, overlapping, outer wall 1-1.5 uw thick,
inner wall 3-4 yu thick, verrucose; aeciospores globoid or broadly
ellipsoid, 15-19 by 19-23 u, wall colorless, I-1.5 w thick, finely and
closely verrucose.
II. Uredinia amphigenous, scattered, round, 0.2-0.5 mm. across,
tardily naked, pulverulent, pulvinate, cinnamon brown, ruptured
epidermis conspicuous; urediniospores subglobose or broadly ellip-
soid, occasionally obovate, 19-21 by 23-29 uw; wall cinnamon brown,
1.5-2 w thick, minutely and closely echinulate, pores 2—3, scattered.
III. Telia amphigenous and petiolicolous, scattered, round, 0.2—
0.8 mm. across, tardily naked, pulvinate, becoming pulverulent,
blackish brown, ruptured epidermis conspicuous; teliospores ellipsoid
or broadly obovoid, occasionally somewhat irregular, 16-20 by 23-32 u,
rounded at both ends, scarcely or not at all constricted, wall chestnut
brown, 1.5—2 u thick, uniform, smooth; pedicel colorless, deciduous.
On CICHORIACEAE:
Nabalus hastatus (Less) Heller—Hood River, May 16, 1915, 2662,
July 24, 1915, 3265, type.
A very distinct species separable from the eastern P. orbicula
Pk. by the smooth teliospores and the presence of a definite peridium
in the aecidium. The two collections were made at the same spot.
The first shows aecia unaccompanied by pycnia, associated with
telia chiefly on the petioles, suggesting strongly an opsis-form. The
second collection, however, shows scattered uredinia and telia with a
few old aecia.
121. Puccinia Ir1pis (DC.) Wallr. in Rabenh. Krypt. Fl. 1: 23.
1844.
Uredo Iridis DC. Encycl. 8: 224. 1808.
ON IRIDACEAE:
Ins tenax Dougl.—Corvallis, June 24, 1914, F. D. Bailey, 1343;
Wren, Benton Co., July 22, 1914, 1473, Ashland, Jackson Co., Sept.
10, 1914, 1904.
122. PucCINIA JONESII Pk. Bot. Gaz. 6: 226. 1881.
Allodus Jones Arth. Résult Sci. Congr. Bot. Vienne 345. 1906.
ON UMBELLIFERAE:
Peucedanum triternata (Pursh) Nutt.—Austin, Grant Co., Aug.
Tors, |..R:. Weir, 240, rF1., *
254 BROOKLYN BOTANIC GARDEN MEMOIRS
123. Puccinia Licustici Ell. & Ev. Bull. Torrey Club 22: 263. 1895.
Puccinia luteobasis Ell. & Ev. Bull. Torrey Club 24: 457. 1897.
ON UMBELLIFERAE:
Ligusticum apufolium (Nutt.) Gray—Corvallis, 1911, 7766, April
15, 1913, F. D. Bailey, 3080, March +23, 1914, G. B. Posey, ;gg2e,
March 22, 1914, G. H. Godfrey, 2549; Orenco, Washington Co.,
June 23, 1913, 3079.
124. PUCCINIA LUXURIOSA Sydow, Monog. Ured. 1: 812. 1904.
Puccinia tosta luxurians Arth. Bull. Torrey Club 29: 229. 1902.
On POACEAE:
Sporobolus airoides Torr.—Andrews, Harney Co., Aug. Igor,
Griffiths & Morris (Griffiths, W. Am. Fungi 304).
Bethel (Phytopath. 7: 93. 1917) has reported successful cultures
of this rust on Sarcobatus vermiculatus, amply supported by field
observations. Arthur (Mycol. 1: 234. 1909) has infected that host
with teliospores of P. subnitens (cf. 167) from Nevada. Bethel,
however, failed to obtain infection on any of the aecial hosts for that
species with teliospores from Sporobolus airoides in Colorado. It is
possible that the two forms represent biological races of the same
species and should be united.
125. Puccinta MAJANTHAE (Schum.) Arth. & Holw. Bull. Lab. Nat.
Hist. Univ. Iowa 5: 188. 1901.
Aecidium Majanthae Schum. Enum. PI. Saell. 2: 224. 1803.
Puccinia sessilis Schneider, in Schréter Abh. Schles. Ges. 49: 19.
1870.
On POACEAE:
Phalaris arundinacea L.—Beulah, Malheur Co., Aug. Igor,
Griffiths & Morris (Griffiths, W. Am. Fungi 26a).
No culture work has been reported in America. In Europe the
aecia have been fouhd on Allium, Arum, Convallaria and various other
related hosts. Various names have been given to the different bio-
logical forms (Klebahn, Die Wirtsw. Rostp. 263-272. 1904; Sydow,
Monog. Ured. 1: 776-784. 1904). Aecia on Smilacina, Polygonatum,
Maianthemum and Uvularia from the central and eastern United
States doubtless belong here in whole or in part.
126. Puccinta MALVACEARUM Bert. Gay’s Hist. de Chile 8:43. 1852.
On MALVACEAE:
Abutilon ? sp. (cultivated shrub)—Corvallis, May, 1914, W. E.
Lawrence, 3364, Nov. 17, 1914, W. E. Lawrence, 3363, Jan. 12, 1915,
2627.
Althaea ficifolia Cav.—The Dalles, Wasco Co., Aug. 25, 1915, E.
3artholomew (Barth. Fungi Columb. 4758, N. Am. Ured. 1559).
JACKSON: UREDINALES OF OREGON 255
Althaea rosea Cav.—Corvallis, April 27, 1907, E. R. Lake, 1299,
Aug. 1909, 1068, Sept. I910, ror4, Jan. 12, 1915, W. E. Lawrence,
2627; Gibbon, Umatilla Co., June 5, I911, comm. 1850; Klamath
Falls, Klamath Co., Sept. 19, 1911, 1031; Salem, Marion Co., May 2,
1913, comm. 1866; Svenson, Clatsop Co., June 5, 1913, comm. 1895;
Grant’s Pass, Josephine Co., July, 1913, 1926; Dayton, Yamhill Co.,
April 6, 1914, comm. 1479; Lents, Clackamas Co., April 27, 1914,
comm. 1936; Ione, Morrow Co., June 26, 1914, comm. 1937; The
Dalles, Wasco Co., July 1, 1914, 1342; Hillsboro, Washington Co.,
July 26, 1914, 1738; Medford, Jackson Co., March 31, 1915, 2556.
Malva rotundifolia L.—Corvallis, Apr. 12, 1909, J. C. Bridwell,
3388, Aug. 30, 1913, 1741, Feb. 14, 1914, 1181; The Dalles, Wasco
ie, July 1, 1914, 7703.
Malva ? sp.—New Pine Creek, Lake Co., July, 1910, comm. 3376.
127. PuccInIA McCLaTcHIEANA Diet. & Holw. Erythea 2: 127.
1894.
ON CYPERACEAE:
Scirpus microcarpus Presl.—Elgin, Union Co., Aug. 19, 1897,
E. P. Sheldon, 8735; Beulah, Malheur Co., Aug. 1901, Griffiths &
Morris (Griffiths, W. Am. Fungi 348); Glenbrook, Benton Co., Aug.
1909, 1190; Hubbard, Marion Co., May 27, 1914, 2518; Gresham,
Multnomah Co., June 6, 1914, F. D. Bailey, 2516; Tualatin, Washing-
ton Co., July 10, 1914, F. D. Bailey, 1356; Hood River, Aug. 5, 1914,
2520; Mary’s Peak, Benton Co., Aug. 15, 1914, 2517; Orenco, Wash-
ington Co., April 2, 1915, 3386; Yaquina, Lincoln Co., July 20, 1915,
3317; Toledo, Lincoln Co., July 19, 1914, 3072.
128. PUCCINIA MELANCONOIDES Ell. & Hark. Bull. Calif. Acad. Sci.
Le.272 (LOor.
Allodus melanconioides Arth. Result Sci. Congr. Bot. Vienne 345.
1906.
ON PRIMULACEAE:
Dodecatheon latifolium (Hook.) Piper—Hills, N. W. Corvallis,
April 5, 1914, 1290, May 1, 1915, 3049, April 13, 1912, F. D. Bailey,
zoo1; Moist Woods (Corvallis?), April, 1897, Moses Craig.
129. PuccinIA MENTHAE Pers. Syn. Fungi 227. 1801.
On LABIATAE:
Mentha canadensis L.—Corvallis, Aug. 10, 1910, 1167, Nov. 4,
1911, 1772; Clatskanie, Columbia Co., Oct. 6, 1914, F. D. Bailey,
3099; Portland, Aug. 21, 1915, E. Bartholomew (Barth. Fungi Columb.
4908).
Mentha canadensis lanata Piper—Toledo, Lincoln Co., Sept 25,
1911, F. D. Bailey, rr62.
18
256 BROOKLYN BOTANIC GARDEN MEMOIRS
Mentha piperata L.—Grant’s Pass, Josephine Co., Sept. 3, 1916,
J. R. Weir, 255.
Mentha spicata L.—Hood River, Aug. 6, 1914, 17452; Elk City,
Lincoln Co., Aug. 20, 1914, 3217; Portland, April, 1914, comm. 3208;
The Dalles, Wasco Co., July I, 1914, 1340.
130. PUCCINIA MESOMEGALA B. & C. in Peck, Rep. N. Y. State Mus.
252 Tidy 2873:
Dicaeoma mesomegalum Kuntze, Rev. Gen. Pl. 3: 469. 18098.
ON CONVALLARIACEAE:
Clintonia uniflora Kunth.—Mt. Hood, Sept. 1, 1901, E. W. D.
Holway, 1016, Road to Mt. Hood, Aug. 7, 1914, 1601; Klamath Co.,
Oct. 7, 1903, E. B. Copeland (Sydow, Ured. 1776); Bridal Veil,
Multnomah Co., Aug. II, 1910, 1079; Parkdale, Hood River Co.,
March 20, 1915, L. Childs, 3788; Sumpter, Baker Co., July 16, 1913,
J. R. Weir, 276.
This very distinct micro-form is very common in the mountains
of the northwestern states on the above host, and on C. borealis in the
northern tier of states from New Hampshire to Minnesota and in
Canada.
131. PUCCINIA MICROMERIAE Dudley & Thomp. Jour. Myc. 10: 54.
1904.
On LABIATAE:
Micromeria chamissonis (Benth.) Greene (M. Douglasit Benth.)—
Mary’s Peak, Benton Co., June 20, 1910, 3163; Corvallis, June, 1910,
1157, May 4, 1912,'F. D. Bailey, 7734; Philomath, Apnl 20,srgre
F. D. Bailey, 7736, Jan. 6, 1914, 1154; Eugene, Lane Co., Julyom,
1914, G. B. Posey, 1292; N. slope Mt. Hood, Aug. 7, 1914, 2560;
Washington Co., July—Aug. 1897, Moses Craig; Grant’s Pass, Jose-
phine Co., Sept. 3, 1916, J. R. Weir, 252.
132. PUCCINIA MICROSORA K6rn.; Fuckel, Fungi Rhenani 2637.
1874.
ON CYPERACEAE:
Carex mirata Dem.—Clatsop, Clatsop Co., Nov. 7, 1913, 1196.
Otherwise known only locally from the eastern United States.
The aecial stage is unknown.
133. PucctntA MILLEFOLII Fckl. Sym. Myc. 55. 1869.
ON CARDUACEAE:
Achillea millefolium L.—Philomath, May 10, 1914, 1834; Yaquina
River, Elk City, Lincoln Co., Aug. 20, 1914, 1625; Hood River, July
22. ORS Fao
This micro-form, evidently introduced from Europe, is otherwise
JACKSON: UREDINALES OF OREGON 257
known only from a few collections made in California and single
collections from Montana and New Mexico.
134. PUCCINIA MONARDELLAE Dudley & Thomp., Jour. Myc. 10: 53.
1904.
On LABIATAE:
Monardella odoratissima (Benth.) Greene—Hilgard, Union Co.,
July 10, 1914, 1531.
Monardella villosa Benth.—Ashland, Jackson Co., Sept. 10, 1914,
2542; Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 2534.
135. PuccINIA MONOICA (Pk.) Arth. Mycologia 4: 61. 1912.
Aecidium monoicum Peck, Bot. Gaz. 4: 230. 1879.
On Poaceae: III.
Koeleria cristata (L.) Pers.—Austin, Grant Co., Aug. 1915, J. R.
Weir, 237.
This species has aecia on Avabis as has been shown by Arthur
(Mycol. 4:59. 1912; 7:75. 1915). No aecial collections have been
seen from Oregon but this stage undoubtedly occurs in the eastern
part of the state. The collection cited under P. Holboelli (cf. 117)
was said to have aecia on a part of the original specimen which un-
doubtedly are to be referred here rather than to A. Barbareae DC. as
was done by Vize in the original notice of P. Barbareae.
136. PUCCINIA MONTANENSIS Ellis, Jour. Myc. 7: 274. 1883.
Aecidium Hydrophylli Pk. Bull. Buff. Soc. 1: 68. 1873.
Aecidium Phaceliae Pk. Bull. Torrey Club 11: 50. 1884.
Aecidium Mertensiae Arth. Bull. Torrey Club 31: 6. 1904.
On HyYDROPHYLLACEAE: I.
Hydrophyllum albtfrons Heller—Corvallis, April 5, 1914, 1280;
Mary’s Peak, Benton Co., May 21, 1915, 3029.
Hydrophyllum capitatum Dougl.—Near Crystal Lake, Corvallis,
May 20, 1899, E. R. Lake, r600.
Hydrophyllum tenuipes Heller—Corvallis, April, 1910, rz7rz, 3105;
April 15, 1912, F. D. Bailey, rr2g.
Hydrophyllum sp.—Horse Creek Canon, Wallowa Co., May 14,
1897, E. P. Sheldon, Sogo.
Mertensia laevigata Piper—Jefferson Lake, Marion Co., Aug. 1892,
Moses Craig; Parmelia Lake, Cascade Mts., July 2, 1914, J. H.
Corsaut, 2554.
ON BORAGINACEAE: I.
- Phacelia heterophylla Pursh—Philomath, April 20, 1912, 1768.
Phacehia leucophylla Torr.—Austin, Grant Co., Aug. 1915, J. R.
Weir, 155.
258 BROOKLYN BOTANIC GARDEN MEMOIRS
On Poaceae: II, III.
Elymus glaucus Buckl.—Glendale, Douglass Co., July 17, 1914,
1347.
The aecial stage of this rust is very common in western Oregon
particularly on Hydrophyllum. The uredinial and telial stages are
doubtless much more common than the single record above would
indicate. The species is very difficult to separate in the uredinial
stage from P. Clematidis (cf. 85) and it is probable that some of the
collections referred to that species belong here.
Arthur (Mycol. 8: 139. 1916) sowed aecia from Hydrophyllum
capitatum on Agropyron tenerum and Elymus virginicus. On the for-
mer uredinia and telia developed, and on the latter a few uredinia only.
This is the only successful culture with this species, though aecia on
other Hydrophyllaceae and on Boraginaceae are referred here on
morphological grounds.
137. PUCCINIA MUTABILIS Ellis & Gal. Jour. Myc. 5: 67. 1889.
ON ALLIACEAE:
Allium Geyerit Wats.—Blue Mts., July 5, 1897, W. C. Cusick, 1827.
The writer is indebted to Professor Holway for the specimen on
which this record is based.
138. PUCCINIA MADIAE Syd. Monog. Ured.1: 121. 1902.
ON CARDUACEAE:
Madia elegans Don.—Corvallis, June, 1910, 2670.
Madia glomerata Hook.—Corvallis, Aug. 1899, E. R. Lake.
Madia sp.—Hood River, June 20, 1914, 3349.
This species is very close to, and possibly identical with, P. Hemi-
zoniae (cf. 113).
139. PUCCINIA OBSCURA Schroet., Pass. Nuov. Giorn. Bot. Ital. I,
OQ: 4250... “1877:
Aecidium Bellidis Thiim. Fungi Austr. 635. hyponym. 1873.
Puccima Bellidis Lagerh. Bol. Soc. Broter. 8: 134. 1890.
On JUNCACEAE:
Juncoides parviflorum (Ehrh.) Coville—Ashland, Jackson Co.,
Sept. 10, 1914, 2519; Bend, Crook Co., Sept. 11, 1916, J. R. Weir,
200; Ukiah, Umatilla Co., Aug. 21, 1903, M. A. Crosby.
No culture work has been conducted in America. Plowright
(Jour. Linn. Soc. Lond. 20: 511. 1884) has shown the aecia to be A.
Bellidis, having cultured the species in both directions. Other Euro-
pean workers have confirmed Plowright’s results (Klebahn, Die
Wirtsw. Rostp. 317. 1904).
JACKSON: UREDINALES OF OREGON 252
140. PUCCINIA OBTECTA Pk. Bull. Buff. Soc. Nat. Hist. 1: 66. 1873.
Aecidium compositarum Bidentis Burrill; DeToni in Sacc. Syll.
Fung. 7: 799. 1888.
ON CYPERACEAE:
Scirpus americanus Pers. (S. pungens Vahl.)—Westfall, Malheur
Co., Aug. 1901, Griffiths & Carter (Griffiths, W. Am. Fungi 353).
Arthur (Jour. Myc. 14: 20. 1908) has cultured this species on
Bidens. Using telia on S. americanus from Indiana successful infec-
tion resulting in pycnia and aecia was obtained on B. frondosa and
B. connata. Aecia have not been collected west of the Rocky Mt.
region. The above collection was issued as P. canaliculata, which is
now interpreted as occurring only on Cyperus having aecia on Xan-
thium and is unknown in Oregon.
141. Puccinia Ortonii sp. nov.
O. Pycnia few, imperfectly known.
I. Aecia chiefly hypophyllous, gregarious, in roundish or elongated
groups 6-8 mm. across, short cupulate, 0.2-0.3 mm. in diameter;
peridium yellowish, the margin erose; peridial cells oblong or rhombic,
20-26 by 29-35 p, slightly overlapping, the outer wall finely striate,
8-10 w thick, the inner verrucose or slightly tuberculate, 4—6 uw thick;
aeciospores globoid or broadly ellipsoid, 18-19 by 19-24 u, wall color-
less, I-1.5 w thick, very closely and finely verrucose.
Il. Uredinia amphigenous, scattered, round, 0.5-I mm. across,
tardily naked, cinnamon brown, cinereous when covered, ruptured
epidermis conspicuous, pulverulent; urediniospores broadly ellipsoid
(or when young obovoid), 19-26 by 23-32 u, wall cinnamon brown,
2-3 mw thick, moderately and very minutely and obscurely echinulate;
pores 3-5, scattered.
Ill. Telia amphigenous, scattered, round, 0.2-1 mm. across,
tardily naked, ruptured epidermis conspicuous, chestnut brown,
cinereous when covered, pulvinate, somewhat pulverulent; teliospores
broadly and somewhat angularly ellipsoid, 18-26 by 30-42 u, rounded
at both ends, slightly or not constricted at the septum, wall chestnut
brown, 1.5-2.5 w thick, uniform, smooth, with hyaline papilla over
pore of apical cell which is usually at the apex but occasionally placed
laterally, pore of lower cell varying in position from near.the pedicel
to the septum; pedicel deciduous, colorless.
ON PRIMULACEAE:
Dodecatheon Hendersonii leptophylla Suks.—Lake of the Woods,
Cascade Range, Aug. 1892, Moses Craig.
The above collection bears aecia and uredinia only. This species
differs from P. melancoides (cf. 128) in the presence of uredinia in the
life cycle. It is to be regarded as a correlated form with that species.
260 BROOKLYN BOTANIC GARDEN MEMOIRS
The species is dedicated to Prof. C. R. Orton who was the first to
separate the material from the opsis-form. The following is a list
of the specimens from other localities in the Arthur herbarium.
Dodecatheon alpinum Greene—Susanville, California, 5,000 ft.,
June 30, 1897, II, iti, M. E. Jones; Bluff Lake, San Bernardino Mts.,
California, 7,400 ft., Sept. 1895, III, Miss Nora Pettibone, 2853; Mt.
Eddy, Siskiyou Co., California, Sept. 7, 1903, 1, 11, III, E. B. Copeland,
(Sydow, Ured. 1774, type).
Dodecatheon Jeffreyi Van Houtte—South of Sitka, Alaska, u, III,
Aug. 29, 1916, J. P. Anderson, 337; Vancouver Island, British Colum-
bia, Aug. 26, 1908, 11, III, E. W. D. Holway.
Dodecatheon (tetrandrum Suks.?)—Mt. Adams, Washington, 6,000-—
7,000 ft., Aug. 31, 1886, W. N. Suksdorf (Barth. N. Am. Ured. 554,1457).
142. PUCCINIA OxyRIAE Fckl. Symb. Nachtr. 3: 14. 1875.
ON POLYGONACEAE:
Oxyria digyna (L.) Hill—Strawberry Mt., 8,o00 ft., Grant Co.,
Sept. 2; 29013) WE. bawrence;nr2:
Evidently a rather rare species represented in the Arthur her-
barium otherwise only by single collections from Colorado, Utah,
Idaho, Alberta and British Columbia on the above host.
143. PucctntA PaLMERI D. & H. Erythea 7: 98. 1899. (Not
Aecidium Palmert And. 1891.)
Allodus Palmert Orton, Mem. N. Y. Bot. Gard. 6: 202. 1916.
(Not A. Palmeri Arth. 1906.)
ON SCROPHULARIACEAE:
Pentstemon Menziesit Hook.—Near Mt. Jefferson, Linn Co., July
3, 1914, F. D. Bailey, 3046; Mt. Hood, 6,000 ft., Sept. 1, 1901, Baye
D. Holway; Horse Creek, Wallowa Co., June 24, 1897, E. P. Sheldon,
S368. .
This is an opsis-form common in the Rocky Mt. and north Pacific
states. Dietel & Holway (I. c.) based their combination on Aecidium
Palmert Anderson. This Aecidium, as has been recently determined
by Orton, working in this laboratory, is the aecial stage of the heter-
oecious rust P. Andropogonis Schw. and not the aecia of this species.
While the name P. Palmeri D. & H. has been misapplied in this way
it seems best to retain it for this species, particularly since the telia
described apply to this fungus.
144. PUCCINIA PARKERAE Diet. & Holw. Erythea 3: 78. 1895.
ON SAXIFRAGACEAE:
Ribes lacustre (Pers.) Poir.—Whitewater Ranger Station, near
Mt. Jefferson, Aug. 28, 1916, H. P. Barss, 3398; Hood River, July
23, I9I5, 3010.
JACKSON: UREDINALES OF OREGON 261
This short-cycled form possesses teliospores which closely resemble
the telia of the Carex rusts having aecia on Ribes (cf. P. Grossulariae,
109) as has been pointed out by Holway (N. Am. Ured. 1: 53. 1906).
It is not to be confused with the rust having a similar life history
in the eastern United States, P. Ribis DC. The latter has verrucose
spores, while in the one under discussion the spores are smooth.
145. PUCCINIA PATTERSONIANA Arth. Bull. Torrey Club 33:29. 1906.
ON POACEAE:
Agropyron spicatum (Pursh) Rydb.—Dufur, Wasco Co., June 19,
1914, 71398; Hilgard, Union Co., July 10, 1914, 1364; Grant’s Pass,
Josephine Co., Sept. 3, 1916, J. R. Weir, 208.
The aecial form of this distinct heteroecious rust is unknown.
146. Puccinta Peckit (DeToni) Kellerm. Jour. Myc. 8: 20. 1902.
Aecidium Oenotherae Pk. Ann. Rep. N. Y. State Mus. 23: 60.
1873. (Not P. Oenotherae Vize, 1877.)
Aecidium Peckit DeToni, in Saccardo Syll. Fung. 7: 790. 1888.
? Puccinia ludibunda E. & E. Proc. Phil. Acad. 1893: 153. 1893.
ON CYPERACEAE:
Carex Hookeriana Dewey—Springbrook, Yamhill Co., May ta,
1914, F. D. Bailey, 3015; Whitewater Station near Mt. Jefferson,
Aug. 12, 1914, H. P. Barss & G. B. Posey, 30006.
The above collections are referred somewhat doubtfully to this
species as no aecia have been collected in the Pacific northwest.
The aecia occur on Onagraceae as was first shown by Kellerman
(1. c.) and later by Arthur (Bot. Gaz. 33:13. 1903; Jour. Myc. 8: 52.
Poa ET 50.) T9055 122. 15:. 1900. > 1324195.) 19075 (Mycol: 1+ 233.
Te, 22222, 1910; Ae T5s) O12): :
147. PuccINIA PENTASTEMONIsS Pk. Bull. Torrey Club 12: 35. 1885.
ON SCROPHULARIACEAE :
Pentstemon diffusus Dougl.—Bridal Veil, Multnomah Co., May 18,
I915, 3267.
148. PUCCINIA PIMPINELLAE (Str.) Mart. Fl. Mosq. Ed. II: 226.
1817.
Uredo Pimpinellae Strauss, Wett. Ann. 2: 102. 1810.
Puccinia Osmorrhizae Cke. & Peck, in Peck, Rep. N. Y. State Mus.
20: 73. 1878.
Puccimia trifoliata E. & E. Bull. Torrey Club 22: 58. 1895.
ON UMBELLIFERAE:
Osmorrhiza brevipes (Coulter & Rose) Suks.—Corvallis, May 4,
1912, F. D. Bailey, 2550, July 3, 1914, G. B. Posey, 1998, Apr. 28,
1915, 3313; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir,
196; Austin, Grant Co., Aug. 1915, J. R. Weir, 289.
262 BROOKLYN BOTANIC GARDEN MEMOIRS
Osmorrhiza divaricata Nutt.—Jackson Co., Sept. 7, 1903, E. B.
Copeland (Sydow, Ured. 1778); Mt. Hood, Hood River Co., Aug. 7,
1914, 3081.
Osmorrhiza Liebergit (Coulter & Rose) Suks.—North of Mt.
Jefferson, Aug. 28, 1916, H. P. Barss, 3400.
Osmorrhiza occidentalis Nutt.—Paisley, Lake Co., Aug. 1914, J. C.
Elder, 3783.
149. PucciniA PipERI Ricker, Jour. Myc. 11: 114. 1905.
On Poaceae: II and III.
Festuca pacifica Piper—Eight Dollar Mt., Oregon, June 12, 1904,
C.V. Piper, 6502, type.
This species is evidently rare, since it is known only from the type
locality listed above and from two localities in California. The life
history is unknown.
150. PUCCINIA PLUMBARIA Pk. Bot. Gaz. 6: 238. 1881.
Aecidium Giliae Pk. Bot. Gaz. 4: 230. 1879. (Not P. Gihae
Hark. 1884.)
Allodus Giliae Orton, Mem. N. Y. Bot. Gard. 6: 199. 1916.
ON POLEMONIACEAE:
Gilia gracilis (Dougl.) Hook.—Mary’s Peak, Benton Co., May 21,
1915, 3310; Hood River, May 14, 1914, 1526, 2514, June 9, 1915,
3274.
Phlox speciosa Pursh—Sherman, Sherman Co., July 1, 1914, 2575.
151. PUCCINIA POCULIFORMIS (Jacq.) Wettst. Verhl. Zool. Bot. Ges.
Wein 35: 544. 1885.
Lycoperdon poculiforme Jacq. Coll. Austr. 1: 122. 1786.
Aecidium Berberidis Pers. in Gmel. Syst. Nat. 2: 1473. 1791.
Puccinia graminis Pers. Neues Mag. Bot. 1: 119. 1794.
Puccima Phlei-pratensis Erikss. & Henn. Zeit. f. Pflanzenkr. 4:
140. 1894.
Uredo quinqueporula Arth. & Fromme, Torreya 15: 265. I915.
On Poaceae: II, III.
A gropyron dasystachyum (Hook.) Vasey—Sherman Sta. O. W. R. R.
& Nav. Co., Sherman Co., July 1, 1914, 1420.
Agrostis ecarata Trin.—Ashland, Jackson Co., Sept. 10, 1914, 1567.
Alopecuris californicus Vasey—Corvallis, Sept. 21, 1914, 1552.
Avena fatua glabrata Peterm.—Philomath, Jan. 16, 1914, 1138.
Avena sativa L.—Corvallis, Aug. 12, 1911, 3161, July 6, 1914,
1667, Aug. 13, 1914, 1661, Aug. 9, 1915, G. H. Godfrey & F. D. Bailey,
3135; Cottage Grove, Lane Co., July 14, 1914, 1674; Pleasant Hill,
Lane Co., Aug: 7, 1953, F) DD: Bailey.auanco,
JACKSON: UREDINALES OF OREGON 263
Beckmannia erucaeformis (L.) Host.—Corvallis, Aug. 8, I915,
30204.
Dactylis glomerata L.—Cottage Grove, Lane Co., July 14, 1914,
1670; Corvallis, July 6, 1914, 1666; Clatsop, Clatsop Co., Nov. 3,
1913, 1155; Philomath, Jan. 6, 1914, 1084.
Deschampsia elongata (Hook.) Munro—Glendale, Douglass Co.,
July 17, 1914, 1410; Ashland, Jackson Co., Sept. 10, 1914, 1566, 1568;
Wren, Benton Co., June 26, 1914, 1329; Garden Home, Multnomah
Po, july 20, 1915, 3157, 3158.
Elymus glaucus Buckl.—Philomath, Jan. I, 1914, 1152; Bend,
rook Co:, Sept. 11, 1916, J. R. Weir, 272.
__ Festuca elatior L.—Wren, Benton Co., June 26, 1914, 1324; Cottage
Grove, Lane Co., July 14, 1914, 1348; Corvallis, July 29, 1915, 3190.
Festuca megalura Nutt.—Corvallis, June 24, 1914, 71390, Aug. 13,
1914, 3791.
Festuca myuras L.—Cottage Grove, Lane Co., July 14, 1914, 1351.
Festuca pacifica Piper—Corvallis, July 6, 1914, 1434, July 29, 1914,
1410.
- Hierochloe macrophylla Thurb.—Glendale, Douglass Co., July 17,
1914, 1411 (type of Uredo quinqueporula).
Hordeum distichon L.—Corvallis, Aug. 20, 1915, H. P. Barss, 3184.
Hordeum vulgare L.—Corvallis, Oct. 5, 1914, 3165.
Lolium multiflorum Lam.—Cottage Grove, Lane Co., July 14,
1914, 1349; Corvallis, Sept. 20, 1914, 1551; Tualatin, Washington
Co., July 10, 1914, F. D. Bailey, 1357.
Lolium subulatum Vis.—Corvallis, March 22, 1915, 3277.
Phleum pratense L.—Cottage Grove, Lane Co., July 14, 1914, 1668;
Hood River, June 19, 1914, 1665; Springbrook, Yamhill Co., June
oe, Toi4, Fk: 1. Bailey, 3736; Briton, Lincoln, Ca:; July 17, 1915,
VanGundia, 3086, 3087; Corvallis, Aug. 13, 1914, 1662; Philomath,
Jan. 1, 1914, 71151; Gresham, Multnomah Co., Aug. 7, 1913, F. D.
Bailey, 1140; Sumpter, Baker Co., June, 1913, J. R. Weir, 97; Grant’s
Pass, Josephine Co., Sept. 2, 1916, J. R. Weir, 246.
Triticum compactum Host.—Moro, Sherman Co., June I1, 1915,
F. K. Ravn & A. G. Johnson.
Triticum vulgare L.—Corvallis, July 29, 1914, 1684; Hood River,
mug. 5, 1914, 1660; Ashland, Jackson €o., Aug. 28, 1913, 1725;
Union, Union Co., Aug. 13, 1915, F. D. Bailey, 3733; Albany, Linn
Co., Aug. 22, 1900, E. B. Townsend, 3384.
Since the classic researches of DeBary, who first demonstrated
heteroecism in rusts by showing that this species has aecia on Berberis,
this rust has received more attention on the part of investigators than
any other species. (Klebahn,. Die Wirtsw. Rostp. 205-235. 1904.)
264 BROOKLYN BOTANIC GARDEN MEMOIRS
In America the most important work has been conducted by
Carleton (Div. Veg. Phys. & Path. U.S. D. A. Bull. 16. 1899;)#Bae
Pl. Industry, U.S. D. A. Bull. 63. 1904); Arthur (Jour. Myce. 8: 53.
1902; II: 57. 1905; 12217. 1906; 13:°198. “1907; "147 1o7 ome
Mycol. 2: 227. 1910; 4:18. 1912); Freeman & Johnson (Bur. PI.
Ind. U.S. D. A. Bull. 216. 1911); Stakman (Minn. Exp. Sta. Bull.
138. 1914; Jour. Agr. Research 4: 193-199. 1915) and Stakman
and Piemeisel (Jour. Agr. Research 6: 813-816. 1916; 10: 429-495.
1917).
In Oregon the rust is apparently not as important on wheat and
other grains as it is in the spring wheat districts east of the Rocky
mountains. No aecial collections have been made.
A number of unrecorded hosts appear in the above list. Since
the publication’ of Uredo quinqueporula by Arthur and Fromme (I. c.),
telia have been found on a duplicate specimen which determines that
that species is properly referred here. The number of pores in the
uredospores of that collection is unusual for this rust, the usual number
being 4, and in the absence of telia was considered of sufficient impor-
tance to separate it as a distinct species.
152. PUCCINIA POLYGONI-ALPINI Cruchet & Mayor, Bull. Herb. Bois
8: 245. 1908.
ON POLYGONACEAE:
Rumex paucifolius Nutt.—Crater Lake, Klamath Co., Sept. 9,
1916, J..R. Weir, 253.
This specimen is referred to the above species on account of the
hyaline umbo covering the pore of the apical cell in the teliospore.
The species is described from material on Polygonum alpinum from
Europe with which our material closely agrees. A collection on that
host from Idaho is also to be referred here. The rust is unlike any
other recorded on Rumex. The only other collection recorded on this
host from North America is the one on which P. uniformis Pammel &
Hume from Wyoming was based, which Holway (N. Am. Ured. 1: 36.
1906) considers to be on Polygonum sp. and refers to P. Bistortae.
153. PUCCINIA POLYGONI-AMPHIBII Pers. Syn. Fung. 227. 1801.
Aecidium Geranit maculati Schw. Schr. Nat. Ges. Leipzig 1: 67.
1822.
ON POLYGONACEAE:
Polygonum amphibium L.—Brandt’s Ranch, Wallowa Valley, Aug.
26, 1897, E. P. Sheldon, 8972; The Dalles, Wasco Co., Aug. 25, 1915,
E. Bartholomew (Barth. N. Am. Ured. 1566):
Polygonum Muhlenbergiu S. Wats. (P. emersum Britt.) —The Dalles,
Wasco Co., Aug. 26, 1915, E. Bartholomew (Barth. Fungi Columb.
JACKSON: UREDINALES OF OREGON 265
4762); Portland, Aug. 21, 1915, E. Bartholomew (Barth. Fungi
Columb. 4861).
Polygonum pennsylvanicum L.—Corvallis, Sept. 20, 1914, 1547;
Clatskanie, Columbia Co., Oct. 10, 1914, F. D. Bailey, 1944.
No collections of aecia referable to this species have been made
west of the Rocky mountains. Tranzschel (Centr. f. Bakt. II, 11:
106. 1903) was the first to show that this species has aecia on Gera-
nium. Arthur working with American material has confirmed
Tranzschel’s results (Jour. Myc. 11: 59. 1905; 12:18. 1906).
154. PUCCINIA PORPHYROGENITA Curt.; DeToni in Sacc. Syll. 7: 703.
1888.
Puccimia porphyrogenita Curt. in Thiim. Myc. Univ. 545 (hypo-
nym). 1876.
Puccinia acuminaia Pk. Rep. N. Y. State Mus. 23: 57. 1872.
(Not P. acuminata Fckl. 1869.)
On CORNACEAE:
Cornus canadensis L.—Near Mt. Jefferson, Aug. 1892, Moses
Craig; Larch Mt., Multnomah Co., Aug. 11, 1910, 1078; South Mt.
Jefferson, Linn Co., July 3, 1914, F. D. Bailey, 7840; Mt. Hood,
Aug. 7, 1914, 1604; Trail to Hanging Valley, Be Jefferson, Aug. II,
1914, H. P. Barss & G. B. Posey, 1622.
155. PUCCINIA PROCERA Diet. & Holw. Erythea 1: 249. 1893.
ON POACEAE:
Elymus arenicola Schrib. & Smith—Umatilla, Umatilla Co., May
1915, 3200, 3201, July 11, 1914, 1374, 1375; Sherman Sta. O. W.
R. R. & Nav. Co., Sherman Co., July I, 1914, 142z.
This species is distinguishable from other forms on Elymus by
the large urediniospores, 26-32 by 32-48 wu. The aecial connection is
unknown.
156. PUCCINIA PUNCTATA Link, Ges. Nat. Freunde Berlin Mag. 7: 30.
1816.
ON RUBIACEAE:
Galium aparine L.—Hood River, July 24, 1915, 3225; Ashland,
Jackson Co., Sept. 10, 1914, 3239; Corvallis, May 1, 1915, 3148.
Galium asperrimum A. Gray—Big Canon, Wallowa Co., Aug. 24,
1897, E. P. Sheldon, 8774.
Galium triflorum Michx.—Mary’s Peak, Benton Co., Aug. 15, 1914,
1513, 1515; Elk City, Lincoln Co., Aug. 20, 1914, 2528; Hood River,
July 24, 1915, 3224.
Galium sp.—Philomath, April 20, 1914, F. D. Bailey, 2570; Cor-
vallis, April 8, 1914, 1524; Dufur, Wasco Co., June 30, 1914, 1335.
266 BROOKLYN BOTANIC GARDEN MEMOIRS
157. PUCCINIA PYGMAEA Erikss. Fungi Par. Scand. 9: 449. 1895.
ON POACEAE:
Calamagrostis aleutica Bong.—Newport, Lincoln Co., July 18, 1915,
3204, Aug. 30, 1914, 1579.
The above collections show uredinia only. The aecial connection
is unknown.
158. PUCCINIA RECEDENS Syd. Monog. Ured. 1: 146. 1902
On CARDUACEAE:
Senecio harfordii Greenman—Bridal Veil, Multnomah Co., May
18, LOL5, 3273:
Senecio sp.—Hilgard, Union Co., July 10, 1914, 7542.
159. PuccINIA RHAMNI (Pers.) Wettst. Verhl. Zool.-Bot. Ges. Wein
35: 545. 1885.
Aecidium Rhamni Pers. in Gmel. Syst. Nat. 2: 1472. 1791.
Puccinia coronata Corda, Icones 1: 6. 1837.
ON RHAMNACEAE: I.
Rhamnus purshiana DC.—Corvallis, July 5, 1911, F. D. Bailey,
1135, May 9, 1914, 1627, May 12, 1914, 17277, July 5; 1014; ieee
Barss, 1940; Hood River, May 14, 1914, 1278; Clatskanie, Columbia
Co., May 20, 1914, F. D. Bailey, 1251; Cottage Grove, Lane Go.,
May I, 1915, C. E. Stewart, 3058.
On PoacEAE: II, III.
Agrostis alba L.—Toledo, Lincoln Co., July 19, 1915, 3779.
Agrostis alba maritima Meyer—Philomath, Jan. 6, 1914, 1140, I150.
Agrostis exarata Trin.—Corvallis, Feb. 14, 1914, 3008, June 29,
1914, G. B. Posey, 2306, 1308, Sept. 20, 1914, 7553, Sept. 5,.memar
1578, Dec., 1915, G. B. Posey; Hood River, June 19, 1904; caam
Philomath, June 26, 1914, 17344; Eugene, Lane Co.,; July 11, 9004
G. B. Posey, 1377; Newport,, Lincoln Co., July 18, 1or5).¢ee0
Garden Home, Multnomah Co., July 20, 1915, 372z; Portland,
Aug. 21, 1915, E. Bartholomew, 5943.
Agrostis foliosa Vasey—Yaquina, Lincoln Co., July 17, 1915, 3722.
Agrostis longiligula Hitchc.—Jetty, Lincoln Co., July 19, 1915,
VanGundia, 32506.
Agrostis microphylla Steud.—Wren, Benton Co., June 26, 1914,
1314; Cottage Grove, Lane Co., July 14, 1914, 1353; Corvallis,
July 6, 1914, F. D. Bailey, 14306.
Avena sativa L.—Newport, Lincoln Co., July 18, 1915, 3132;
Briton, Lincoln Co., July 19, 1915, G. VanGundia, 3088; Marshfield,
Coos Co., July, 1916, C. E. Owens; Myrtle Creek, Douglass Co., June
9, 1914, F. D. Bailey, 3762.
Calamagrostis canadensis (Michx.) Beauv.—Clatskanie, Columbia
Co., May 20, 1914, F. D. Bailey, 1580.
JACKSON: UREDINALES OF OREGON 267
Calamagrostis hyperborea Lange—Clatsop, Clatsop Co., Nov. 7,
POLL, FI0O.
Festuca elatior L.—Elk City, Lincoln Co., Aug. 20, 1914, 1380.
Festuca subulata Trin.—Ashland, Jackson Co., Sept. 10, I914,
1561; Mary’s Peak, Benton Co., Aug: 15, 1914, 1572; Elk City,
Lincoln Co., Aug. 20, 1914, 1382.
Holcus lanatus L.—Canby, Clackamas Co., July 21, 1911, 1182;
Pilomath, Jan. 6,° 1914, 3715, 1133; Cottage Grove, Lane Co.,
July 14, 1914, 3114; Mouth of Salmonsberry River, Tillamook Co.,
July 17, 1915, G. VanGundia, 3085, 3129; Yaquina, Lincoln Co.,
July 17, 1915, 3212; Jetty, Lincoln Co., July 19, 1915, G. VanGundia,
3130; Eddyville, Lincoln Co., Aug. 10, 1915, Hoerner, 3090; Grant’s
Pass, Josephine Co., Sept. 2, 1916, J. R. Weir, 227; Portland, Jan. 9,
1914, 1139; Elk City, Lincoln Co., Aug. 20, 1914, 1370.
Lolium multiflorum Lam.—Near Gray Station, Linn Co., July 4,
1914, 1419; Corvallis, July 6, 1914, 1435, Sept. 20, 1914, 1554; New-
port, Lincoln Co., July 21, 1915, 3728.
Lolium perenne L.—Corvallis, Aug. 3, 1914, 1412, Sept. I, 1914,
B27.
Panicularia elata Nash—Clatskanie, Columbia Co., Aug. II, 1913,
F. D. Bailey, 1705.
Panicularia pauciflora (Presl.) Kuntze—Orenco, Washington Co.,
June 13, 1914, 2388; Neah-Kah-Nie Mt., Tillamook Co., Sept. 17,
1915, F. D. Bailey, 3258; Portland, Aug. 21, 1915, E. Bartholomew,
5942 (Barth. Fungi Columb. 4973).
This coronate-spored grass rust is evidently very common through-
out western Oregon on native grasses. It is, however, not common in
the Willamette valley on oats. All of the collections on that host are
from near the sea coast.
DeBary (Monat. Akad. Wiss. 211. 1866) was the first to conduct
cultures indicating the genetic connection with aecia on Frangula
and Rhamnus in Europe. Since that time many European investi-
gators have conducted culture experiments (Klebahn, Die Wirtsw.
Rostp. 254-262. 1904).
In America this species has been cultured by Arthur (Bull. Lab.
Nat. Hist. State Univ. Iowa 4: 398. 1898; Jour. Myc. 11:58. 1905;
Mycol. 4: 18. 1912) and Carleton (Div. Veg. Phys. & Path. U. S.
Dept. Agr. 16: 48. 1899; Bur. Pl. Industry, U. S. Dept. Agr. Bull.
63:15. 1904).
The only culture made with Pacific coast material was made in
1916 in this laboratory under the writer’s direction, using telial
material on Agrostis exarata sent to the writer from Corvallis by
G. B. Posey. This was used to inoculate Rhamnus Purshiana, with
268 BROOKLYN BOTANIC GARDEN MEMOIRS
the development of pycnia and aecia. This host is the only one on
which aecia have been collected in Oregon, and they are very abundant,
as the number of collections indicates.
160. Puccinia Romanzoffiae sp. nov.
O. Pycnia not seen.
III. Telia chiefly hypophyllous and petiolicolous, crowded on con-
fluent groups, 0.5-I mm. across or covering extensive areas on the
petioles, early naked, pulverulent, chestnut brown, ruptured epi-
dermis noticeable; teliospores somewhat irregularly ellipsoid or oblong,
19-24 by 34-42 u, rounded above and below, not or scarcely con-
stricted; wall chestnut brown, 2-3 wu thick, marked by large sparsely
distributed irregular tubercles, thickened at apex by a low sub-hyaline
umbo to 4-5 p, pore of lower cell at septum similarly thickened; pedicel
colorless, short deciduous.
‘(ON HYDROPHYLLACEAE:
Romanzoffia sitchensis Bong.—Mt. Jefferson, 8,000 ft., Aug. 14,
1914,,11. PB. Barss, 2530, ype:
This species is distinguished from other species on this family of
hosts by the character of the markings of the teliospore. In P.
Hydrophylli Pk. the teliospores are closely and finely verrucose while
in P. Phaceliae Syd. & Holw. they are smooth. The character of the
teliospores in the latter species suggests a correlation with P. mon-
tanensis (cf. 136) which has aecia on Phacelia and other members of
the family Hydrophyllaceae.
161. PUCCINIA RUBEFACIENS Johans. Bot. Centr. 28: 394. 1886.
ON RUBIACEAE:
Galium boreale L.—Hilgard, Union Co., July 10, 1914, 1540;
Austin, Grant Co., Aug. 1916, J. R. Weir, 247.
The teliospores of this micro-form are very similar in shape and
size to those of the opsis-form P. ambigua (cf. 61) and the eu-form
P. punctata (cf. 156). These three species on Galiwm doubtless repre-
sent a series of correlated forms.
162. PUCCINIA RUGOSA Billings, King’s Rep. 4oth Par. 914. 1871.
(Not P. rugosa Speg. 1886.)
Puccinia Troximontis Pk. Bot. Gaz. 6: 227. 1881.
Puccinia Columbiensis E. & E. Proc. Phil. Acad. 1893: 153. 1893.
ON CICHORIACEAE:
Agoseris laciniata (Nutt.) Green—Corvallis, July 10, 1915, 3275.
163. PUCCINIA SAXIFRAGAE Schlecht. Fl. Berol. 2: 134. 1824.
Puccima curtipes Howe, Bull. Torrey Club 5:3. 1874.
JACKSON: UREDINALES OF OREGON 269
ON SAXIFRAGACEAE:
Saxtifraga Marshalliit Greene—Hood River, May 16, 1915, 3268;
Mary’s River, E. of Wren, Benton Co., April 17, 1915, 2617.
Saxifraga odontoloma Piper—Corvallis, May 1, 1915, 3260.
164. PUCCINIA SHERARDIANA Koern. Hedw. 16: 19. 1877.
Puccimia Malvastri Peck, Bull. Torrey Club 12: 35. 1885.
ON MALVACEAE:
Sidalcea virgata Howell—Corvallis, May 31, 1892, A. T. Mulford,
BAer. 13, 1912, F. D. Bailey, 3354, June 23, 1913, F. D: Bailey, 1726,
Apr. 8, 1914, 3352, Apr. 29, 1914, G. B. Posey, 3353, Apr. 30, 1915,
3071; Newburg, Yamhill Co., Apr. 9, 1915, F. D. Bailey, 3072.
165. PUCCINIA SIDALCEAE Holw. N. Am. Ured. 1: 67. 1907.
On MALVACEAE:
Sidalcea oregana Gray—Klamath Co., July 10, 1903, E. B. Cope-
land, type.
This collection was distributed as P. Sphaeralceae E. & E. in
Sydow’s Uredineen 1782.
166. PuccINIA STIPAE Arth. Bull. Iowa Agr. College Dept. Bot.
1884: 160. 1884.
On Poaceae: II and III.
Stipa comata Trin. & Rupr.—Umatilla, Umatilla Co., July 11,
1914, 1369, May 11, 1915, 3205; Hermiston, Umatilla Co., May 12,
1915, 3200.
This species has aecia on various genera of Carduaceae including
Aster, Solidago, Grindelia and Senecio, as has been shown by Arthur
Went. Myc. :t1r: 63. 1905; -Mycol..4>,10. 19123. 72-72" 1915):
No aecial collections have been made in Oregon, though that stage
is doubtless not uncommon in the eastern part of the state (cf. 69).
167. PUCCINIA SUBNITENS Dietel, Erythea 3: 81. 1895.
? Aecidium Sarcobati Pk. Bot. Gaz. 6: 240. I881.
ON CHENOPODIACEAE: I.
Sarcobatus vermiculatus (Hook.) Torr.—Eastern Oregon, Aug.
1902, D. Griffiths (Vestergren, Micro. Rar. Sel. 852).
On Poaceae: III.
Distichlis spicata (L.) Greene—LaGrand, Union Co., March, 1915,
C. C. Cate, 3278; Umatilla, Umatilla Co., July 11, 1914, 1367, 1373;
Moro, Sherman Co., Aug. 4, 1914, C. R. Ball, 1856.
This remarkable species has aecia on a large number of hosts in
the Polygonaceae, Chenopodiaceae, Amaranthaceae, Cruciferae, etc.
as was first shown by Arthur (Bot. Gaz. 35: 19. 1903; Jour. Myc.
Ei; 54.) 1905, 12 16.: 1906) 13:.197) 1907; Ta205: (1908; Mycol.
270 BROOKLYN BOTANIC GARDEN MEMOIRS
I: 234. 1909, 2:.225. I910, 4: 18. 1912). Bethel((Phytopathaa,
92-94. 1917) has also conducted very extensive cultures and proven
the genetic connection with aecia on many hosts.
In 1915, Arthur (Mycol. 8: 135. 1916), using telial material
sent by the writer collected by Mr. C. C. Cate at LaGrand, Ore.,
obtained the development of aecia on Chenopodium album. This is
the only culture made with material from the Pacific coast.
The aecia on Sarcobatus are included here on the strength of
culture work conducted by Arthur, in which he obtained aecia on that
host using telial material on Distichlis from Nevada. The matter is
complicated by the fact that Bethel (I. c.) finds that the aecia on this
host in Colorado go to P. luxuriosa (cf. 124) and P. subnitens can not
be made to infect Sarcobatus. It is possible that the two species repre-
sent closely related biological forms.
168. PUCCINIA SYMPHORICARPI Hark. Bull. Calif. Acad. Sci. 1: 35.
1884.
ON CAPRIFOLIACEAE:
Symphoricarpos albus (L.) Blake—Corvallis, Oct. 17, 1909, F. L.
Griffin, 1047, July, 1910, 1086, Aug. 1910, 1050, May 4, 1912, 1087,
May 19, 1913, F. D. Bailey, 7758; Sheridan, Yamhill Co., July 7,
1914, H. P. Barss, 1291, July 8, 1916, H. P. Barss, 3396; North slope
Mt. Hood, Aug. 7, 1914, 7611; Grant’s Pass, Josephine Co., Sept. 3,
1916, J. R. Weir, 234; Hood River, July 24, 1915, 3064.
This micro-form is exceedingly abundant in western Oregon. This
species is morphologically correlated with the telial stage of P. abun-
dans (cf. 59), which has aecia on the same host, as has been pointed
out by Travelbee (Proc. Ind. Acad. Sci. 1914: 233. 1915).
169. PuccInIA TARAXACI (Reb.) Plowr. British Ured. & Ustil. 186.
1889.
Puccinia Phaseolht var. Taraxact Reb. Fl. Noem. 356. 1804.
ON CICHORIACEAE:
Taraxacum Taraxacum (L.) Karst.—Corvallis, June, 1910, 11717;
Bonneville, Multnomah Co., Aug. 10, 1910, 3078; Ashland, Jackson
Co., Sept. 10, 1914, 3074; The Dalles, Wasco Co., July 1, 1914, 3073;
Talent, Jackson Co., June 22, 1915, G. B. Posey, 3077; Newburg,
Yamhill Co., Apr. 13, 1914, F. D. Bailey, 3076.
170. Puccinia Toumeyi Syd. in Sacc. Syll. Fung. 16: 299. Feb.
1902.
Puccinia circinans Ell. & Ev. Bull. Torrey Club 27: 61. Ig00.
(Not P. circinans Fckl. 1869 or Dietel 1897.)
Puccinia chasmatis Ell. & Ev. Jour. Myc. 8: 15. May, 1902.
JACKSON: UREDINALES OF OREGON 271
ON SCROPHULARIACEAE:
Pentstemon sp.—Canyon City, Grant Co., Aug. 26, 1914, W. E.
Lawrence, 3185.
171. PUCCINIA TRAUTVETTERIAE Syd. & Holw. in Sydow, Monogr.
Ured. i: 552. “1903:
ON RANUNCULACEAE:
Trautvetteria grandis Nutt.—S. W. slope Mt. Hood, July 23, 1915,
B25.
This interesting micro-form, known only from a few collections
from the mountains of the northwestern states, has also been reported
from Japan.
£72, PUCCINIA TRITICINA Erikss. Ann. Sci. Nat. VIII. 9: 270. 1899.
On POACEAE:
Triticum aestivum L.—Hood River, June 20, 1914, 1399.
Triticum ovatum Rasp.—Myrtle Creek, Douglass Co., June 9,
1914, F. D. Bailey, 1407.
Triticum vulgare L.—Lebanon, Linn Co., Aug. 2, 1913, F. D. Bailey,
1126; Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, z947;
Cottage Grove, Lane Co., July 14, 1914, 1676; Corvallis, July 6, 1914,
maa 3100, july 10, 1914, F. D. Bailey, 7677, July 29,1914, 1685;
Bend, Crook Co., Sept. 11, 1916, J. R. Weir, 207.
This, the common leaf rust of wheat, is very abundant in western
Oregon. The life history is unknown. In morphological characters,
it resembles closely the forms on native grasses commonly referred to
P. rubigo-vera, most of which are now included in P. Clematidis (cf. 85).
173. PUCCINIA UNIVERSALIS Arth. Jour. Myc. 11: 21. 1908.
Aecidium Dracunculi Thiim. Bull. Soc. Nat. Moscow. 58: 212.
1878. (Not P. Dracuncult Auers. 1850.)
On CARDUACEAE: I.
Artemisia sp.—White Pine,Baker Co., June, 1913, J. R. Weir, 720.
On CYPERACEAE: II, III.
Carex multicaulis Bailey—Grant’s Pass, Josephine Co., May 5,
1887, Thomas Howell.
Carex praegracilis W. Boott. (C. marcida Boott.)—Redmond,
Crook Co., July 2, 1914, 1425.
Carex Rossti Boott.—Hood River, July 23, 1915, 3289.
Carex umbellata Schk.—Hood River, July 23, 1915, 3282.
This species has aecia on Artemisia as has been shown by Arthur
Hour. Myc, 14:21; “1908; Mycol.\2a:2e4. ora, 42:16: “1912).
174. PUCCINIA URTICATA (Lk.) Kern, Mycol. 9: 214. 1917.
Caeoma urticatum Link, in Willd. Sp. Pl. 67: 62. 1825.
19
272 BROOKLYN BOTANIC GARDEN MEMOIRS
Puccinia Urticae Lagerh. Mitt. Bad. Ver. 2: 72. 1889. (Not P.
Urticae Barcl. 1887.)
Puccinia Garrett Arth. Bull. Torrey Club 32: 41. 1905.
ON URTICACEAE: I.
Urtica Lyallit S. Wats.—Philomath, April 26, 1914, 1829, May 10,
1914, 2569; Corvallis, May I, 1915, 3052.
On CYPERACEAE: II, III.
Carex Barbarae Dewey (C. laciniata Boott.)—Grant’s Pass,
Josephine Co., Sept. 3, 1916, J. R. Weir, 250.
Carex magnifica Dewey—Newberg, Yamhill Co., April 13, 1914,
F. D. Bailey, 3009; Clatskanie, Columbia Co., May 20, 1914, F. D
Bailey, 3002; Neah-Kah-Nie Mt., Tillamook Co., Sept. 17, 1915,
Dt Bailey, 6357.
Carex nebraskensis Dewey—Andrews, Harney Co., Aug. 1901,
Griffiths & Morris (Griffiths, W. Am. Fungi 339).
Carex rostrata Stokes (C. utriculata Boott.)—Clatskanie, Columbia
Co., May 20, 1914, F. D. Bailey, 30z0; Redmond, Crook Co., July 7,
1914, 1433.
Carex sp.—Hood River, May 14, 1914, 3021.
The connection of this Carex rust with aecia on Urtica was first
shown by Magnus in 1872 (Vehr. Bot. var. Prov. Brandb. 14: IT.
1872). Many other European investigators have confirmed Magnus
results (Klebahn, Die Wirtsw. Rostp. 293. 1904).
In America Arthur has conducted numerous successful culture
experiments with this species (Bot. Gaz. 29: 270. 1900; 35: 16.
1903; Jour. Myc. ‘8: 52: 19023 “12: 15." 1906; “14s 14e eee
Mycol. 2: 223. 1910; 4:17. 1912): Kellerman has also conducted
successful culture experiments (Jour. Myc. 9: 9. 1903). None of
the culture work, however, has been conducted with Pacific coast
material.
175. PUCCINIA VERATRI Duby, Bot. Gall. 2: 890. 1830.
Puccinia Verairi Clinton, in Peck, Rep. N. Y. State Mus. 27: 103.
1875.
On ONAGRACEAE: I.
Epilobium sp.—Parkdale, Hood River Co., May 14, 1914, 1511.
ON LILinceAn: Tih
Veratrum californicum Durand—The Meadows, Wallowa Co.,
Aug. 18, 1897, E. P. Sheldon, 8774.
Veratrum viride Ait.—Calloway Station, Benton Co., June 28,
1901, E. R. Lake, 7737; Parkdale, Hood River Co., May 14, 1914,
1279; Hilgard, Union Co., July 10, 1914, 1934.
The aecial collection is referred here with some confidence. It
was made in the immediate vicinity of Veratrum plants showing fresh
JACKSON: UREDINALES OF OREGON 273
uredinia. The aecia were somewhat old and no uredinia were found
on other Epilobium plants in the vicinity. Tranzschel (Ann. Mye. 7:
182. 1909) established the connection of aecia on Epilobium with P.
Veratri, obtaining his clew from the close morphological resemblance
of the teliospores of this species to those of P. Epilobii DC. Bisby
(Am. Jour. Bot. 3: 527-561. 1916) has pointed out the morpho-
logical similarity of this species with Uromyces plumbarius (cf. 201),
P. Epilobu, P. Epilobu-tetragoni (cf. 99) and P. Epilobii-Fleischert.
176. PuccinIA VIOLAE (Schum.) DC. FI. Fr. 6: 62. 1815.
Aecidium Violae Schum. Enum. PI. Saell. 2: 224. 1803.
ON VIOLACEAE:
Viola adunca J. F. Smith—Mary’s Peak, Benton Co., May 21,
IOUS, 3223.
Viola glabella Nutt.—Corvallis, Linn Co., April 16, 1912, 1081;
Mocvyalis, May 19, 1912, F. D. Bailey, 3708, April 29, 1914, F: D.
Bailey, 3164, July 14, 1914, H. P. Barss, 2548; Hood River, May 14,
1914, 3197; Portland, Aug. 30, 1915, E. Bartholomew, 5978 (Barth.
N. Am. Ured. 1677); Sumpter, Baker Co., July 16, 1913, J. R. Weir,
180; Mary’s Peak,- Benton Co., Aug. 15, 1914, 2547.
Viola nephrophylla Greene—Hilgard, Union Co., July 10, 1914,
2557:
Viola rugulosa Greene—Horse Creek Canyon, Wallowa Co.,
June 4, 1897, E. P. Sheldon, 8258.
Viola sp.—N. slope Mt. Hood, Aug. 7, 1914, 2553.
177. PuCCINIA WULFENIAE Diet. & Holw. Erythea 3: 79. 1895.
Puccinia Syntheridis Ell. & Ev. Bull. Torrey Club 27: 61. Igoo.
ON SCROPHULARIACEAE:
Synthyris rotundifolia Gray—Philomath, April 20, 1912, 1146;
Corvallis, April 8, 1914, 12806.
178. TRANZSCHELIA PUNCTATA (Pers.) Arth. Résult Sci. Congr. Bot.
Vienne 340. 1906.
Aecidium punctatum Pers. Ann. Bot. Usteri 20: 135. 1796.
Puccima Pruni-spinosae Pers. Syn. Fung. 226. 1801.
ON ROSACEAE:
Amygdalus Persica L.—Kiger’s Island, Benton Co., Oct. 5, 1913,
C.M. Scherer, 7825.
Prunus domestica L. (Italian Prune)—Salem, Marion Co., Aug.
1909, 1062; Yamhill Co., Sept. 9, 1911, 1040; Corvallis, Oct. 29,
1914, G. B. ‘Posey, 3770.
- This is not an uncommon disease of the prune, though apparently
doing little damage. It is less common on the peach. No aecial
collections have been made in the northwest.
274 BROOKLYN BOTANIC GARDEN MEMOIRS
Tranzschel (Trans. Bot. Acad. St. Petersb. 11: 67-69. 1905) was
the first to culture this species showing that aecia occur on Anemone.
In America Arthur (Jour. Myc. 12: 19. 1906; 13: 199. 1907)
has shown that the aecia on Hepatica common in the eastern United
States are genetically connected.
179. UROMYCES AEMULUS Arth. Bull. Torrey Club 38: 373. IgI1I.
Nigredo aemula Arth. N. Amer. Flora 7: 241. 1912.
On ALLIACEAE:
Allium validum S. Wats.—Paisley, Lake Co., Aug. 1914, J. S.
Elder, 1987.
180. UROMYCES AMOENUS Syd. Ann. Myc. 4: 28. 1906.
ON CARDUACEAE:
Anaphalis margaritacea occidentalis Greene—Hood River, July 23,
1915, 3243; Crater Lake, Klamath Co., Sept. 9, 1916, J. R. Weir, 235.
Anaphalis margaritacea subalpina Gray?—N. slope Mt. Hood,
Aug. 7, 1914, 1673.
181. URoMyYCES ARMERIAE (Schlechtd.) Lev. Ann. Sci. Nat. III, 8:
375. 1847.
Caeoma Armeriae Schlechtd. Fl. Berol. 2: 126. 1824.
ON PLUMBAGINACEAE:
Statice armeria L.—Newport, Lincoln Co., May 16, 1914, C. E.
Owens, 1999, July 18, 1915, 3078.
This species differs from U. Limonzi in the shorter, broader telio-
spores and the short mostly deciduous pedicel. The first collection
mentioned bears aecia accompanied by uredinia, the second, uredinia
and telia only. The rust is abundant on a cliff near the seashore.
So far as we are aware this is the first record of this species in America.
182. Uromyces Beckmanniae sp. nov.
O and I. Pycnia and aecia unknown.
II. Uredinia amphigenous, scattered, elliptical, 0.5-0.8 mm. long,
soon naked, pulverulent, cinnamon brown, ruptured epidermis notice-
able; paraphyses none; urediniospores globoid or broadly ellipsoid,
19-24 by 23-30 yw, wall colorless or pale yellow, 2—2.5 uw thick, finely
verrucose-echinulate, pores 8-10, scattered.
Ill. Telia amphigenous and culmicolous, scattered or crowded,
oblong, 0.4-0.7 mm. across, often confluent to form crusts or lines,
tardily naked, blackish brown; teliospores obovoid or ellipsoid, angular,
20-26 by 29-40 », apex rounded or angular, narrowed below; wall
chestnut brown, 1-2 thick, smooth, but showing distinct longi-
tudinal ridges, apex thickened, 3-6 4, pedicel colorless or slightly
tinted next to the spore, equalling the spore or usually deciduous.
JACKSON: UREDINALES OF OREGON 275
ON POACEAE:
Beckmannia erucaeformis (L.) Host.—Corvallis, Sept. 21, IgII,
1183; south Mary’s River, Sept. 30, 1914, 3144, Oak Creek, July 29,
1915, 3145 type, Aug. 14, 1915, 3026, May, 1916, G. B. Posey.
Evidently the most common rust in Oregon on this host. It differs
from U. Hordei, which is in general a southern form not known on the
Pacific coast, in the larger teliospores which show distinct longi-
tudinal ridges. From U. Jacksonii (cf. 192) this species differs in the
thickened apices of the teliospores. No clue is available as to the
aecial host. The rust is difficult to separate from Puccinia Rhamni
(cf. 159) in the uredinial stage.
183. Uromyces Broprgsk Ell. & Hark., Harkness, Bull. Cal. Acad.
Sel. 1:.28; 1884.
Uromycopsis Brodieae Arth. Result Sci. Congr. Bot. Vienne 345.
1906.
On ALLIACEAE:
Brodiaea sp.—Corvallis, May 4, 1912, F. D. Bailey, 3304, April
25, 1915, 2625.
This opsis-form is evidently common in western Oregon. The rust
is usually found attacking the tips of the leaves early in the spring.
Aecia usually predominate, the telia being inconspicuous and easily
overlooked.
184. UROMYCES CARNEUS (Nees) Hariot, Jour. de Bot. 7: 376. 1893.
Aecidium carneum Nees; Funk. Krypt. Gew. Ficktelgeb. 25: 4.
1818.
Uromyces lapponica Lagerh: Bot. Nat. 1890: 274. 1890.
Uromycopsis lapponica Arth. Result Sci. Congr. Bot. Vienne 345.
1906.
On LEGUMINOSAE:
Astragalus sp.—Austin, Grant Co., Aug. 1915, J. R. Weir, 768.
185. UROMYCES CARYOPHYLLINA (Schrank.) Wint. in Rab. Krypt. FI.
B49. TS8i.
Lycoperdon caryophyllinum Schrank. Baier. Fl. 2: 668. 1789.
Nigredo caryophyllina Arth. N. Am. Flora 7: 246. 1912.
On CARYOPHYLLACEAE :
Dianthus Caryophyllus L.—Portland, Sept. 30, 1912, F. D. Bailey,
1089, Dec. 19, 1912, F. D. Bailey, 1744; Corvallis, Dec. 1910, 3781.
186. UROMYCES FABAE (Pers.) DeBary, Ann. Sci. Nat. IV, 20: 80.
1863.
Uredo Fabae Pers. Neues Mag. Bot. 1: 93. 1794.
Nigredo Fabae Arth. N. Am. Flora 7: 251. 1912.
276 BROOKLYN .BOTANIC GARDEN MEMOIRS
ON LEGUMINOSAE:
Lathyrus obovatus (Torr.) White?-—Sumpter, Baker Co., July 16,
1916, J. R. Weir, 792; Austin, Grant Co., Aug. 1915, J. R. Weir, 197.
Lathyrus oregonensis White—Andrews, Harney Co., Aug. Igot,
Griffiths & Morris (Griffiths, West. Am. Fungi 349a); Spencer Creek,
Klamath Co., July 10, 1903, E. B. Copeland (Syd. Ured. 1764).
Lathyrus pauciflorus Fern.2—Klamath Falls, Klamath Co., Sept.
8, 1916, J. R. Weir, 203.
Lathyrus polyphyllous Nutt.—Mt. Hood, Aug. 31, 1901, E. W. D.
Holway.
Lathyrus sulphureus Brewer—Corvallis, May 9, 1914, 3226;
Ashland, Jackson Co., Sept. 10, 1914, 3320.
Lathyrus sp.—Scotts, 7 mi. N. of Fort Klamath, Klamath Co.,
Sept. 20, 1913, E. P. Meinecke, Cr D 10; Glendale, Douglass Co..,
July 17, 1914, 1506; N. Mt. Hood, Aug. 7, 1914, 1490, Aug. 9, 1914,
1486; Whitewater Forest Station, Aug. 12, 1914, H. P. Barss, 3248;
Garden Home, Multnomah Co., July 20, 1915, 3249.
Vicia americana Muhl.—N. slope Mt. Hood, Aug. 7, 1914, 1489,
Aug. 9, 1914, 1485; Corvallis, Sept. 21, 1914, 1545.
Vicia linearis (Nutt.) Greene—Mary’s Peak, Benton Co., June 20,
1910, 7501; Philomath, June 20, 1910, 1502; Newberg, Yamhill Co.,
April 13, 1914, 1525; Springbrook, Yamhill Co., June 25, 1914, F. D.
Bailey, 3236.
Vicia truncata Nutt.—Bonneville, Multnomah Co., Aug. 10, 1910,
1179; Hood River, May 14, 1914, 1527.
Vicia sp.—Dothan, Douglass Co., Sept. 8, 1914, G. B. Posey,
1550.
187. UROMYCES FALLENS (Desmaz.) Kern, Phytopath. 1: 6. I9ITI.
Uredo fallens Desmaz. Pl. Crypt. 1325. 1843.
Nigredo fallens Arth. N. Am. Flora 7: 254. 1912.
ON LEGUMINOSAE:
Trifolium pratense L.—Mary’s Peak, Benton Co., June, 1910,
3094; Springbrook, Yamhill Co., June 22, 1914, G. B. Posey, 3234;
Corvallis, July 15, 1914, G.. B: Posey; 3003, Oct: 26, Toi4; Geae:
Posey, 1983; Parkdale, Hood River Co., Aug. 5, 1914, 3701; Portland,
Aug. 23, 1915, E. Bartholomew (Barth. Fungi Columb. 4788).
188. UROMYCES HETERODERMUS Syd. Ann. Mycol. 4: 29. 1906.
On LILIACEAE:
Erythronium parviflorum (Wats.) Goodding—Corvallis, April 13,
1912, F. D. Bailey, zrrz.
A short-cycle form not uncommon in the Rocky Mt. and Pacific
coast regions.
JACKSON: UREDINALES OF OREGON Lie
189. Uromyces Hotwayt Lagerh. Hedwigia 28: 108. 1899.
Uromyces Lilia G. W. Clinton; Peck, Ann. Rep. N. Y. State Mus.
27: 103. 1875. (Not U. Lili Kunze. 1873.)
Nigredo Lilu Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
On LILIACEAE:
Lilium parviflorwm (Hook.) Holzinger—Wren, Benton Co., July,
1911, W. E. Lawrence, 1144; Hood River, May 9, 1915, 3044; May
16, 1915, 2061; Bridal Veil, Multnomah Co., May 18, 1915, 2650;
Portland, June 21, 1915, 3060.
190. UROMyCES HyYPERICI-FRONDOsI (Schw.) Arth. Bull. Minn. Acad.
Mat. sel..22:. 55; - 1es3.
Aecidium Hyperici-frondosi Schw. Schr. Nat. Ges. Leipzig 1: 68.
1822. A
Nigredo Hyperici-frondosi Arth. Résult Sci. Bot. Vienne 344.
1906.
ON HYPERICACEAE:
Hypericum Scoulert Hook.—Corvallis, June 24, 1914, F. D. Bailey,
‘ 1628, July 29, 1914, 1476; Hood River, June 20, 1914, 3372.
This species has not been previously reported west of the Missis-
sippi valley.
EOL. UROMYCES INTRICATUS Cooke, Grevillea 7: 3. 1878.
Uromyces Eriogont Ell. & Hark.; Harkness, Bull. Cal. Acad. Sci.
Te 204 L804.
Nigredo iniricata Arth. N. Am. Flora 7: 244. 1912.
ON POLYGONACEAE:
Eriogonum compositum Dougl.—The Dalles, Wasco Co., July 3,
1914, 1300; Hood River, July 22, 1915, 3140.
Eriogonum microthecum Nutt.—Redmond, Crook Co., July 2, 1914,
2537:
Eriogonum stellatum Benth.—Wren, Benton Co., June 26, 1914,
1326; Hilgard, Union Co., July 10, 1914, 1439.
Eniogonum umbellatum Torr.—Mt. Hood, 7,000 ft. elev., Sept. 1
1901, E. W. D. Holway, 6,500 ft., Aug. 9, 1914, 1481, 1493.
Eriogonum vimineum Dougl.—Elgin, Union Co., Aug. 15, 1899,
C. L. Shear (Ell. & Ev. Fungi Columb. 1470).
Enogonum virgatum Benth.—Grant’s Pass, Josephine Co., July
13, 1887, Thomas Howell, from Phanerogamic specimen in the her-
barium Missouri Bot. Gard.
Eriogonum sp.—Waloupi Canyon, Wallow Co., Aug. 18, 1897,
E. P. Sheldon, 8727; Hermiston, Umatilla Co., May 12, 1915, 3039,
3250; Hood River Co., July 22, 1915, 3140.
?
278 BROOKLYN BOTANIC GARDEN MEMOIRS
192. Uromyces JAcksoni Arth. & Fromme, Torreya 15: 260. 1915.
ON POACEAE:
Agrostis Halli Vasey—Corvallis, Sept. 4, 1914, 1576.
Agrostis maritima Lam.—Hood River Co., Aug. 26, 1915, E.
Bartholomew, 5971 (Barth. Fungi Columb. 4992).
Deschampsia caespitosa (L.) Beauv.—Toledo, Lincoln Co., July
19, 1915, 3194.
Deschampsia elongata (Hook.) Munro—Orenco, Washington Co.,
June 13, 1914, 2396; Corvallis, July 6, 1914, 2658, type, July eo,
1914, 1438; Glendale, Douglass Co., Aug. 17, 1914, 1408.
Hordeum jubatum L.—Umatilla, Umatilla Co., July 11, 1914, 1376.
Hordeum nodosum L.—Portland, May 21, 1914, F. D. Bailey, 1583.
In addition to the above collections this species is now recognized
on Agrostis pallens from California and Muhlenbergia Lemmoni from
Arizona and New Mexico. Collections have also been made on
Deschampsia elongata in Washington and on Hordeum nodosum in
Washington and California.
193. URoMyYcES JUNCI (Desmaz.) L. Tul. Ann. Sci. Nat. IV, 2: 146.
1854.
Puccinia Junct Desmaz.:Pl. Crypt. 81. 1825.
Nigredo Junci Arth. N. Am. Flora 7: 238. 1912.
ON CARDUACESE: I.
Arnica cordifolia Hook.—Austin, Grant Co., Aug. 1915, J. R. Weir,
190.
On JuncacEsE: JJ, III.
Juncus balticus Willd.—Redmond, Crook Co., July 2, 1914, 1426;
Umatilla, Umatilla Co., July 11, 1914, 1372; Toledo, Lincoln Co.,
July 19, 1915, 3391.
This species develops its aecia on various Carduaceae. In Europe
cultures have been conducted by various authors, according to Klebahn
(Die Wirtsw. Rostp. 329. 1904), showing that the aecia occur on
Pulicaria dysenterica (Inula dysenterica).
In America Arthur (Mycol. 4: 22. I912,'7:77. 1915) has shown
by culture experiments that aecia occur on Carduus and Ambrosia.
The aecia on Arnica are referred here on morphological grounds.
194. UROMYCES JUNCI-EFFUSI Sydow, Monog. Ured. 2: 290. IgI0.
Puccinia Junct Schw. Trans. Am. Phil. Soc. II. 4: 295. 1832.
(Not P. Junci Desmaz. 1825.)
Uromyces effusus Arth. Jour. Myc. 13: 193. 1907. (Not U.
effusus DeToni. 1888.)
Nigredo Junci-effust Arth. N. Am. Flora 7: 239. I912.
JACKSON: UREDINALES OF OREGON 219
ON JUNCACEAE:
Juncus Bolanderi Engelm.—Ashland, Jackson Co., Sept. 10, 1914,
2523.
Juncus ensifolius Wikstr.—Minum River, Wallowa Co., Aug. 11,
1897, E. P. Sheldon, Aug. 20, 1897, 875za; Corvallis, Aug. 10, I9II,
1188, July 29, 1914, 2522; Philomath, Oct. 28, 1911, 1784, 1185, 1186,
wan. 6, 1914, 7700; Clatsop, Clatsop Co.,, Nov. 7, 1913, 17909; Hood
River Co., Aug. 5, 1914, 2521; Ashland, Jackson Co., Sept. 10, 1914,
2524; Newport, Lincoln Co., July 17, 1915, 3394.
Juncus Mertensianus Bong.—Big Creek, Waldport, Lincoln Co.,
mie. 22, 1915, F. D. Bailey, 3367.
Juncus orthophyllus Cov.—Silver Lake, Lake Co., (?) Aug. 20,
1891, J. B: Lieburg, from Phan. spec. in N. Y. Bot. Gard. 765.
Juncus oxymeris Engelm.—St. Johns, Multnomah Co., July 28,
1902, E. P. Sheldon, from Phanerogamic specimen in National Museum
IIOIQ.
This species has not been connected with any aecial form. From
field observations made by the writer in Oregon it seems probable
that the aecia are to be looked for on Aster.
195. Uromyces LupIni Berk. & Curt. Proc. Am. Acad. 4: 126. 1858.
Nigredo Lupini Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
On LEGUMINOSAE:
Lupinus laxiflorus Dougl.—Garden Home, Multnomah Co., Aug.
1909, 1828, July 20, 1915, 3240.
Lupinus rivularis Dougl.—Springbrook, Yamhill Co., May 14,
1914, F. D. Bailey, 1528.
Lupinus sp.—Mt. Hood, Aug. 31, 1901, E. W. D. Holway, Aug.
9, 1914, 3227; Bonneville, Multnomah Co., Aug. 11, 1910, 1074,
1088; Philomath, May 10, 1914, 3708; Springbrook, Yamhill Co.,
June 22, 1914, F. D. Bailey, 3zzz; Jetty, Lincoln Co., July 19, 1915,
VanGundia, 3731; Newport, Lincoln Co., July 20, 1915, 3264; Hood
River, July 23, 1915, 3020; Eddyville, Lincoln Co., Aug. 9, 1915,
Hoerner, 3180.
196. UROMYCES MEDICAGINIS Pass. in Thiim. Herb. Myc. Oecon. 156.
1874.
Nigredo Medicaginis Arth. N. Am. Flora 7: 256. 1912.
On LEGUMINOSAE:
Medicago lupulina L.—Albany, Linn Co., Aug. 1907, David
Griffiths; Medford, Jackson Co., June 26, 1915, G. B. Posey, 3057.
The aecia of this species in Europe have been shown by Schroeter
(Krypt. Fl. Schl. 3!: 306. 1887) and by Trebaux (Ann. Myc. 10: 74.
1912) to occur on various species of Euphorbia.
280 BROOKLYN BOTANIC GARDEN MEMOIRS
No aecia in America have been found which can be referred to this
species. There is, however, no evidence at present available for be-
lieving the American species different from the European.
197. UROMYCES MINIMUS Davis, Bot. Gaz. 19: 415. 18094.
Nigredo minima Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
ON POACEAE:
Muhlenbergia comata (Thurnb.) Benth.—Wallowa Valley, Wallowa
Co., July 28, 1900, Wm. C. Cusick.
Muhlenbergia racemosa (Michx.) B.S.P.—Wallowa Valley, Wal-
lowa Co., July 28, 1900, Wm. C. Cusick.
198. UROMYCES OBLONGA Vize, Grev. 5: I10. 1877.
Uromyces minor Schrot. Krypt. F. Schles. 3': 310. 1887.
Uromycopsis minor Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
On LEGUMINOSAE:
Trifolium albopurpureum T. & G.—Corvallis, E. R. Lake, 3229.
Trifolium dubium Sibth.—Corvallis, Apr. 10, 1914, F. D. Bailey,
1522; Orenco, Washington Co., June 13, 1914, 3228; Yaquina,
Lincoln Co., July 17, 20, 1915, 3100; Gerlinger, June 22, 1914,4G. Bs
Posey, 3231.
Trifolium eriocephalum Nutt.—Corvallis, July, 1910, 1980.
Trifoium Hallit Howell—Corvallis, June 6, 1899, E. R. Lake,
3272. May 12, 1903; 4. Hl. Post, 42370-
Trifolium microdon H. & A.—Corvallis, May 11, 1907, E. R. Lake,
1408.
Trifolium oliganthum Steud.—Corvallis, May 11, 1914, F. D. Bailey,
Pile ye
Trifolium procumbens L.—Corvallis, June 22, 1901, L. C. M.,
1833, July, 1910, 1176.
Trifolium tridentatum Lindl.—Philomath, June 24, 1914, 1345;
Corvallis, May 28, 1903, E. R. Lake, 3236, May 11, 1914, Fs
Bailey, 3235.
This opsis-form is very common on native Tyvifolium sp. The
original collection by Harkness was reported as occurring on “ Bur
cloves” now considered an error for Trifolium.
199. UROMYCES OCCIDENTALIS Diet. Hedwigia Beibl. 42: 98. 1903.
Nigredo occidentalis Arth. N. Am. Flora 7: 252. 1912.
ON LEGUMINOSAE:
Lupinus sp.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R.
Weir, 248.
200. UROMYCES PERIGYNIUS Halsted, Jour. Myc. 5: 11. 1889.
Nigredo perigynia Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
JACKSON: UREDINALES OF OREGON 281
ON CYPERACEAE:
Carex arthrostachya Olney—Corvallis, July, 1910, rrg7, 1192.
This species is morphologically indistinguishable from P. Asterum
(cf. 69) in all spore stages except in the possession of one-celled telio-
spores. Like that species the aecia occur on Aster and Solidago. The
genetic connection was established by Arthur in repeated experiments
Cour Myc.10: 16. 1904; Mycol.4: 21. 1912, 7:75. 1915,.7: 83.
1915). Fraser (Mycol. 4: 181. 1912) has confirmed Arthur’s results
in part.
The above collection is the only one so far known from the Pacific
coast.
201. UROMYCES PLUMBARIUS Peck, Bot. Gaz. 4: 127. 1906.
Nigredo plumbaria Arth. Résult Sci. Congr. Bot. Vienne 344.
1906.
ON ONAGRACEAE:
Pachylophus marginatus (Nutt.) Rydb.—Snake River, E. Oregon,
June 3, 1901, W. C. Cusick (Phan. spec. 2542).
Pachylophus montanus (Nutt.) A. Nels.—Crooked River, Crook
Co., July 3, 1901, W. C. Cusick (Phan. spec. 2633).
This species is correlated in morphological characters with P.
Epilobu-tetragont (cf. 99).
202. UromMyCcEs POLyGonl! (Pers.) Fuckel, Symb. Myc. 64. 1869.
Puccinia Polygoni Pers. Neues Mag. Bot. 1: 119. 1794.
Nigredo Polygoni Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
ON POLYGONACEAE:
Polygonum aviculare L.—Corvallis, Sept. 4, 1914, 1929; Clatskanie,
Columbia Co., Oct. 6, 1914, F. D. Bailey, 1945; Medford, Jackson
Co., June 20, 1915, G. B. Posey, 3055.
203. Uromyces porosus (Peck) comb. nov.
Aecidium porosum Peck, Bot. Gaz. 3: 37. 1878.
Uromyces coloradensis Ellis & Ev. Erythea 1: 204. 1893.
Uromycopsis porosa Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
ON LEGUMINOSAE:
Vicia americana Muhl.—Orenco, Washington Co., April 23, 1915,
Berror, June'13,.1914, Ill, 37237:
The two collections were made at the same spot on different dates.
The first consists of aecia only and the second of telia only.
204. UROMYCES PROEMINENS (DC.) Pass.; Rabh. Fungi Eur. 1795.
1873.
Uredo proeminens DC. FI.-Fr. 2: 235. 1805.
282 BROOKLYN BOTANIC GARDEN MEMOIRS
Nigredo proeminens Arth. N. Am. Flora 7: 259. Ig12.
ON EUPHORBIACEAE:
Euphorbia glyptosperma Engelm.—Wasco Co., July 23, 1885, W.
N. Suksdorf (from Phan. spec. in N. Y. Bot. Gard.).
Euphorbia oregonensis Millsp.—Horse Creek Canyon, Wallowa
Co., May 20, 1897, E. P. Sheldon, 8715.
205. UROMYCES PUNCTATUS Schrét. Abh. Schles. Ges. 48: 10. 1870.
Nigredo punctata Arth. N. Am. Flora 7: 253. I9g12.
ON LEGUMINOSAE:
Astragalus Purshit Dougl.—Austin, Grant Co., Aug. 1915, J. R.
Weir, 236.
The aecia of this species have been shown by European authors
to occur on Euphorbia cyparissias. No aecial collections have been
made in America.
206. UromyYcEs Scirpt (Cast.) Burrill, Bot. Gaz. 9: 188. 1884.
Uredo Scirpt Cast. Cat. Pl. Mars. 214. 1845.
Nigredo Scirpi Arth. Résult Sci. Congr. Bot. Vienne 344. 1906.
ON CYPERACEAE:
Scirpus paludosus A. Nels.—Waldport, Lincoln Co., Aug. 23,
1915, F. D. Bailey, 3323.
This species was first shown by Dietel (Hedwigia 29: 149. 1890)
to have its aecia on Sium latifolium and Hippurus vulgaris. Other
investigators have added other Umbelliferous hosts to the list.
In America. Arthur (Jour. Myce. 13199... 1907; 14s 172eeeeee
Mycol. 1: 237. 1909) has shown that Cicuta maculata is an aecial
host. Fraser (Mycol. 4: 178. 1912) has confirmed Arthur’s work.
Aecia on other hosts are properly referred here on morphological
grounds. The species can doubtless be separated into a number of
biological forms when more extensive culture work has been conducted.
207. Uromyces SILpui (Burrill) Arth. Jour. Myc. 13: 202. 1907.
Aecidium compositarum Silphii Burrill; DeToni in Sacc. Syll.
Fung. 7: 798. 1888.
Uromyces Junci-tenuis Sydow, Monog. Ured. 2: 289. 1910.
Nigredo Silphti Arth. N. Am. Flora 7: 239. I912.
On JUNCACEAE:
Juncus occidentalis (Cov.) Wieg.—Corvallis, Aug. 10, 1911, 1187,
June 24, 1914, F. D. Bailey, 1387, July 29, 1914, 1445; Philomath,
Jan. 6, 1914, 1108, May 10, 1914, 3393; Hood River, July 24, 1915,
3392.
Arthur (Jour. Myc. 13: 202. 1907; 14:17. 1908) has shown that
this common species has its aecia on Silphium. Using telial material
on J. tenuis from Indiana, West Virginia and Nebraska, five successful
JACKSON: UREDINALES OF OREGON 283
infections of Silphium perfoliatum were obtained, all of which resulted
in the development of pycnia and aecia. The aecia on Silphium have
been collected, so far as known to the writer, only in the Mississippi
Valley from Ohio to Wisconsin, Kansas and Missouri, on three species
of Silphium. The range of the telial collections referred here, how-
ever, is much greater including nearly the entire United States and
Canada except the south Pacific slope. It seems probable that some
plants other than Silphium, at present unrecognized, also serve as
aecial hosts for this species.
208. UROMYCES SOLIDAGINIS (Sommf.) Niessl, Verh. Natur.-Ver.
Brum: T0162. 1872.
Caeoma Solidaginis Sommerf. Suppl. Fl. Lapp. 234. 1826.
Telospora Solidaginis Arth. Résult Sci. Congr. Bot. Vienne 346.
1906.
ON CARDUACEAE:
Solidago sp.—Dufur, Wasco Co., June 30, 1914, 13306.
This is the only micro-Uromyces occurring in both Europe and
America. This species shows a morphological correlation with P.
Asteris (cf. 70).
209. UROMYCES SPRAGUEAE Hark. Bull. Calif. Acad. Sci. 1: 36. 1884.
Uromycopsis Spragueae Arth. Résult Sci. Congr. Bot. Vienne 345.
1906.
ON PORTULACEAE:
Calyptridium roseum Wats.?—Crater Lake, Klamath Co., 7,000 ft.,
Sept. 22, 1913, E. P. Meinecke, CrPkD (2) 11.
Spraguea multiceps Howell—Strawberry Mt., Grant Co., Sept. 2,
1913, W. E. Lawrence, 1177.
210. UROMYCES SUBSTRIATUS Sydow, Ann. Myc. 4: 30. 1906.
Nigredo substriata Arth. N. Am. Flora 7: 253. 1912.
ON LEGUMINOSAE:
Lupinus sp.—Austin, Grant Co., Aug. 1915, J. R. Weir, r5o.
211. UroMyYcES TrRIFOLII (Hedw. f.) Lev. Ann. Sci. Nat. III, 8: 371.
1847.
Puccima Trifolu Hedw.f:; DC. Fl. Fr. 2: 225. 1805.
Nigredo Trifolii Arth. Result Sci. Congr. Bot. Vienne 344. 1906.
On LEGUMINOSAE:
Trifolium hybridum L.—Corvallis, Aug. 1909, 3092; Hood River
Co., May 14, 1914, 3091, Aug. 5, 1914, 3095; Portland, June 11,
1914, 3233; Springbrook, Yamhill Co., June 22, 1914, G. B. Posey,
3103; Garden Home, Multnomah Co., July 15, 1914, F. D. Bailey,
1508; Grant’s Pass, Josephine. Co., Sept. 3, 1916, J. R. Weir, 277.
284 BROOKLYN BOTANIC GARDEN MEMOIRS
212. UROPYXIS SANGUINEA (Peck) Arth. N. Am. Flora 7: 155. 1907.
Uromyces sanguineus Peck, Bot. Gaz. 4: 128. 1879.
Puccinia mirabilissima Peck, Bot. Gaz. 6: 226. 1881.
ON BERBERIDACEAE:
Berberis aquifolium Pursh—Sauvies Island, Multnomah Co., Apr.,
1882, Joseph Howell; Siskiyou, Jackson Co., May 31, 1894, E. W. D.
Holway (Barth. N. Am. Ured. 1400); Corvallis, March 26, 1908,
C. C. Cate, 3362, April 13, 1908, F. L.. Griffin, 3963, Mareh T4,a8os
1137; Philomath, Jan. 1, 1914, 1153; Tualetin, Washington Co.,
March 25, 1915, F. D. Bailey, 2616; Grant’s Pass, Josephine Co.,
Sept. 3; 1920; Jo RK. Weir, 12a.
Berberis pumila Greene, Pokegama, Klamath Co., July 9, 1903,
E. B. Copeland (Sydow, Ured. 1777; Baker, Pacific Coast Fungi
3708). ;
FORM GENERA
213. AECIDIUM ALLENII Clinton in Peck, Rep. N. Y. State Mus. 24:
Og L672.
ON ELAEAGNACEAE:
Lepargyrea canadensis (L.) Greene—Strawberry Mt., Grant Co.,
Sept. 2, 1913, W. E. Lawrence, 1773; Sumpter, Baker Co., June, 1913,
J. R. Weir, 6; Gold Center, July, r914, H. F. Wilson, 1842.
214. AECIDIUM COLLINSIAE Ell. & Ev. Bull. Washb. Lab. 1: 4. 1884.
Aecidium Tonellae D. & H. Erythea 3: 77. 1895.
ON SCROPHULARIACEAE:
Collinsia parviflora Lind|.—Philomath, April 20, 1912, 7169.
This species is evidently an heteroecious form known otherwise
only from Washington on the above host and on C. Rattoni and C.
tenella.
It is apparently distinct from P. Collinsiae P. Henn. (Hedwigia
37: 269. 1898) as stated by Hennings. The aecia of the latter,
judging from the description, arise from a limited mycelium. There
is no evidence of telia in any of the collections of A. Collinsiae ex-
amined. P. Collinsiae has evidently been collected but once and
material is not available for examination.
215. AECIDIUM COLUMBIENSE Ell. & Ev. Erythea 1: 206. 1893.
On CICHORIACEAE:
Mieracium albiflorum Hook.—Hood River, road to Lost Lake,
May 16, 1915, 3245.
Hieracium sp.—Bridal Veil, Multnomah Co., May 18, 1915, 3291.
The aecia arise from a distributed mycelium and are not followed
by any other stage. Puccinia Hieracii may however occur on the
JACKSON: UREDINALES OF OREGON 285
same plants and even on the same leaves. Sydow (Ann. Myce. 1: 326.
1903) described P. sejuncta on such a mixture.
216. AEcIDIUM DELPHINII Barth. Jour. Myc. 8: 173. 1902.
Aecidium Batesianum Barth. in E. & E. Fungi Col. 1901. 1904.
ON RANUNCULACEAE:
Delphinium depauperatum Nutt.—Mary’s Peak, Benton Co., May
i. T9015, 3210.
Delphinium sp.—Corvallis, April 11, 1915, 2615; Redmond,
rook Co., May 15, 1915, 3327:
This species is possibly identical with aecia on other Ranuncu-
laceous hosts referred to P. Clematidis (cf. 85). For purposes of this
list it is retained as a separate form as no cultures have been conducted.
217. AECIDIUM GRAEBNERIANUM Henn. Hedwigia 37: 273. 1808.
Aecidium Alaskanum Trelease, Harr. Alaska Exp. 5: 37. 1904.
ON ORCHIDACEAE:
Limnorchis dilatata (Pursh) Rydb.—Horse Lake, Cascade Mts.,
Aug., 1909, J. C. Bridwell, 3322.
This unconnected Aecidium is doubtless heteroecious since no
other stages have been found following the aecia on any of the col-
lections examined. The species is known otherwise only from Alaska
and‘in the mountains of British Columbia, Montana and California.
218. PERIDERMIUM COLORADENSE (Diet.) Arth. & Kern, Bull. Torrey
Club 33: 426. 1906.
On PINACEAE:
Picea Engelmanii Parry—Whitman National Forest, Wallowa Co.,
Waly, 1973, J. R-Weir, 277:
This species forms large witches’ brooms.
219. PERIDERMIUM ORNAMENTALE Arth. Bull. Torrey Club 28: 665.
19Ol.
On PINACEAE:
Abies concolor (Gord.) Parry
J. R. Weir, 745.
Alies nobilis Lind|.—Larch Mt., Multnomah Co., Aug., 1910, 3293.
White Pine, Baker Co., June, 1913,
220. Uredo Phoradendri sp. nov.
O. Pycnia not seen.
II. Uredinia amphigenous, gregarious, not crowded, spots not
conspicuous, punctate; rounded or slightly elongated, 0.4—0.8 mm.
across, tardily naked, somewhat pulverulent, bright orange, dehiscent
by an elongate or irregular fissure of the epidermis, ruptured epidermis
conspicuous and persistent; peridium membranous, at first hemi-
spherical, remaining closely adherent to the ruptured epidermis,
286 BROOKLYN BOTANIC GARDEN MEMOIRS
made up of colorless isodiametric cells, 14-19 w across, sometimes
somewhat rhomboidal, smooth, wall 1I-1.5 4 thick; urediniospores
ellipsoid or obovoid, 17-33 by 26-32 yw, wall colorless, 1.5—2.5 uw thick,
very closely and minutely echinulate, pores very indistinct, 10-12,
scattered.
III. Telia unknown.
On LORANTHACEAE:
Phoradendron villosum Nutt.—Corvallis, Sept. 21, 1915, C. E.
Owens, 3377 type.
INDEX TO SPECIES
Figures following the names refer to the species numbers in the list.
Synonyms are in italics. New names and combinations are in bold-face type.
Aecidium abundans 59 Mertensiae 136
Alaskana 217 monoicum 135
Allenii 213 Myrtilli 27
Aqutlegiae 85 nilens 45
asperifolit 67 Oenotherae 146
Asterum 69 Ornithogalum 63
Barbarae 135 Palmeri 143
Batesianum 216 Peckii 146
Bellidis 139 Phaceliae 136
Berberidis 151 Pirolae 22
Blasdaleanum 35 porosum 203
carneum 184 pseudo-balsameum 32
Cinerariae 101 punctatum 178
Clematidis 85 Pyrolae 29
columbiense 116, 215 Rhamnzi 159
columnare 5 Sarcobati 167
Collinsiae 214 Solidaginis 69
compositarum Bidentis 140 Sorbi 43
composttarum Silphiu 207 Tonellae 214
conorum Piceae 22 Tussilaginis 100
Delphinii 216 Urticae 174
Dracunculi 173 Violae 176
elatinum 20 Allodus ambigua 61
Fendleri 103 asperior 68
fuscum 57 Calochorti 76
Galit ambigum 61 claytoniata 84
Geranii-maculati 153 commutata 89
Giliae 150 Dichelostemmae 96
Graeberianum 217 Douglasit 97
Grossulariae 109 Giliae 150
Helianthi-mollis 112 Jonestt 122
Holboellit 117 melanconioides 128
Hydrophylli 136 oregonensis 68
Hyperici-frondosi 190 Palmeri 143
Ligulariae 101 Aregma speciosa 33
Majanthae 125 Ascophora disciflora 47
JACKSON: UREDINALES OF OREGON
Bullaria Angelicae 62
Caeoma Armeriae 181
asteratum 69
claytontata 84
erigeronatum 69
Galit 25
hieraciatum 115
nitens 45
occidentalis 13
Solidaginis 208
urticatum 174
Calyptospora columnaris 5
Geoppertiana 5
Chrysomyxa Abietis 6
Pirolae 22
Weirii 6
Coleosporium Adenocaulonis 1
asterum 4
Madiae 2
occidentale 3
Solidaginis 4
Cronartium coleosporiodes 7
Comandrae 8
filamentosum 7
pyriforme 8
Dicaeoma asarinum 66
mesomegalum 130
Earlea speciosa 33
Gymnoconia interstitialis 45
Gymnosporangium asiaticum 40
Betheli 34
Blasdaleanum 35
chinense 40
Haraeanum 40
Harknessianum 36, 39
juniperinum 37
juvenescens 38
Kernianum 39
koreaense 40
Libocedri 35
Nelsoni 41
nootkatensis 42, 43
Sorbi 43
tubulatum 44
Hyalopsora Aspidiotus 9
laeviuscula 10, 23
Polypodii 11
20
Kunkelia nitens 45
Lycoperdon caryophyllinum 185
epiphyllum 100
poculiforme 151
Melampsora albertensis 13, 18
arctica 14
Bigelowii 15, 16
confiuens 16
Lini 17
Medusae 18
occidentalis 18
Piscariae 19
Sparsa 30
Spal
Melampsorella Cerastii 20
elatina 20
Melampsoropsis Piperiana 21
Pyrolae 22
Milesia Polystichii 23
Nigredo aemula 179
caryophyllina 185
Fabae 186
fallens 187
Hyperici-frondosi 190
intricata 191
Junct 193
Junci-effusi 194
Lilit 189
Lupini 195
Medicaginis 196
minima 197
occidentalis 199
perigynia 200
plumbaria 201
Polygoni 202
proeminens 204
punctata 205
Scirpt 206
Silphit 207
substriata 210
Trifolit 211
Nyssopsora echinata 46
Peridermium acicolum 4
balsameum 31, 32
Betheli 8
californicum 2
coloradense 218
conorum Piceae 22
287
288 BROOKLYN BOTANIC GARDEN MEMOIRS
Peridermium filamentosum 7 Puccinia Boisduvaliae 99
Harknessii 7 Calochorti 76
montanum 4 Campanulae 77
ornamentale 219 canaliculata 140
Peckii 27 Caricts-A steris 69
pseudo-balsameum 32 Caricis-Erigerontis 69
pyriforme 8 Caricis-Solidaginis 69
Stalactiforme 7 chasmatis 170
Phragmidium affine 49 Chelonis 78
disciflorum 47 Chrysanthemi 79
imitans 48 Cichorii 80
Ivesiae 49 Cicutae 81
Jonesii 50 cinerea 85
montivagum 51 circaeae 82
occidentale 52 circinans 170
Potentillae 53 Cirsii 83
Rosae-acicularis 54 Cirsit-lanceolatt 87
Rosae-californicae 55 cladophila 111
speciosum 33 Clarkiae 99
Pileolaria brevipes 56 claytoniata 84
Toxicodendri 56 Clematidis 85, 107, 136, 172, 216
Polythelis fusca 57 Clintonii 86
Puccinia Absinthii 58, 90 Cnici 87
abundans 59, 168 Columbiensts 162
Acetosae 60 Collinsiae 214
acuminata 154 Comandrae 88
Agropyri 85 compacta 94
albiperidium 109 commutata 89
alternans 85 conferta 58, 90
ambigua 61, 161 congregata 114
Andropogonis 143 Convolvuli 91
Angelicae 62, 98 coronata 88, 159
anomala 63 Crandallit 59
Antirrhini 64: Crepidis-acuminatae 92
Archangelicae 62 curtipes 163
arnicalis 65 Cyani 93 7
Asari 66 DeBaryana 94
asarina 66 Dentariae 95
asperifolii 67 Dichelostemmae 96
asperior 68 difformis 61
Angelicae 98 dis persa 67
Asteris 70, 208 Douglasii 97
atro-fusca 71 Dracunculi 173
Asterum 69, 200 Ellisi 62, 98
atropunctata I10 Epilobii 175
Bakeriana 98 Epilobii-Fleischeri 175
Balsamorrhizae 72 Epilobii-tetragoni‘99, 175, 201
Barbareae 117, 135 Eriophori 101
Bellidis 139 epiphylla 100
bicolor 73 Eriophyllii 102
Bistortae 74 extensicola 69
Blasdalei 75 Fendleri 103
JACKSON: UREDINALES OF OREGON 289
Puccinia Garrettii 174
Gayophytt 99
Gentianae 105
gemella 104
Giliae 106
glabella 99
glumarum 107
gramints 103, 151
granulispora 108
Grossulariae 109, 144
grumosa IIO
Harknessi 111
Heliantht 112
Helianthi-mollis 112
Hemizoniae 113
Heucherae 114
hieraciata 115
Hieracii 116, 215
Holboellii 117, 135
holcina 118
Hordeti 63
Hydrophylli 160
Hypochoeridis 119
inclusa 83
insperata 120
Iridis 121
Koeleriae 103
Jonesii 122
Junci Desmaz. 193
Junci Schw. 194
Kreageri 59
Ligustici 123
ludibunda 146
luteobasis 123
luxuriosa 124, 167
Madiae 138
Magnusii 109
Majanthae 125
Malvacearum 126
Malvastri 164
Mariae-Wilsont 84
~McClatchieana 127
melanconoides 128, 141
Menthae 129
mesneriana 88
mesomegala 130
micromeriae 131
microsora 132
Millefolii 133
mirabilissima 212
monardellae 134
monoica 135
Puccinia montanensis 136
mutabilis 137
neglecta 107
obliterata 85
obscura 139
obtecta 140
Oenotherae 99
orbicula 120
oregonensis 68
Ortonii 141
Osmorrhizae 148
Oxyriae 142
Palmeri 143
Parkerae 144
patruelis 115
Pattersoniana 145
Peckiana 45
Peckii 146
Pentastemonis 147
Phaceliae 160
Phaseoli var. Taraxaci 169
Phlei-pratensis 151
Pimpinellae 148
Piperi 149
plumbaria 106, 150
Poarum 100
poculiformis 151
Polygoni 202
Polygoni-alpini 152
Polygoni-amphibii 153
porphyrogenita 154
Potentillae 53
Pringsheimiana 109
procera 155
Prunt-spinosae 178
punctata 61, 156, 161
pustulata 88
pygmaea 157
recedens 158
recondita 90
Rhamni 88, 118, 159
Ribis 144
Richardsonti 97
Romanzoffiae 160
rubefaciens 161
rubigo-vera 172
rugosa 162
Saxifragae 163
sejuncla 116, 215
sesstlas 125
Sherardiana 164
Sidalceae 165
290 BROOKLYN BOTANIC GARDEN MEMOIRS
Puccinia similis 58
simplex 63
Sphaeralceae 165
spreta 114
Stipae 166
straminis simplex 63
subnitens 124, 167
Symphoricarpi 168
Syntheridis 177
Taraxaci 169
Tiarellae 114
tomipara 85
tosta luxurians 124
Toumeyi 170
Trautvetteriae 171
Treleasiana 104
trifoliata 148
Trifolit 211
triticina 172
Troximontis 162
untporula 109
universalis 173
Urticae 174
Urticatum 174
Veratri 175
Violae 176
Wulfeniae 177
Pucciniastrum Abieti-Chamaenerii 24
Galil 25
Goodyerae 26
Myrtilli 27
pustulatum 24, 28
Pyrolae 29
sparsum 30
Vacciniorum 27
Roestelia Betheli 34
Harknessiana 36
koreaensis 40
lubulata 44
Telospora Solidaginis 208
Tranzschelia punctata 178
Tremella juniperina 37
Trichobasis Balsamorrhizae 72
glumarum 107
Triphragmium echinatum 46
Uredinopsis Atkinsonit 31
Copelandi 31, 32
Pteridis 32
Uredo Acetosae 60
Angelicae 62
Artemisi 58
Aspidiotus 9
Betae Convolvuli 91
Chamaecyparidis-nutkaensis 42
Chimaphilae 29
Cichorit 80
coleosporiodes 7
confluens 16
Cyant 93
Dentariae 95
Fabae 186
fallens 187
Gentianae 105
glumarum 107
Goodyerae 26
Heucherae 114
Hieracu 116
Tridis 121
laeviuscula 10
miniata Lint 17
nootkatensis 42
Phoradendri 220
Pimpinellae 148
Polygoni Bistortae 74
Polypodit 11
proeminens 204
pustulata 28
quinqueporula 151
Scirpt 206
Solidaginis 4
vagans a Epilobii-tetragont 99
Uromyces Acetosae 60
aemulus 179
amoenus 180
Armeriae 181
atro-fuscus 71
Beckmanniae 182
Brodieae 183
carneus 184
caryophyllinus 185
coloradensis 203
effusus 194
Eriogoni 191
Fabae 186
fallens 187
heterodermus 188
Holwayi 189
Hordei 182
Hyperici-frondosi 190
intricatus I9I
JACKSON: UREDINALES OF OREGON
Uromyces Jacksonii 182, 192
Junci 193
Junci-effusi 194
Junci-tenuis 207
lapponica 184
Lilit 189
Lupini 195
Medicaginis 196
minimus 197
minor 198
oblonga 198 |
occidentalis 199
perigynius 200
plumbarius 175, 201
Polygoni 202
porosus 203
Abies amabilis 32
balsamea 5, 14, 24
concolor 5, 219
Fraseri 5
grandis 5, 20, 24, 25, 31, 32
lasiocarpa 5, 20, 24
magnifica 5
nobilis 31, 219
pectinata 5
Abutilon? sp. 126
Achillea millefolium 133
Adenocaulon bicolor 1
Agoseris glauca 115
laciniata 162
Agropyron dasystachyum 85, 151
lanceolatum 85
spicatum 85, 145
tenerum 136
Agrostis alba 159
alba maritima 159
exarata I5I, 159
foliosa 159
Hallii 192
longiligula 159
maritima 192
microphylla 159
pallens 192
Alopecuris californicus 151
Althaea ficifolia 126
_ rosea 126
Alliaceae 75, 96, 108, 137, 179, 183
Allium acuminatum 75
attenuifolium 75
Uromyces proeminens 204
punctatus 205
sanguineus 212
Scirpi 206
Silphii 207
Solidaginis 208
Spragueae 209
substriatus 210
Toxicodendri 56
Trifolii 211
Uromycopsis Brodieae 183
lapponica 184
minor 198
porosa 203
Spragueae 209
Uropyxis sanguinea 212
Host INDEX
Allium Geyeri 137
nevii 108
validum 179
Amaranthaceae 167
Amelanchier alnifolia 36
florida 35
Amygdalus Persica 178
Anacardiaceae 56
291
Anaphalis margaritacea occidentalis 180
margaritacea subalpina? 180
Anemone Drummondii 94
oregana 57
quinquefolia 57
Angelica atropurpurea 62
genuflexa 62, 98
Lyallii 62
tomentosa 98
Antirrhinum majus 64
Aquilegia formosa 85
truncata 85
Arabis sp. 135
Arbutus Menziesii 30
Arctostaphylos alpina 30
manzanita 30
nevadensis 30
Aristolochiaceae 66
Arnica cordifolia 65, 193
Artemisia dracunculoides 58
frigida 58
ludoviciana 58, 90
rigida 58
tridentata 58
Asarum caudatum 66
292 BROOKLYN BOTANIC GARDEN MEMOIRS
Aspidium Thelepteris 31
Aster acuminatus 69
conspicuus 4, 70
Cuseckii 4
Douglasii 4
folicaeus frondeus 4
Hallii 4
laevis geyeri 4
Astragalus sp. 184
Purshii 205
Athyrium cyclosorum 31
Avena fatua glabrata 151
sativa 151, 159
Balsamorrhiza deltoidea 72
sagittata 72
Beckmannia erucaeformis 151, 182
Berberidaceae 103, 212
Berberis aquifolium 103, 212
nervosa 103
pumila 212
Bidens connata 140
frondosa 140
Boisduvalia densifolia 99
glabella 99
stricta 99
Boraginaceae 136
Brodiaea sp. 183
congesta 96
Bromus carinatus 85
carinatus californicus 85
grandis 85
hordeaceus 85
hordeaceus leptostachys 85
marginatus 85
secalinus 85
tectorum 85
villosus 85
vulgaris 85
Calamagrostis aleutica 157
canadensis 159
hyperborea 159
Calochortus macrocarpus 76
Caltha biflora 104
Calyptridium roseum 209
Campanulaceae 77
Campanula rotundifolia 77
Scouleri 77
Caprifoliaceae 59, 168
Carduaceae 1, 2, 3, 4, 58, 65, 69, 79, 72,
79; 33, 37, 90, 93, IOI, 102, 112, 113,
119, 133, 138, 158, 173, 180, 193,
208
Carex athrostachya 69, 200
Barbarae 174
canescens 69
Deweyana 69
Douglasii 71
festiva 69, 109
Goodenowli 69, 109
Hookeriana 146
Kelloggii 109
laciniata 174
magnifica 109, 174
marcida 69, 115, 173
mertensii 109
mirata 132
monile 109
multicaulis 173
nebraskensis 109, 174
phyllomanica 109
praegracilis 69, 115, 173
pratensis 115
rostrata 174
Rossii 173
scoparia 69
spectabilis 109
stipata 69
straminis 69
subfusca 69
trisperma 69
umbellata 173
utriculata 174
vulgaris 69
Caryophyllaceae 20, 185
Castilleja miniata 7
Centaurea cyanus 93
Cerastium oreophilum 20
viscosum 20
vulgatum 20
Chamaecyparis nootkatensis 42
Chamaenerion angustifolium 24
Chleone nemorosa 78
Chenopodiaceae 167
Chenopodium album 167
Chimaphila occidentalis 29
umbellata 29
Chrysanthemum sinense 79
Cichoriaceae 73, 80, 92, III, 116, 120,
162, 169, 215
Cichorium intybus 80
Cicuta maculata 206
occidentalis 81
JACKSON: UREDINALES OF OREGON
Circaea pacifica 82
Cirsium americanum 83
edule 83
lanceolatum 87
undulatum 83
Clarkia pulchella 99
Claytonia lanceolata 84
Clematis Drummondii 85
Douglasii 85
hirsutissima 85
ligusticifolia 85
Clintonia uniflora 130
borealis 130
Collinsia parviflora 214
Rattoni 214
tenella 214
Comandra umbellata 8, 88
Convallariaceae 130
Convolvulaceae 91
Convolvulus atriplicifolius 91
Cornaceae 154
Cornus canadensis 154
Crataegus Douglasii 34, 35, 44
Crepis gracilis 92
Cruciferae 95, 167
Cydonia vulgaris 35, 40
japonica 35
Cyperaceae 69, 7I, IOI, 109, IIS, 127,
132, 140, 146, 173, 174, 200, 206
Dactylis glomerata 151
Delphinium depauperatum 216
Dentaria tenella 95
Deschampsia caespitosa 192
elongata I51, 192
Dianthus caryophyllus 185
Distichlis spicata 167
Dodecatheon alpinum 141
Hendersonii 141
Jeffreyi 141
latifolium 128
tetrandrum? 141
Elaeagnaceae 213
Elymus arenicola 155
condensatus 85
glaucus 85, 107, 136, I51
triticoides 85
* virginicus 136
Epilobium adenocaulon 28
brevistylum 28
minutum 99
Epilobium paniculatum 99
Ericaceae 21, 30
Erigeron speciosus 69
Eriogonum compositum 191
microthecum 191
stellatum I9gI
umbellatum Igt
vimineum IgI
virgatum IQI
Eriophorum angustifolium 1o1
polystachyon ror
viridi-carinatum 101
Eriophyllum lanatum 102
leucophyllum 102
Eremocarpus setigerus 19
Erythronium parviflorum 188
Euphorbia cyparissias 205
glyptosperma 204
oregonensis 204.
Euphorbiaceae 19, 204
Euthamia occidentalis 69
Festuca confinis 59
elatior 151, 159
idahoensis 59
megalura I51
myuras I51
pacifica 149, 151
rubra 59
subulata 59, 159
Filix fragilis 11
Galium aparine 61, 156
asperrinum 156
boreale 161
triflorum 25, 156
Gaylussacia baccata 27
resinosa 27
Gayophytum ramosissimum 99
Gentiana oregana 105
Gentianaceae 105
Gilia gracilis 150
Godetia amoena 99
Goodyera Menziesii 26
Grindelia sp. 166
Grossulariaceae 16, 109
Helianthus annuus 112
Hemizonia truncata 113
Heuchera micrantha 114
Hieracium albiflorum 116, 215
cinereum 73, 116
293
294 BROOKLYN BOTANIC GARDEN MEMOIRS
Hieracium gracile 116
Scouleri 73, 116
Hierochloe macrophylla 151
Hippurus vulgaris 206
Holcus lanatus 118, 159
Hordeum distichon 151
Gussoneanum 107
jubatum 192
montanense 63
murinum 63
nodosum 63, 192
vulgare 63, 107, 151
Hookera pulchella 96
Hydrophyllaceae 136, 160
Hydrophyllum albifrons 136
capitatum 136
tenuipes 136
Hyperiaceae 190
Hypericum Scouleri 190
Hypochaeris radicata 119
Inula dysenterica 193
Iridaceae 121
Iris tenax 121
Ivesia Baileyi 50
Juncaceae 139, 193, 194, 207
Juncoides parviflorum 139
Juncus balticus 193
Bolanderi 194
ensiformis 194
mertensianus 194
occidentalis 207
orthophyllus 194
oxymeris 194
tenuis 207
Juniperaceae 34, 35, 37, 38, 39; 49 41, 42
Juniperus chinensis 40
communis 41
occidentalis 34, 39, 41
scopulorum 34, 38, 41, 44
sibirica 37
Koeleria cristata 103, 135
Labiatae 129, 131, 134
Lagophylla ramosissima 112
Larix decidua 15
europea 15
occidentalis 15
Lathyrus obovatus 186
oregonensis 186
Lathyrus pauciflorus 186
polyphyllus 186
sulphureus 186
Leguminosae 184, 186, 187, 195, 196,
198, 199, 203, 205, 210, 211
Lepargyrea canadensis 213
Leptaxis Menziesii 114
Leptotaenia dissecta 68
Libocedrus decurrens 35
Ligularia sibirica 101
Ligusticum apiifolium 123
Cusickii 46
purpureum 46
Liliaceae 76, 110, 175, 188, 189
Lilium parviflorum 189
Limnorchis dilatata 217
Linaceae 17
Linum Lewisii 17
usitatissimum 17
Lolium multiflorum 151, 159
perenne 159
subulatum I51
Loranthaceae 220
Lupinus 199, 210
laxiflorus 195
rivularis 195
Lycopsis arvensis 67
Lygodesmia juncea III
Madia citriodora 2
elegans 138
exigua 2
glomerata 2, 138
racemosa 2
ramosa 2
sativa 2
Malaceae 34, 35, 39, 37, 38, 49 43, 44
Malus floribundus 35
Malva rotundifolia 126
Malvaceae 126, 164, 165
Medicago lupulina 196
Mentha canadensis 129
canadensis lanata 129
piperata 129
spicata 129
Mertensia laevigata 136
Micromeria chamissonis 131
Douglassvt 131
Mitella Breweri 114
ovalis 114
Monardella odoratissima 134
villosa 134
JACKSON: UREDINALES OF OREGON 295
Muhlenbergia comata 197 Poa fertilis 100
Lemmoni 192 Kingii 59
racemosa 197 macrantha 100
nemoralis 100
Nabalus hastatus 120 pratensis 100
Navarettia intertexta 106 triflora 100
trivialis 100
Onagraceae 24, 28, 82, 99, 175, 201 Poaceae 59, 63, 67, 85, 103, 107, 118,
Orchidaceae 26, 217 D2A 2 5yelS5, LeO 045s Aone nse
Ornithogalum umbellatum 63 ey UG WES) wWslop,. ier, 7/5 ite
Narbonense 63 192, 197
Osmorrhiza brevipes 148 Polemoniaceae 97, 106, 150
divaricata 148 Polygonaceae 60, 74, 142, 152, 153, 167,
Liebergii 148 IQI, 202
occidentalis 148 Polygonum alpinum 153
Oxycoccus macrocarpus 27 amphibium 153
Oxyria digyna 142 aviculare 202
emersum 153
Pachylophus marginatus 201 imbricatum 74
montanus 201 Muhlenbergii 153
Panicularia elata 159 Newberryi 74
pauciflora 159 pennsylvanicum 153
Pedicularis bracteosa 86 Polypodiaceae 9, 10, II, 23, 31, 32
racemosa 86 Polypodium occidentale 10
Pentstemon sp. 170 Polystichum munitum 10, 23
diffusus 147 Populus acuminata 18
Menziesii 143 alba 12
Peucedanum sp. 81 angustifolia 18
triternata 122 balsamifera 18
Phalaris arundinacea 125 candicans 18
Phacelia heterophylla 136 tremuloides 13
leucophylla 136 trichocarpa 18
Phegopteris Dryopteris 9 Portulaceae 84, 209
Phleum pratense 151 Potentilla aracnoides 53
Phlox condensata 97 blaschkeana 49
diffusa 97 glomerata 49
speciosa 150 gracilis 49
Phoradendron villosum 220 Hippiana 53
Picea canadensis 22 Primulaceae 128, 141
Engelmanii 6, 22, 218 Prunus domestica 178
excelsa 30 Pseudotsuga mucronata 13
mariana 22 Pteridium aquilinum pubescens 32
Pinaceae 4, 5, 6, 7, 8, 13, 15, 20, 22, 24, Ptiloria paniculata 111
31, 32, 218, 219 Puccinellia nuttalliana 85
Pinus contorta 4, 7 Pulicaria dysenterica 193
ponderosa 8 Pyrus baccata 35
pungens 8 communis 35
rigida 4 diversifolia 35
Piscaria setigera 19 ioensis 35
Plumbaginaceae 181 malus 35
Poa ampla 85, 100 rivularis 35
annua 100 sinensis 35, 40
296
Pyrola americana 22
elliptica 22
secunda 22, 29
Pyrolaceae 22, 29
Ranunculaceae 57, 85, 94, 104, I7I, 216
Rhamnaceae 159
Rhamnus purshiana 159
Rhododendron californicum 21
Rhus diversiloba 56
Ribes divaricatum 109
lacustre 16, 109, 144
sanguineum 109
saxosum 16
vallicola 16
Romanzoffia sitchensis 160
Rosa gymnocarpa 33, 51, 55
nutkana 54, 55
pisocarpa 51, 55
Rosaceae 33, 45, 47, 48, 49, 50, 51, 52,
53, 54, 55, 100, 178
Rubiaceae 25, 61, 156, 161
Rubus leucodermis 48
neglectus 48
nigrobaccus 45
parviflorus 52
strigosus 48
vitifolius 45
Rumex acetosella 60
hastatulus 60
paucifolius 152
Salicaceae 12, 14, 15, 16, 18
Salix amygdaloides 15
argophylla 16
Bebbiana 14, 15
corda mackenzieana 15
cordata 15
discolor 14
fendleriana 14
lutea 14
Piperi 15
pseudocordata 15
scouleriana 15, 16
sitchensis 14
Sanicula bipinnata 68
Santalaceae 8, 88
Sarcobatus vermiculatus 124, 167
Saxifraga Marshallii 163
odontoloma 163
Saxifragaceae II4, 144, 163
Scirpus americanus 140
BROOKLYN BOTANIC GARDEN MEMOIRS
Scirpus microcarpus 127
paludosus 206
pungens 140
Scrophulariaceae 7, 64, 78, 86, 143, 147,
170, 177, 214
Secale cereale 67
Senecio aureus IOI
ductaris 101
harfordii 158
hydrophiloides 3
palustre IOI
triangularis 3
Sidalcea oregana 165
virgata 164
Silphium perfoliatum 207
Sitanion californicum 85
glabrum 85
hystrix 107
jubatum 85, 107
velutinum 85
Sium latifolium 206
Solidago sp. 208
canadensis 4
caurina 4
elongata 4
missouriensis 4
rugosa 4
tolmieana 4
Sorbus americana 37
aucuparia 35
hybrida 35
occidentalis 37, 43
sambucifolia 35
Sphaerostigma Boothii 99
dentatum 99
Sporobolus airoides 124
Spraguea multiceps 209
Statice Armeria 181
Stellaria borealis 20
Stenanthium gramineum 110
occidentale 110
Stipa comata 166
Symphoricarpos albus 59, 168
racemosus 59
Synthyris rotundifolia 177
Taraxacum Taraxacum 169
Tellima grandiflora 114
Thalictrum occidentale 85
Tiarella unifoliata 114
Trautvetteria grandis 171
Trifolium albopurpureum 198
JACKSON: UREDINALES OF OREGON
Trifolium dubium 198
eriocephalum 198
Hallii 198
hybridum 211
microdon 198
oliganthum 198
pratense 187
procumbens 198
tridentatum 198
Triticum aestivum 107, 172
compactum 107, I5I
diococcum 107
ovatum 172
vulgare 107, I51, 172
Tsuga canadensis 27
heterophylla 27
Tussilago farfara 100
Umbelliferae 46, 62, 68, 81, 98, 122, 123,
148
Urtica Lyallii 174
Urticaceae 174
Vacciniaceae 5, 27
Vaccinium caespitosum 27
297
Vaccinium canadense 27
macrophyllum 5, 27
myrtilloides 5
ovalifolium 5, 27
ovatum 5
parviflorum 5
pennsylvanicum 5
scoparium 5
Vitis-idaea 5
Valeriana occidentalis 89
Valerianaceae 89
Veratrum californicum 175
viride 175
Vicia americana 186, 203
linearis 186
truncata 186
Viola adunca 176
glabella 176
nephrophylla 176
rugulosa 176
Violaceae 176
Xanthium sp. 140
Zygadenus elegans 110
EVOLUTION BY HYBRIDIZATION
EDWARD C. JEFFREY
Harvard University
Not long ago we were told that the investigation of the problems
of evolution had left behind the inexact if broad phase of study in the
field and had now entered upon the more accurate and satisfactory
stage of quantitative elaboration under laboratory conditions. Leav-
ing aside the question whether whatever exactitude in connection
with this tendency has not been more than offset by a corresponding
narrowness of outlook, it is now quite apposite to inquire if the experi-
mental methods of the physiologist are in reality in the position to
supply final light upon the fundamental problems of evolution. The
judging of living beings by what they do rather than by what they
are, has made notable progress in recent years. We are often told for
example that an organ is the tool of a function and consequently
should be defined by its performance rather than by its organization.
I need not point out the essential fallacy of the physiological definition
of an organ cited above. It obviously breaks down the moment it is
used on any wide range of facts.
Perhaps the most striking illustration of depending overmuch
upon physiological data is supplied in connection with present investi-
gations upon the all important question of the origin of species. It
is practically universally assumed in genetical studies, that the capacity
to breed true under exacting experimental conditions is the most
reliable criterion of good species. It has for example assumed that
breeding results obtained with Oénothera and Drosophila are of funda-
mental importance for the science of biology. By those of us who
have neither been intoxicated with the cult of the evening primrose
nor bowed the knee in the temple of the god of flies, this conclusion
will in general be held undemonstrated. We must obviously know a
good deal more about the antecedents of those forms which have
been raised in recent years to the dignity of veritable biological touch-
stones, before we can admit the validity of the far-reaching con-
clusions drawn from their genetical behavior.
The question of the origin of the species is as much with us today
as it was at the time of the publication of Darwin’s epoch-making
work. Darwin himself ultimately ventured no explanation of the
causes of the changes concerned in the formation of new species, but
298
JEFFREY: EVOLUTION BY HYBRIDIZATION 299
contented himself mainly with pointing out that a general process of
variation has been going on from age to age in matter endowed with
life. He emphasized the fact that the struggle for existence on the
one hand and the selection exercised by environment on the other
provided an important directing influence upon the development of
new species of plants and animals. In recent years a doctrine of old
standing has been revived, namely the hypothesis of mutation. It has
been maintained that new forms or elementary species arise spon-
taneously from formerly existing species. This doctrine has been
particularly advanced by the activities of the Dutch physiologist De
Vries and his disciples in this and other countries. It is a general
observation in connection with the activities of the lower organisms
that in the process of their often extremely active development they
give rise to inhibiting substances. In the case of the common yeast
for example we have the formation of alcohol, which finally, by a high
degree of concentration in fermenting sugary solutions, brings the
activity of the yeast organism to a close. It is of interest to note in
this connection that it is precisely in Holland that scientific opposition
to the mutation hypothesis of De Vries has recently appeared. To
Dr. Lotsy we owe a recent volume on Evolution by Means of Hybrid-
ization, which attacks the mutation hypothesis at its very base
through the contrary hypothesis that all changes in living matter are
due to crossing or hybridization and are not the consequence of
spontaneous internal or mutational phenomena. The author argues
that since hybrids are notoriously variable all variability must be
due to hybridism. This appears to be reasoning in a vicious circle.
Clearly the most definite evidence in regard to hybridism as the cause
of new species should be demanded before the possibility of the
appearance of new types in this manner can be admitted. We
fortunately have extremely good testimony on this subject from the
earlier investigations published by Kerner in Austria and Brainerd in
this country. Kerner in his well-known Pflanzenleben as well as in
an earlier publication in the Oesterreiche Botanische Zeitung has
brought forward much evidence as to the origin of new species as the
result of hybridization in the mountainous regions of eastern central
Europe, where the floras of the Pontic, Mediterranean and Baltic
areas meet. It is impossible within the time at my disposal to make
more than a very brief reference to the results reached by this writer.
He has made it clear that the members of different floras are very
apt indeed to produce new species by hybridization in nature and that
these species, where they are advantageously equipped as compared
with the parent forms, flourish within the same region. In case they
have qualities which enable them to live where the parental species
300 BROOKLYN BOTANIC GARDEN MEMOIRS
are not able to survive successfully they are found to the exclusion of
one or both of these. This last conclusion upsets the conventional
assumption that hybrids can only exist where their originating species
occur side by side. It is clear from the general results of the highly
important systematic and geographical investigations of Kerner that
new species may appear as the result of spontaneous hybridization.
The more recent evidence supplied by the investigations of Brainerd
upon the violets and certain Rosaceae point equally positively in the
same direction. This author has made it clear that a number of
recognized species of Viola and Rubus are in reality hybrids in their
origin. A particularly interesting result reached by Dr. Brainerd is
that these hybrid species may become absolutely fixed in spite of their
mode of origin and respond not only to recognized systematic but also
to genetical criteria for species.
It is too often assumed at the present time that the best criteria
of species are physiological. On this basis the capacity to breed true
in cultures and to produce offspring which comply with the tests ‘of
genetical analysis is regarded as of paramount importance. Since
many known hybrids comply equally with recognized species with
these tests it has become clear that what a plant does in cultures can-
not be accepted as an infallible evidence of its antecedents. Where
physiological criteria fail, we turn to the more constant ones furnished
by morphological characters. It has been recognized for nearly a
century that sterility is often a marked feature of hybrids, especially
when they result from the crossing of somewhat incompatible species.
The causes of incompatibility are apparently unknown as often
species more different in their external characteristics and more
widely separated in geographical range can be crossed with greater
success than those nearly related on the evidence of external features
and geographic coincidence. For example the horse and the zebra
produce fertile hybrids, while the horse and the ass, when crossed, give
rise ordinarily to infertile mules. Similarly our common canoe, yellow
and -black birches, which often grow side by side without hybridizing
all apparently cross with a considerable degree of readiness with the
more isolated shrubby birch of swamps, Betula pumila, according to
the investigations of Jack and Rosendahl in this country.
Hybrids may present in the case of plants a number of interesting
morphological characteristics. For example we frequently find a
high degree of imperfection in their gametic cells, male and female,
with the emphasis of sterility nearly always on the male. This feature
is often so marked that it is impossible to fertilize a hybrid with its
own pollen, even when the ovules present a considerable degree of
fertility. The morphological imperfection in pollen grains can obvi-
JEFFREY: EVOLUTION BY HYBRIDIZATION 301
ously be most easily estimated and varies in proportion from a small
percent to complete sterility. The sterility resulting from hybrid-
ization should not be confused, as sometimes happens, with sterility
arising from purely physiological causes. For example the common
horse radish, Lilium bulbiferum and L. candidum, under ordinary
conditions do not set seed, by reason of the fact that the assimilates
are too strongly determined to the subterranean parts to permit of
the necessary materials being set free for the formation of seeds. It
has been found however that the girdling the top of the subterranean
stem in the horse radish or cutting off the flowering axis in the case of
the lilies, brings about the formation of normal seeds. Similarly very
marked climatic change or subjection to starvation or other extremely
unfavorable physiological conditions results in the degeneracy of
reproductive as well as other parts. Conditions like these are, how-
ever, very easily distinguished from the sterility normally resulting
from hybridization.
Sterility in hybrids is of particular interest from the genetical
standpoint because it more or less completely upsets the expectancy of
Mendelian ratios in cultures of the offspring of species hybrids. This
is doubtless one of the causes why the Mendelians have in general
manifested so little interest in the genetical study of hybrids between
natural species. Obviously however if the crossing of species in nature
is a common cause of the origin of new species this part of the evolu-
tionary field cannot be safely neglected if we are to reach any broad
and permanently valid conclusions as the modus operandi of the origin
of species.
Another feature in the organization of hybrids is the frequent in-
crease in the typical generic chromosome number as a consequence of
crossing. For example we find in the much crossed oriental species of
Chrysanthemum a number of chromosomes in the gametic nuclear
divisions varying from 9 (the normal) to 18, 27, 36, and even 45, in
other words, two, three, four and five times the normal gametic
number. Similarly in another compositaceous genus Dahlia we find
in the species D. coronata sixteen chromosomata in gametophytic
divisions while in the hybrids between D. variabilis and D. coccinea
thirty-two chromosomes have been enumerated. One further example
will point the situation. In the monotypic Liriodendron and in certain
species of Magnolia, nineteen gametophytic chromosomes have been
counted, while in M. soulangeana (suspected of hybrid origin) as well
as M. Yulan and M. grandiflora twice that number or more of chromo-
somes have been observed. If we contrast the situation in these
examples with that presented by the genera Pinus and Lilium, which
are not at all prone to hybridism, we note a curious contrast. In the
302 BROOKLYN BOTANIC GARDEN MEMOIRS
species of the pine or lily, the chromosomes are always of the same
number and do not vary as in the examples cited above, as occurring
in connection with hybridism. Frequently the addition of chromo-
somes under hybrid conditions is not simply a doubling, tripling, etc.,
of the original number but a mere arithmetical addition. Among the
vascular cryptogams and the mosses similar cases of multiplication
of the normal number of chromosomes are frequently found in species
growing in very damp situations or actually in the water and in which
hybridization is accordingly favored.
Having enumerated a few of the morphological characteristics of
hybrids we are now in the position to apply the facts elucidated to the
case of the Oenotheras, which have been brought particularly into
prominence in connection with the mutation hypothesis of De Vries.
In the so-called mutants of Oenothera lamarckiana as well as in that
species itself, we find all the cited stigmata of hybridism as presented
by incompatible species, namely a high degree of sterility, amounting
in some cases to complete impotency, particularly in the male gametic
cells, failure to segregate in accordance with Mendelian ratios and the
multiplication of the number of chromosomes beyond the normal
gametophytic number seven, or sporophytic fourteen. In O. gigas
the gametophytic chromosomes are 14 instead of the normal seven,
while in O. semigigas there are 21 in the sporophyte instead of the
normal fourteen and in O. lata, O. semilata and O. rubricalyx, fifteen.
Similar conditions have been described in many other species of
Oenothera and their so-called mutants. It accordingly appears
abundantly clear when morphological considerations are brought into
court as well as the physiological data derived from experimental
breeding that the genus Oenothera is obviously affected by contamina-
tion through hybridization in its various species and their so-called
mutants. The conclusion may accordingly be drawn that so far as
the genus Oenothera throws light upon the origin of new species at all,
it vouches rather for the multiplication of species as a consequence of
hybridization than for their appearance as a result of the mysterious
process of mutation.
In conclusion we may turn to an objection which has been raised
by De Vries and other mutationists to the interpretation of morpho-
logical sterility as an evidence of hybridization. It has been claimed
that this feature is an accompaniment of mutation. It is most un-
fortunate for the mutationists that a phenomenon so generally recog-
nized as associated with the crossing of species should at the same
time occur in mutating forms. The burden of proof that such forms
are not of hybrid origin distinctly lies with the mutationists. We
have however positive evidence that this is not a possible interpre-
JEFFREY: EVOLUTION BY HYBRIDIZATION 303
tation. In monotypic genera such as Ginkgo, Liriodendron, Calla,
Spathyema, etc., the pollen grains under normal physiological condi-
tions of development are all alike and perfect. In Fig. 1, Plate V,
is shown the pollen of Zannichellia palustris, a species isolated in our
northern North American flora. It is clear that the grains are strik-
ingly uniform and are all well developed. For comparison with the
genus just mentioned which has very few species and consequently
cannot be considered as highly variable, let us take the common
pondweed Potamogeton, of which there are very many species. Fig. 2
shows the situation in the large genus just mentioned. The cells are
not by any means all perfectly developed and are conspicuously char-
acterized by extreme variations in size. It might be maintained on
this basis of the illustrations furnished in Figs. I and 2 alone that
variability in size of pollen grains is associated with the multiplication
of species or in other words with the phenomenon of mutation.
Against this view in the forms under discussion may properly be urged
the fact that many natural hybrids between species of Potamogeton
are known which manifest the usual morphological features of such
forms.
A clearer elucidation of the situation is furnished by the conditions
in large genera, where a number of the species coincide both in geo-
graphical distribution and in the time of flowering. As a first illus-
tration in this connection may be taken the genus Rubus, which has
recently been investigated by Dr. Hoar. Fig. 4 shows the condition
of the contents of the anther in R. villosus. Clearly the pollen varies
greatly in size and perfection of development. A similar condition
has been described by the author just cited in a large number of the
species of Rubus. The general situation might be interpreted in view
of the very numerous and at the same time very variable species of
Rubus as an argument for the correlation of mutation and pollen
sterility. When however the facts in species of the genus, which are
in some manner isolated, are examined quite a different light is thrown
on the subject. Fig. 3, Plate V, illustrates the pollen of R. odoratus,
the flowering raspberry, which opens its blossoms at a considerably
later period than the mass of the species of the genus. Care has been
taken to include as large a number as possible of the grains in the field
of view. It is obvious that the variation in size and frequent defective
development of unisolated species of Rubus, are conspicuous by their
absence. If irregularities in the development of the contents of the
anthers were a feature correlated with mutation in the genus Rubus
then we ought to find it equally present in isolated and unisolated
species. Since that is not the case, the natural inference is that the
sterility present in the pollen of species subject to hybrid contamina-
21
304 BROOKLYN BOTANIC GARDEN MEMOIRS
tion by reason of their coincidence of flowering periods, is actually the
result of previous specific crossing. This view of the matter is strongly .
confirmed by the fact that the investigations of Brainerd and others
on that genus have revealed a large number of natural hybrids.
For a parallel case we may now turn to the genus Ranunculus.
If any of the species which flower in the early summer are examined,
such as R. acris, R. repens, R. aquatilis, R. Cymbalaria, a considerablé
proportion of imperfect pollen is usually present and frequently the
grains vary extremely in size. This situation is shown for R. acris
in Fig. 6, Plate V. Obviously there is a great range of size in the
grains and some are imperfect. This condition is most naturally
interpreted as a consequence of previous hybridization. Fig. 5, Plate
V, illustrates the condition of the pollen in R. rhomboideus, a species
which flowers in the very early spring long before the other species
of the genus have opened their blossoms. The numerous grains
shown in the illustration are clearly well developed and somewhat
uniform in size. In the species under consideration as well as in R.
odoratus, perfection in development of pollen is clearly correlated with
isolation from possibility of contamination with other species.
In view of the facts supplied in the above instances, which might
be almost indefinitely multiplied in representatives of other angio-
spermous families, it appears clear that the obvious interpretation
of pollen sterility is to be adopted, namely as an indication (where
it occurs under normal conditions of growth) not of mutability but
of previous hybridization. The large situation which is so briefly
illustrated by the accompanying photographs, indicates the necessity
of bringing morphological criteria relating to the cytology and develop-
ment of the gametic cells (pollen and embryo sacs) into court, as well
as the data derived from genetical behavior, in speculations in regard
to the origin of species.
It seems clear from the evidence supplied on the systematic and
phytogeographical sides on the one hand and that from the standpoint
of morphology on the other, that the crossing of species in nature is
an extremely common cause of the multiplication of species. It is
further obvious that physiological and genetical criteria must not be
given greater weight than the more reliable ones supplied by actual
history and by morphological structure, in the all-important biological
question of the origin of the species. It is finally apparent that the
genetical status of the Oenotheras is so dubious that they cannot be
brought into court to furnish decisive evidence in favor of the muta-
tion hypothesis of De Vries. It may be added in conclusion that the
multiplication of species by hybridization does not by any means
invalidate the Darwinian hypothesis but merely supplies an additional
BROOKLYN BoTANIC GARDEN Memoirs. VotumeE I, PLATE V.
JEFFREY: EVOLUTION BY HyYBR!DIZATION
ety
JEFFREY: EVOLUTION BY HYBRIDIZATION 305
agency for the formation of species. It appears moreover logically
impossible to regard hybridization as the universal and sole cause of
the appearance of new species, as has been recently maintained by
Lotsy in his Evolution by Means of Hybridization, since the original
species must have come into existence by some other means than by
hybridization. The adaptation of the floral structures of the Angio-
sperms to cross fertilization, emphasized many years ago by the
Austrian botanist Kerner is doubtless of significance in connection
with the ever-increasing volume of evidence for the wide occurrence
of natural hybrids in this large and successful group of seed-plants
which have to so notable a degree furnished the facts for the existing
general biological theories.
DESCRIPTION OF PLATE V
Fic. 1. Pollen of Zannicheliza palustris, showing great uniformity in a species
unable to hybridize. X 400.
Fic. 2. Pollen of Potamogeton diversifolius, showing great diversity of size and
development in pollen of a species subject to hyridization. > 400.
Fic. 3. Pollen of Rubus odoratus, a species which flowers late and consequently
is not subject to crossing. > 400.
Fic. 4. Pollen of Rubus villosus, a species flowering with a number of others
and consequently subject to hybridization. X 400.
Fic. 5. Pollen of Ranunculus rhomboideus, showing uniformity in a species
exempt from crossing by early date of flowering. X 400.
Fic. 6. Pollen of Ranunculus acris, a species flowering at the same time as a
number of others and consequently exposed to hybridization. > 400.
A METHOD OF OBTAINING ABUNDANT SPORULA-
TION IN CULTURES OF MACROSPORIUM
SOLANI E. & M.
L. 0. KUNKEL
Bureau of Plant Industry, U. S. Department of Agriculture
Although the early blight fungus, Macrosporium solani, often
fruits abundantly when growing as a parasite on potato leaves, it
usually does not bear very many spores when grown in pure culture.
Jones (4) reports that some of his cultures when old fruited rather
freely. Jones and Grout (5) state, however, in their technical de-
scription of the organism as Alternaria solani (E. & M.) Sorauer that
it sporulates ‘‘sparsely in pure cultures.”
Enough spores may be obtained by growing it in the ordinary way
on culture media to test its parasitism to the potato plant. Galloway
(3) performed this experiment as early as 1893 and Jones (4) repeated
it a few years later. Nevertheless, the failure to obtain spores in
quantity from pure cultures has made it impossible to perform ex-
tended infection experiments with this important parasite.
The writer found M. solani doing considerable damage in the
potato fields of Aroostook County, Maine, last August, and in the
hope of obtaining a strain of this fungus that would fruit abundantly
in pure cultures, a considerable number of isolations were made.
Cultures were in each case made from single spores. The organism
was isolated from fifty-four different potato plants selected at random
in half a dozen potato fields in the vicinity of Presque Isle, Maine.
All of these single-spore strains were grown on a number of different
culture media, including potato agar, string-bean agar, prune agar,
and glucose agar. The several strains showed considerable differences
in the appearance of their growth in culture, but none of them pro-
duced more than an occasional spore on any of the media tested.
In a former paper the writer (6) has described a method of retarding
the growth of Monilia sitophila (Mont.) Sacc. by lowering the vapor
tension of the atmosphere above pure cultures. It was recalled that
by checking the mycelial growth in this way the fungus could be made
to fruit more abundantly than when grown in a moist atmosphere.
In the hope that this method’ might serve to induce sporulation,
cultures of M. solani were subjected to like treatment. More spores
306
KUNKEL: SPORULATION IN MACROSPORIUM SOLANI 307
were obtained in these cultures than when the atmosphere above the
fungus growth was allowed to remain near saturation. It was found,
however, that this method of treatment would not bring about very
abundant sporulation in the case of the early blight organism. A
number of other methods were tested, and the idea of wounding the
mycelium was finally hit upon. This seems to be the stimulus neces-
sary to bring about abundant sporulation in cultures of M. solant.
It will fruit profusely on any of the media above mentioned if the
mycelium is thoroughly wounded after the culture is two or three days
old and has made a good statt. So long as the mycelium is undisturbed
it grows very vigorously through and over the surface of most culture
media. In these cultures very few conidiophores are produced and
very few spores are developed. Undisturbed cultures often fail to
Fic. 1. A wounded hypha bearing conidiophores of Macrosporium solani,
X 650.
Fic. 2. A wounded hypha of Macrosporium solani showing the production of
conidiophores near the point at which the hypha was cut. X 900.
bear a single spore. If, however, the radiating mycelial strands are
severed at the proper stage in the life of the culture, thousands of
conidiophores, each bearing a spore, will develop from the cells of these
hyphae. The wounding may be accomplished by scraping the surface
of the culture with a sterile scalpel or even with a strong platinum
needle. The more thoroughly it is done, the greater will be the
quantity of spores produced.
Conidiophores arise abundantly near the point at which the
mycelium is broken. Sometimes each consecutive cell of the mycelium
for a considerable distance produces one or more conidiophores. Such
a series of fruiting cells is shown in Fig. 1. At the distal end of some
308 BROOKLYN BOTANIC GARDEN MEMOIRS
of these conidiophores may be seen a scar. This indicates the point
at which the spore was attacked. The production of conidiophores
near a wound is shown in Fig. 2. In Fig. 2 may be seen a portion of a
cell that was killed when the hypha was cut. Each of the photo-
micrographs (Figs. 3 and 4) shows portions of the surface of two
different string-bean agar cultures of the early blight fungus. These
photographs indicate the abundance of spore production when the
mycelium is properly wounded. Many thousands of spores may
Fic. 3. Fruiting culture of Macrosporium solani. This photograph of the
surface of a string-bean agar culture shows the abundance of spore production when
the mycelium is properly wounded. X 15.
thus be obtained from a single Petri dish culture. Spores produced
in this way were sprayed with an atomizer onto potato plants growing
in a greenhouse. They produced good infection not only on the older
leaves but on young leaves as well.
The method of stimulating spore production by wounding was
tested out on each of the single spore strains isolated from the potato
KUNKEL: SPORULATION IN MACROSPORIUM SOLANI 309
fields of Maine. They all responded in like manner, producing spores
in great numbers. The method has also been applied to other Macro-
sporiums that do not fruit readily in culture. M. tomato, Cooke, a
parasite of the tomato and M. daturae, Fautr., a parasite of the jimson
weed, Datura Stramonium L. respond in the same way to the wound-
stimulus. Although M. daturae fruits quite freely without this
stimulus, the number of spores produced in any culture can be greatly
increased by wounding. M. tomato fruits even more sparingly than
Fic. 4. Fruiting culture of Macrosportum solani. This picture shows the
spores more highly magnified and indicates their abundance in a wounded string-
bean agar culture. XX 250.
M. solani in unwounded cultures. When thoroughly wounded it
bears spores in great numbers:
The abundant sporulation of MM. solani in culture makes more
easy the study of its fruiting stages. The successive steps in the
development of conidiophores and spores can easily be observed.
The mature conidiophores are always several-celled. They may
arise singly or in whorls, as is shown in Fig. 2. The typical conidio-
310 BROOKLYN BOTANIC GARDEN MEMOIRS
phore bears a single spore. Successive production of spores on the
same conidiophore, such as has been described by Miyabe (7) for
M. parasiticum Thiim., has not been observed for M. solani. Occa-
sionally the spores are borne in chains of two. This may occur quite
frequently when the atmosphere above the culture is saturated with
water vapor and only a few conidiophores are being produced by a
vigorously growing mycelium. It occurs very seldom or not at all in
cultures that are sporulating abundantly. The occurrence of spores
in chains of two is exceptional for M. solani. The writer has never
observed longer chains and is inclined to the view that Duggar (1)
is correct in leaving this fungus in the genus Macrosporium rather than
to put it into the closely related genus Alternaria as Jones and Grout
(5) have done. Duggar’s objection that the catenulate method of
spore production has not been seen except in artificial culture does
not hold, however, for the writer has more than once observed the
spores in chains of two on the potato leaf. M. daturae when grown
under very moist conditions also produces its spores in chains of two.
This fungus in rare instances even produces spores in chains of three.
Under more normal conditions, however, the spores are always borne
singly. As one finds it growing on its host or observes it in pure cul-
ture, the catenulate method of spore production is exceptional, and
it seems doubtful whether any useful purpose would be served by
transferring it to the genus Alternaria. The writer has examined
many fruiting cultures of the tomato parasite, but has never observed
it producing spores in chains even under very moist conditions.
Besides usually producing their spores singly, both the early
blight fungus and the jimson-weed fungus show other characteristics
which seem to put them with the Macrosporiums. They both possess
a coarser mycelium than is usual for the genus Alternaria. Their
spores are larger and under most conditions are produced in smaller
numbers than is common for an Alternaria. Moreover, their spores
are normally attenuated into a beak similar to that on the spores of
other species of Macrosporium, such as M. catalpae, E. & M., M.
cucumerinum, E. & E., M. caudatum, C. & E., M. concentricum,
Winter, M. brassicae, Berk., M. porri, Ellis, and M. tomato, Cooke.
M. caricinum, one of the four species mentioned by Fries (2) at the
time he established the genus, bears spores which are, according to
his description, attenuated at both ends. No such beaks are to be
observed on spores that are borne in chains.
Since M. solani, M. tomato and M. daturae are parasitic on closely
related plants, they have by some authors been considered identical.
Sorauer (9) seems to have had this notion and Duggar (1) states that
Cy
M. solani “is found not only upon the potato but also upon tomatoes
KUNKEL: SPORULATION IN MACROSPORIUM SOLANI Sal
and upon the jimson-weed (Datura stramonium).’’ By growing the
Macrosporiums obtained from ‘these three hosts side by side in pure
culture it is easy to observe that they are by no means alike. Not
only are they different culturally, but the spores they produce are
quite unlike morphologically. The spores of M. daturae have an
attenuated beak that is very much longer than the beak on the spores
of M. solani or M. tomato. The beak on the spores of M. tomato is
finer than the beak on the spores of the other two species. The
mycelium of M. tomato is also finer than the mycelium of the other
two forms. The spores of M. daturae are larger and the spores of M.
tomato are smaller than those of M. solant. On such media as string-
bean agar and glucose agar the three fungi show wide differences.
M. solani produces a gray felty growth on string-bean agar and
usually colors it red. No such color is to be observed in the case of
M. tomato and M. daturae on the same substratum. The growth of
M. solani on glucose agar is a rusty gray color. Colonies of M.
tomato and M. daturae on this same medium are blue in color. M.
daturae on many different media produces colonies showing marked
zonation, such as is not to be observed with the other two forms.
These three parasites are unlike in so many different ways that the
writer believes they should be considered separate species, rather
than strains of a single species. It is, of course, probable that either
fungus may infect more than one host. Sorauer (8) reports that he
was able to infect tomato leaves with the Macrosporium from the
potato. This does not prove, however, that the three hosts as they
grow in nature are attacked by one and the same fungus. An examina-
tion of the spores of the three fungi when grown side by side in wounded
cultures brings evidence that this is not the case.
The method of obtaining abundant sporulation in cultures of M.
solani here described makes possible more extended infection-experi-
ments than have hitherto been undertaken. It is believed that it
will also be of service in any study of the systematic relationships of
the genus Macrosporium. ‘The principle involved in the response of
the early blight fungus to a wound-stimulus is one well known to
science. Conditions unfavorable to vegetative growth often lead to
fruiting in unprolific plants. The orchardist recognizes this truth when
he prunes his trees or feeds them with fertilizers poor in nitrogen
compounds.
BIBLIOGRAPHY
1. Duggar, B. M. Fungous Diseases of Plants. Ginn and Co., New York, 1909.
2: Fries, E. Syst. Myc. 3: 373. 1829.
3. Galloway, B. T. The Macrosporium Potato Disease. Agric. Science 7: 370-
352. 1603;
312 BROOKLYN BOTANIC GARDEN MEMOIRS
4. Jones, L. R. Potato Blights. Vt. Agric. Exp. Sta. R.g: 88. 1896.
5. Jones, L. R. and Grout, A. J. Notes on Two Species of Alternaria. Bul. Tor.
Club 24: 254-258. 1897.
6. Kunkel, L. O. Physical and Chemical Factors Influencing the Toxicity of
Inorganic Salts to Monilia sitophila (Mont.) Sacc. Bul. Tor. Club 41: 265-
293. I914.
7. Miyabe, K. On the Life History of Macrosporium parasiticum Thiim. Ann.
Bot. 3: 1-26. 1889.
8. Sorauer, P. Auftreten einer dem amerikanischen ‘‘ Early Blight ’”’ entsprech-
enden Krankheit an den deutschen Kartoffeln. Zeits. f. Pflkr. 6: 1-0.
1896.
9. —— Handbuch der Pflanzenkrankheiten. 2: 455, Berlin, 1908.
SYNCHRONISM IN PLANT STRUCTURES
‘JOHN MUIRHEAD MACFARLANE
University of Pennsylvania
In all departments of botanical inquiry it is becoming increasingly
evident that wide observation and exactness of record are indispens-
able, if we are to reach wide and exact conclusions as to plant life.
So the carefully tabulated experiments of Koelreuter, Gaertner,
Herbert, Darwin, Mendel, Vilmorin and others regarding plant cross-
ing during the past century were the appropriate starting points for
the more extended and exact results that are now being secured by
plant breeders.
The characteristic also of exact and correlated behavior on the part
of plant organisms powerfully impressed the writer as he advanced in
his studies of parent and hybrid types, from 1889 onward. Not the
least striking of his results were those bearing on the relation of
plants to environal atmospheric agents or stimuli, such as light, heat
and water supply.' So alike as regards constitutional vigor and
period of blooming as for chemical nature, color, and odor of hybrids,
it was concluded that each detail was more or less exactly between
that of the parents; “while some vary to a greater or less degree from
one or other parent.’’ Impressed, therefore, by such conditions, the
writer has observed closely, during a period of twenty-five years, the
action of those environal agents which we speak of collectively as
- climatic conditions, and the reaction of plant parts to these agents,
with a view to determining how exactly each plant organism is corre-
lated with its environment. This line of inquiry has received con-
siderable attention during the past seventy-five years, under the
term ‘‘phytophenology.’’ But the study, as well as the results se-
cured, have been very largely ignored by botanists, or even ridiculed
by some as yielding no conclusions of value. We would emphatically
assert that few lines of investigation will compare with this if the
studies are prosecuted in exact manner, and are planned so as to cover
a definite field.
The present communication may be suggestive in connection with
future possible developments at such an experimental institution as
the Brooklyn Botanic Garden, which has already had so successful a
history under its able director.
1 Gard. Chron. 93: 753. 1891. Trans. Roy. Soc. Edin. 37: 255. 1892.
313
314 BROOKLYN BOTANIC GARDEN MEMOIRS
Plant variation as due to varying environal stimuli is a phenomenon
witnessed everywhere around us. The difference in size and color
between similar plants growing in shade and sunshine; the difference
in growth and habit between plants exposed to moist rich soil and to
light dry sand; the difference in time of leafing, blooming, and fruiting
between plants situated at lower and higher levels, are facts that are
familiar to all. But the fundamental causes of such differences, as
well as the important conclusions to be drawn, have hitherto been too
much overlooked. Even the records of leafing, the blooming and
fruiting of flowering plants, the shedding of spores by pteridophytic
and bryophytic species, or the conjugation period of algoid and fungoid
types have often been given in most haphazard, or totally misleading,
manner in many of our local floras and manuals.
We desire, therefore, to inquire how far such great seasonal con-
ditions as the above can be reduced to exact limits, and if possible to
ascertain what fundamental principles underlie their expression. The
writer selects first the blooming period of higher plants as a phenome-
non that all can witness and verify to greater or less degree in daily
life. Given that some one locality is chosen where a certain number of
individuals of a species are exposed to as exactly like environal con-
ditions as possible, it may then be asked how nearly synchronous may
the blooming periods be amongst these, and how correctly can we
define these for any region. In illustration, the following may be
cited from amongst many others that the writer has watched 2
Neglecting the wayward skunk-cabbage—that nevertheless can be
reduced to system—the first plant to bloom each season is the silver
maple. This year (1917) hundreds of trees opened many flowers
synchronously on March 11, instead of on the 13th, as is averagely
the case. Furthermore, the opening took place about nine to eleven
A.M. Favored by bright suns the expansion continued upward along
the branches, as is averagely the case, for a period of nine days, and
by this time the earliest flowers were beginning to push out their
green fruits. If we compare now the same trees for previous years
it may be said that during 1912, and as a result of continued snows
and frosts, the unfolding occurred with equal abundance and exactness
on March 17. In 1913 a remarkable record was made. The weeks
of fall weather during 1912 were balmy and mild, and even at times
warm. Asa result many heat units over the average were absorbed
by the trees and caused precocious though unobserved preparation for
spring unfolding in 1913. And here we would emphasize again, con-
trary to views previously expressed by many, that record must be
kept of environal conditions continuously throughout years, if true
* The results recorded are given for West Philadelphia unless otherwise stated.
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 315
results are to be secured as to the action of environmental agents. To
attempt to start the record from such an artificial period as the first of
any year is to set an arbitrary limit to the continuity of changes and
activity in vegetation.
The short-lived snow and slight frosts of late November started
the apparently needful winter maturation of tissues, and_ this
was followed by an almost continuous series of genial days until the
third week of January. So on the morning of January 19 abundant
first flowers expanded on all observed trees, but a cold wave on the
2Ist split the flowering period in half, and only on March 2, with the
advent of a bright day and warm sun did the opening again proceed
until March 10. This striking result had not been paralleled through a
previous period of at least thirty-five years. As a contrast, in 1914 ex-
pansion occurred only on March 16, owing to the frosts and late tem-
peratures occurring throughout February and on to March 15. In
the neighborhood of Wayne, Pa., with an elevation of 475 feet, with
greater. exposure to cold winds and less influenced by the heat of a
great city, the opening did not take place until the 22d of March.
Records like the above that extend over more than twenty years would
suggest that floral expansion is not a somewhat haphazard and irregular
event, but is rather an exact reaction of an organism to definite and
cumulated environal actions or stimuli. If such be true, we should
expect it to extend probably throughout flowering plants as a group.
Partial proof is subjoined.
The red or swamp maple (Acer rubrum) each year succeeds the
silver species in blooming at an average interval of twelve days. This
year, eleven trees, observed in like locality, all opened on the morning
of March 26, while the climax of blooming was reached on the 4th of
April.
According to the valuable statistics secured by Dr. Mackay and
his committee of observers, it may be instructive here to point out
that the same species in Nova Scotia has expanded averagely on May 5,
or 41 days later than in the Philadelphia region.
The white poplar (Populus alba) is of exceptional interest from the
standpoint of the present communication. Staminate catkins an-
nually mature, and lengthen synchronously, amid like environment
on a definite day, and the shedding of abundant pollen proceeds for
one or at most two days. Thereafter the catkins soon shrivel and
within a week have mostly fallen. The average blooming date is
April 7, but this year, stimulated by the warmth of mid-March days
the tassels suddenly lengthened on March 28. Pollen was completely
shed by the 29th, and sidewalks over wide areas were covered with
fallen tassels by the 2d of April. But though a comparatively rare
316 BROOKLYN BOTANIC GARDEN MEMOIRS
tree, the pistillate plant of this dioecious species has been proved to
mature its stigmas synchronously with the shedding of pollen from
the staminate trees. The yearly variations from the above dates
vary according to environal—mainly heat—stimuli. Thus in 1912,
after a cold February and early March, succeeding favorable days
caused complete pollen dispersal on the 27th and in part on the 28th.
During the precocious springtime of 1913, the flowering occurred
suddenly and uniformly on the morning of March 18, while in the suc-
ceeding year like expansion took place on March 31. The Carolina
poplar (Populus deltoides) is like the last species specially abundant
in the staminate trees, rare in the pistillate ones. Its average bloom-
ing period is April 16. In 1913 its behavior was arresting in the sud-
den and exact procedure shown. Here, these trees under similar
environment lengthened their catkins fully and started to disperse
abundant pollen on March 24, between 9 and 10 A.M. of a bright
warm day. The young catkins were almost emptied by 5 P.M.,
only a few of the smaller terminal flowers still retaining a quantity.
By next day, scant remnants could alone be secured. But on that
day four trees which grew on wind-swept and shaded street corners
were found only beginning to dehisce.
In 1914, owing to prolonged cool periods and warmer ones alter-
nating, blooming occurred on April 6. Many catkins matured only
on the 7th. During the present year, the action and reaction of
environal energy and of organismal tissue have been most suggestive.
For stimulated by the bright warm suns of March 31 and April I
some catkins lengthened gradually during these and succeeding days,
and began to discharge pollen extensively on April 4. But rather
low temperatures on that day and those succeeding, accompanied
often by rain, prolonged reaction fitfully until the roth of the month.
So we learn from such statistics, that have frequently been verified
for the above, as well as for other species, that if a sufficient environal
stimulus act quickly and continuously, an extensive synchronous
blossoming may ensue, that is completed within a few hours each year.
On the other hand, if temperature units be more gradually expended,
and specially if such be combined with wetting conditions that prevent
establishment of tissue tension in anthers, pollen discharge may be
prolonged over a considerable period.
The alder (Alnus incana) that averagely lengthens and opens its
catkins on March 24, and that shows crimson papillose stigmatic
surfaces in exactly synchronous manner, was in like state on the 25th,
but in 1913, March 12 was the date. Pollen dispersal is usually com-
pletely effected in three or at most four days, this lengthened period
being due to differences in position and so in time maturation of the
catkins on each twig.
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 317
The above are all shrubby or arborescent forms, and so are more
directly subject to changes of temperature than are species that
perennate by subterranean parts. The energizing factors are less
complex in the former; while in the latter, warmth, moisture, porosity
and chemical composition of the soil, become highly important govern-
ing factors. So the digestion and transfer of the reserve foods in
rhizomes, corms and bulbs is effected more gradually and the flowering
period is usually more prolonged, though the exactly synchronous
unfolding of the first blooms is as striking as in any of the shrubs
or trees. Of our three commonest spring flowers, Hepatica triloba,
Sanguinaria canadensis and Claytonia virginiana, the two first aver-
agely appear in bloom on April 9, and open successive flowers for a
period of 18 days in Hepatica, 10 to 12 days in Sanguinaria and about
25 days in Claytonia. Claytonia appears in bloom on April 12, aver-
agely. But in 1913, our woods showed a sudden unfolding for the
first two on March 18 and of the last on March 20 or 21.
Another herbaceous plant deserves special notice here as illustrating
an interesting phase in synchrony, namely, the dandelion. Like the
English daisy (Bellis perennis), this is a hardy plant which retains
wintergreen leaves and shelters amid close grassy sod. So very slight
changes of temperature in winter will cause both of these to unfold
their earliest flower-heads in apparently regular manner, specially if
growing in sheltered sunny places. But such by no means represents
the first exact growth period for the season, which for the dandelion
occurs averagely on April 23. Then, instead of the scant or occasional
heads of earlier date, our lawns show a sudden yellow coloring by
g A.M. that is continued for almost a month thereafter, as successive
heads expand and as the florets in each successively open. During
1913, the behavior was noteworthy. For lawns were abundantly
yellowed over from January 16 to 20. But all suddenly closed and
were destroyed by frosts that succeeded from January 21 to March 2.
Then came a warm stimulating March, with the result that from the
15th of the month onward dandelions were abundant.
In connection with his graduate class on the Gymnospermia, the
writer became interested from 1898 onward in the behavior of the
Japanese ginkgo tree (G. biloba), at first only with the aim of securing
appropriate material, but as the years passed the phenological relation
became of equal interest. Two large staminate trees grow near the
historic old Hamilton Mansion, adjoining the University Botanic
Garden. These suddenly and synchronously lengthened their catkins
in .1898 on the morning of May 2, and when visited on the succeeding
day few were still polleniferous. This suggested to the writer a
closer study of the subject from the standpoint of individual and species
318 BROOKLYN BOTANIC GARDEN MEMOIRS
behavior. During the previous two years he had secured scant
supplies of good seeds from a large pistillate tree fronting the old
Jones Home at 65th and Callowhill Sts. Throughout the summer of
1898 he examined it from time to time, and noted that the seeds ma-
tured wholly on the southeast side, though no staminate tree was
then known to exist for miles around. In the succeeding spring the
two staminate trees matured between 9 and 10 A.M. on May 5, and
by the 6th were equally free of pollen as before. But the possible
synchronous relation of these to the pistillate tree was now determined.
For, accompanied by one of his students and laden with staminate
branches, a visit to the latter tree was made on the 5th. A ladder
was secured and examination of the small green ovules on the branches
clearly revealed that each was exuding a shining viscous droplet for
pollen entanglement. The staminate branches were hung over the
western side of the tree and then shaken. As summer advanced the
abundant maturing ovules alike supplied wealth of material for study,
and by their structure showed that perfect pollination had been
effected. Continued study of the above trees in succeeding years
showed that synchronous maturation and rapid pollen discharge
annually took place as early as April 20 in 1913 and as late as May 22
in 1904, according to the stimulating amount of heat units, of rain
condition and of soil moisture. In time also as the above statistics
became known, the writer learned of staminate trees on the grounds of
Girard College, and Laurel Hill Cemetery, which doubtless had con-
tributed pollen to the pistillate tree, widely removed from them, during
previous years.
The peach (Prunus persica) deserves attention as being an intro-
duced woody plant, that bears attractive flowers and valuable eco-
nomic fruit. Trees under like environment open averagely on April
22 and then in considerable numbers. Climax of blooming is reached
five days thereafter and within another four days the flowers have all
fallen. But in 1913 a sudden wealth of bloom appeared on April 1.
It need scarcely be added here that a synchronous activity amongst
pollinating bees was a feature of the event.
The common field daisy, that like the dandelion and English daisy,
are all European and introduced weeds, differs markedly nevertheless
from the other two in that it shows no unseasonable flowers through-
out the winter months and does not even unfold as a harbinger of
spring. For averagely a sudden wealth of flower heads expands on
May 24, and for a month thereafter added heads appear in what may
be—did we only know accurately enough—regulated succession.
Almost exactly a month after the last, the first flowers of the intro-
duced moth mullen (Verbascum blattaria) come into bloom on June
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 319
26, and in wonderfully exact and graded succession, with definite time
interval between each, later flowers open along the elongating axis.
But just three days before, or on June 23, the central flowers of the
cymes on our common Indian bean (Catalpa bignonioides) expand, and
for about 16 days thereafter successive blooms open in exact ratio, if
weather conditions are favorable. But a decided retardation may
occur, if cold winds and wetting rains interfere.
We would draw attention now to some cases of synchrony in floral
parts. From the time of Linnaeus onward descriptions of floral clocks
have been frequent, and no matter what value we attach to such, the
very device points to a surprisingly exact time during the twenty-four
hours when the blooms of each species open. But the maturation and
opening of each flower, as well as the behavior of such parts as the
stamens and styles of it, may vary according to the degree of thermic
energy, or lumic energy, or both that act on these. Such variation
seems to explain apparently contradictory results that have been
recorded by different botanists. Thus Kerner’s attractive state-
ments® regarding Silene nutans and its successive maturation of one
row of stamens in each flower, on one evening, of a second row on a
succeeding evening, and of the styles on the third, have called forth
adverse comment along with the statement that no such exact suc-
cession occurs. But first about twelve years ago along the Trafoi
Ravine in the Tirol, and nearly seven years ago by Morgarten Field
in Switzerland, the writer was able to prove that both statements
might apply according to environal conditions. For when the days
and nights in August are warm, with clear sky overhead, the exact
succession noted by Kerner can readily be traced. But if the days,
and even more the nights, be cold, raw and at times accompanied by
rains, then a nearly simultaneous maturation of all ten stamens and
at times even of the styles may occur.
By careful observation, however, during warm dry days we were
able to determine an even more exact and synchronous behavior of the
ten stamens than that noted above. For on the first evening from
about 6 to 7.30 the anthers of the earlier 5 stamens matured in circular
succession with a clear time interval between each, and then were ready
to scatter pollen; while on the succeeding evening, the second five
ripened similarly. Such again calls to mind the time period shown in
maturation of the five anthers on the stamens of the giant cow parsnip
(Heracleum lanatum). It was found even that the dropping of the
anthers in Silene mentioned by Kerner takes place not together, or
irregularly, but in correlated succession if environal stimuli are favor-
Wat Hist.,of Pliv2, prr54:
99
aa
320 BROOKLYN BOTANIC GARDEN MEMOIRS
able. But here again this is duplicated by Heracleum and other
Umbellifers, as well as by many Araliads.
The writer can never forget his first knowledge obtained for the
almost exactly synchronous floral expansion in Oenothera grandiflora.
Two enthusiastic botanical lady friends had cultivated many of these
handsome plants along with numerous other species nearly twenty-
three years ago. He was asked to pay a visit about 7 P.M. on a mid-
July evening. Chairs were set out amongst a group of the Oenotheras
and he was asked to watch and listen. From 7.15 to 7.50 a constant
succession of “‘pfuffs’’ was heard, that indicated the bursting of the
sepals and unfolding of the petals, which rapidly took place before
one’s eye. From the condition where scarcely a flower was open to
begin with, to that seen at 7.50 the change was striking, for now the
plants were gay with large expanded blossoms. Six years afterward
the writer was carried by train from Botzen to Meran in the Tirol
toward 7 P.M. and running for miles by the banks of the Adige River
he witnessed the same synchronous series of events for plants of
Oenothera that had been introduced there.
In connection with genetical studies increasing attention has been
paid during the past fifteen years to the behavior of varieties and
hybrids. But extremely little has been published as to the relative
period of flowering, fruiting or like phenological conditions for each
parent and for the hybrid. But a very wide field for exact observa-
tion is here awaiting study. The writer has drawn attention to some
results and has since accumulated others. Thus, the relative pro-
duction or not in the wild state of hybrid Sarracenias is almost wholly
determined by the synchronous or asynchronous relation of the
flowers. So the scarcity of wild hybrids of S. rubra with other species
is in part due to difference in locality, but in large measure to later
blooming period of that species. Under cultivation by placing plants
in greenhouses of different temperatures a synchronous blooming can
be effected, and such striking hybrids as S. Popei and S. Chelsont
represent the progeny. In such cases then an exact expenditure or
retardation of definite heat units effects a synchrony that in their
natural environment does not exist. The practical application of such
methods in the prosecution of hybridization experiments will ensure
success where failure might otherwise result.
The writer has watched with interest the phenological behavior of a
wild hybrid between Myrica cerifera and M. carolinensis, that his
former graduate student, Dr. Youngken, has described under the
name of M. Macfarlanei. The first of these is a narrow-leaved ever-
green shrub or low tree, that has its northern limit in New Jersey
round the mouth of the Great Egg Harbor River, and there the rather
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 321
dull deep-green leaves are quite green even in mid-April of each suc-
ceeding year alongside other and deciduous vegetation. These ever-
green leaves defoliate in May or early June after the young leaves and
flowers have well expanded. The species moreover invariably occurs
only where facing sea breezes or along the edges of ocean inlets, as
observed along a stretch of the New Jersey coast-line from the mouth
of the Great Egg Harbor River to Cape May Point, and is invariably a
swamp-loving plant.
Myrica carolinensis—the common Waxberry or Bayberry—is a
deciduous species of much wider range and greater hardihood, which
drops its elliptic obovate shiny leaves by the end of November, at
latest, and whose bare twigs throughout the winter show only the
small protruding staminate catkins on one plant and the even smaller
pistillate buds on another. The species occurs often many miles
removed from the influence of ocean winds or brackish water and
nearly always in dry sandy soil.
The hybrid—WM. Macfarlanei—is of semi-evergreen habit. It grows
frequently interspersed with both parents along the area of the New
Jersey coast already named and doubtless will be recognized south-
ward to Florida. Its lanceolate leaves of rather shining aspect remain
green to the end of March, or only become in part brown and fall
during April. More extensive and exact studies made as to the
occasional retention of leaves on low young shoots of M. carolinensis
into mid-winter and the retention of the evergreen leaves on M.
cerifera to an even later date than the writer has indicated, may yet
make our knowledge much more perfect regarding this striking hybrid
and the possible synchrony of floral events, as well as leaf duration on
individuals of the parent and of the hybrid. It might be added that
the hybrid inhabits soil areas which are fairly intermediate between
the swamps of the one species and the dry sandy soil of the other.
If we consider now germination of seeds, equally suggestive syn-
chronous procedure is observed. Only two, amongst many studied,
need be mentioned as having recently been closely examined side by
side with each other. The seeds of the little annual Floerkea pro-
serpinacoides germinated this year in-immense quantities over several
moist shrubby valleys on March 17. The radicle had protruded and
the cotyledons had become swollen on March 25, the first or trifoliate
leaf was uniformly mature on April 7 and the second leaf was unfolding
by April 11. In contrast, subterranean seeds of Amphicarpaea monoica
alongside the above, were still dormant on April 11, but on April 14
many had simultaneously begun to germinate.
Moreover, but in line with all of the above, the annual trans-
formations that occur in our woodlands in spring and throughout
322 BROOKLYN BOTANIC GARDEN MEMOIRS
summer, when compared from year to year, are as exact in relation to
time and energy expenditure as are the flowering periods. Thus, if
we compare the vegetative growth of the Yellow Adder’s Tongue during
1913 with that of the present year the tips of the leaves were simul-
taneously emerging from the ground over wide areas of a valley on
March 27, while this year they appeared on the 11th of April. In
both cases these leaves were 23 inches high 3 days thereafter, and so
comparatively suddenly transformed wide woodland areas from a
bare unclothed aspect into rich showy brown-green verdure.
_ If we consider now a few naked eye details that depend on definite
histological changes, it may be said that botanists are aware that for
any given species of shrub or tree a fairly definite period arrives
when easing and separation of the epidermis along the stem is effected,
after cork formation has replaced it functionally. Some species show
this change in the latter part of the first year, many in the early part
of the second, while others may be delayed until the following autumn
or even later. A more pronounced though related occurrence is seen
annually in the oriental plane. The extensive flakes of old dull-gray
cork start to separate synchronously on the average about June 28
and so reveal the white younger cork underneath with increasing
effect during the next few days, but variation as to date of this event
may occur from year to year according to environal stimuli. Oppor-
tunity has twice occurred for comparing this with the behavior of the
same species round Kew Gardens, England, and there a like change
starts on July 29. This comparative result agrees closely with other
data obtained as to floral maturation.
Closely related again to the above studies is one that has scarcely
been touched in this country, but which has been investigated by
Hoffmann-Ihne in their observations at Giessen. This is an exact
comparison of flowering periods according to longitude and latitude,
particularly the former. Exceptional facilities exist for the prose-
cution of such an inquiry in this country, for were thirty or forty
observational stations established under competent workers, and the
whole correlated at a central office, valuable results would accrue after
a period of ten to twelve years. A feature of interest here is that a
considerable number of plants of the eastern seaboard extend their
range from central or northern Florida to Newfoundland or even
Labrador. One of these which the writer has shortly referred to
elsewhere* is our native pitcher plant (Sarracenia purpurea). In
northern Florida, as for example round Ponce de Leon, it starts to
bloom in the last week of March and continues until April 10. In the
Charleston region, as at Summerville, it is averagely five days later;
* Engler’s Pflanzenreich, vol. 4 (1908), p. 23.
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 323
round Wilmington, N. C., it begins on April 18; in central New
Jersey, it opens on the 20th of May; in northern New York and
Minnesota it blooms from the 14th to the 26th of June. In eastern
central Maine the period is from the 8th to the 20th of July, while in
Labrador—the northern limit of the species—it finishes in mid-August.
Thus a period of fully five months is represented, and a longitudinal
area of about 2,000 miles is covered, in the floral maturation of this
one species. In connection with such records, and probably due to
the more gradual and even expenditure of environal heat stimuli is
the much more extended floral period of species in the cool north than
in the warm south. Thus while the double Crimson Rambler and
Dorothy Perkins roses show floral attractiveness from June 10 to
June 25 averagely round Philadelphia, on eastern Mt. Desert the
period extends from July 15 to August 30.
Were the valuable records, inaugurated in 1892 for Canada by
Mackay, to be linked up with like records from widely distributed
stations in this country, and were all to be correlated with tempera-
ture or thermotactic and moisture or hydrotactic stimuli, as has in
part been done by the Canadian observers, a most valuable foundation
for the establishment of facts regarding the action of definite environal
stimuli would be made.
A very wide field for exact study, still left practically untouched,
is the observation and recording of sporangial ripening and spore
dissemination in pteridophytic and bryophytic genera and species.
One or two references need only be made here.
For years the writer was puzzled to know when spore-dissemination
took place in the sensitive fern (Onoclea sensibilis). Though the
green sporophylls shot up in late July and became greenish-brown in
autumn, opening of the modified pinnae and dissemination of spores
clearly did not take place before winter. Passing through a swampy
patch of this on March 24 of five years ago his clothes became browned
over with the shed product. Subsequent study has shown that this
event occurs averagely on March 25, and in any one patch or locality
with surprising synchronous exactness. Like observations should be
made for Onoclea Struthiopteris.
The sudden and simultaneous elongation of the sporophores and
the subsequent rupture of the sporangia in such hepatics as Pellia
endiviaefolia is familiar to all in mid-April, but we still lack exact day
and hour records through succeeding years for the entire group of
scale mosses.
The predicable manner in which, when fresh horse manure is
placed under bell jars in the now familiar laboratory experiment with
Pilobolus, an abundant crop of the black sporangia is shot forth on a
324 BROOKLYN BOTANIC GARDEN MEMOIRS
morning after a definite number of days of growth, is as indicative for
that fungus as is the previous growth of Mucor on the same medium,
within a shorter period.
What conclusions, it may now be asked, can be drawn from data
such as the few above given?
Time, space, energy and matter are the four great interrelated
phenomena of the world, as of the universe generally. Not a few
physicists now question the existence of the last of these, but inert
and mobile ether particles as focal centers and pathways for “tubes
of energy’’ seem to be helpful—even necessary requirements. For by
their gradual aggregation under increasing condensations of energy we
can explain the origin of the elements, and equally the compounds of
these. But the fundamentally important consideration is how, when,
and to what extent in given times, do definite tubes of energy distrib-
ute themselves.
In the foregoing pages a set of simple facts has been recorded that
any average observer might accumulate. But the real value of many
of them has been overlooked, because we have not fully realized the
significance of the causes that bring them about. For in the past we
have largely viewed biological phenomena as static or semistatic exhi-
bitions of so much material substance. But we have in great measure
failed to realize that matter as such is physically passive or inert, and
that the fundamental moving, transforming, upbuilding, and dis-
integrating agency in all of the above phenomena of phytophenology
consists in definite expenditures of definite amounts of energy along
definite material pathways. Or to use Faraday’s phrase as applied
to inorganic changes, we are dealing with “‘tubes of energy”’ that are
distributed along definite material pathways, at stated climatic periods,
and that are marvelously exact for any one species, or any one organ
of a species.
In the process these tubes of energy are exactly expended so as
to stimulate the inert material particles to take up water, to digest
or metabolize reserve products, to convey the metabolized products
to definite cells or cell walls, to build these up into new material
linkages or combinations, and in the process to effect growth of leaves,
opening of flowers, dehiscence of anthers or of sporangia, maturation of
ovules and extrusion therefrom or from some accessory part at exactly
appropriate time of viscous entangling secretions that strand the
pollen grains, and that in time aid in the germination of these; or
again that start like initial changes in dormant seeds, once so many
units of heat, moisture, and oxygen have cumulated as summated
tubes of energy after a definite period of time; or that develop new
cells or transform older ones, so as to effect shedding of epidermis or
MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 325
peeling of bark on twigs of trees and shrubs at exact time; or that
effect passage of food material from elaborating to storing centers, and
from the latter in turn to young developing organs.
Even such irregular and delayed occurrences as already noted re-
garding the flowering during the present season of the Carolina poplar,
or during 1913 of the silver maple (A. saccharinum) represent fitful
and prolonged results due to weakened or cancelled tubes of energy-
stimulation expended over an extended period, as compared with the
normal succession of events that may be consummated within a few
hours on a definite day or days. So we might summarize as follows
our conclusions drawn from study of phenological and related events:
1. For any one locality, under like environal surroundings, the
average annual period of seed-germination, leaf-formation or unfolding;
first period of blooming, dissemination of pollen, and other responses
by flowering plants, seem to be synchronous often to a day, and even
to certain hours of one day.
2. In monoecious and dioecious flowering plants, under like en-
vironment, all evidence tends to indicate that maturation of compli-
mental floral organs is effected in exactly synchronous relation, and
so abundant pollination usually ensues.
3. A like principle apparently applies to the maturation and
dispersal of spores and organs of conjugation.
4. The behavior of plant hybrids strongly suggests that each is a
blended combination of parental characters as to period of leafing and
defoliation, of blooming and pollination, capacity for climatic resist-
ance and other phenomena. So each shows synchronous behavior in
its organs, that is a mean—all environal factors being considered—
between those of the parents.
5. The principle, advocated by the writer for the past six years,
of environal action and organismal reaction, seems to hold true in the
organic as in the inorganic world, and only needs to be amplified and
demonstrated by increasingly accurate and extended observations on
plants over wide areas.
* 6. In the evolution of all plants and of all plant parts, the funda-
mental and important consideration is the exact distribution of lines
r ‘tubes of energy”’ (Faraday) along otherwise inert material path-
ways; the lines of inflowing energy constituting stimulation actions,
the lines of outflowing energy constituting reactions on the part of the
organism.
7. Such actions and reactions show an optimum, as well as a maxi-
_ mum and minimum of interrelation. The optimum for the indi-
viduals of each species, and for the organs of this, under like environ-
ment, often constitutes a phase-relation that recalls like phase-rela-
tions amongst inorganic bodies.
326 BROOKLYN BOTANIC GARDEN MEMOIRS
8. The material constituents of each plant organism—in them-
selves inert—constitute the gauge or measure by which tubes of in-
flowing or stimulant energy, and outflowing or reaction energy can
best be estimated. The structure of all plant organisms, therefore,
is a cumulated expression of the continued flow of definite tubes of
energy, and the resultant placing, in definite and orderly manner, of
otherwise inert material particles, when moved by such streams of
energy.
THE PROBLEM OF THE IMPORTED PLANT DISEASE
AS ILLUSTRATED BY THE WHITE PINE
BLISTER RUST
HAVEN METCALF
Bureau of Plant Industry, U. S. Department of Agriculture
Within the last few weeks the civilized world has stood aghast at
the stories told by survivors of the devastation wrought by the German
army in its retreat from Northern France. Particularly schrecklich
are the stories and pictures showing the rows of fruit trees cut down
without being utilized for fuel, apparently with no purpose other than
wanton destructiveness. But if we pause to think we realize that the
Germans did not do the worst that they could have done. The fruit
trees are cut down, to be sure, but others can be planted in their places.
We may expect in future years to see the old orchards completely
rehabilitated. But let us suppose that instead of simply cutting
down trees in Europe, any enemy should see fit to leave them standing
but thoroughly incoulated with diseases which would not only destroy
the present stand of trees but would prevent their future profitable
culture in the same territory. To take an extreme example, suppose
that some malignant person or nation should see fit to introduce into
Europe from America, the Colorado beetle, the pear blight, the chest-
nut bark disease, and the citrus canker. Surely the world would stand
aghast at this if its significance was generally realized, because this
would not only destroy or seriously damage the present generation of
plants concerned but would tax the agricultural resources of Europe
with a perpetual burden. No more fiendish blow to the economic
resources of a country could be imagined. Yet this is exactly what
we have allowed foreign nations to do to us without resistance, through
our lax laws regarding the importation of live plants, or rather through
our virtual absence of laws on this subject. We have permitted our
country to be invaded by not one but many—perhaps hundreds—of
diseases and pests which constitute a permanent tax upon our agri-
cultural and forest resources, and up to the present time we have made
scarcely any serious effort to prevent further invasion.
The Bureau of Entomology has issued a large bulletin entitled
‘““A Manual of Dangerous Insects Likely to be Introduced in the
United States Through Importations.”” I understand that this
327
328 BROOKLYN BOTANIC GARDEN MEMOIRS
bulletin lists about 2,700 such insects and that it also lists 130
important insects which have already been introduced into the
United States. If a thorough canvass was made of foreign literature
I have no doubt that as many plant diseases could be located and
described which are likely to be introduced into the United States,
and many of them produce as much devastation as the chestnut bark
disease, the asparagus rust, the potato blight, the citrus canker, or
many other diseases that could be named. Unfortunately, we have
at present no corresponding manual of plant diseases that are likely
to be introduced into the United States.
There has never been a time when the danger from imported
diseases and pests was so great as now. Commerce in living plants
has in recent years extended to the ends of the earth. I have in mind
one nursery company which makes a specialty of novelties from the
Orient. This company is distributing throughout the United States
plant material from all parts of Asia. Most of the things that they
bring in are woody plants, many related to our American species, and
on account of our comparative ignorance of the botany and zoology
of the Orient we have no idea what diseases and pests are coming in
with Oriental material. The San Jose scale, the chestnut blight, and
the citrus canker are only a part of those that have come in already.
Not only is commerce being carried on with countries from which
hitherto there have been only scattering importations of live plant
material, but material brought in now is much more miscellaneous and
reaches this country in a much shorter time. There is at present a
limited amount of port inspection but too limited to be efficient and
the canker diseases and many insects can not be detected by any
sort of inspection. The roots of plants imported with earth about
them can not even be inspected and such plants constitute a par-
ticularly dangerous class of imported material.
The white pine blister rust (caused by Cronartium ribicola Fischer),
which I am here considering as a fairly typical example of the imported
disease, has long been known in Europe. It apparently originated in
Asia and spread in Europe upon Pinus cembra. When the American
white pine (Pinus strobus) was introduced into Europe it proved sub-
ject to the disease. The first authenticated record of importation of
white pine transplants from Europe to the United States dates back
only to 1899. From that time until prohibited by law such importa-
tion was extensive, as such transplants could be imported more cheaply
than they could be grown in America. There was, furthermore, a
prevalent belief among nurserymen that white pine seedlings could
not be successfully grown in America, a belief which has since been
proved erroneous.
METCALF: WHITE PINE BLISTER RUST 329
The disease was first positively reported in America in 1906, on
Ribes. No notice appears to have been taken of this warning. In
1909 enormous quantities of diseased pine nursery stock were im-
ported. Probably 95 percent of all diseased seedlings imported into
America came from a single nursery, that of J. Hein’s Sohne at Halsten-
bek, Germany. This nursery, on account of its use of Rabes hedges,
was curiously well adapted to distribute the disease. In June, 1909, a
meeting was held in New York City of pathologists and foresters of
New England and the Middle Atlantic States, at which a further
alarm was definitely sounded. With one exception, all states repre-
sented discouraged importation of white pine from that time, but
commercial nurseries continued to import extensively until such
importation was made illegal in 1912. Unfortunately, no studies of
the white pine blister rust have been made in Europe by any American
investigator, but if European accounts of the behavior of the disease
can be trusted, the disease has apparently spread more rapidly and
with greater virulence in New England than it did in Europe. Prob-
ably the new climatic and host relations are more favorable to the
disease. In any case the problem of invasion presented by this
disease makes an interesting study. The black currants, especially
the cultivated varieties, are particularly subject to the disease and in
areas of scattering infection are reliable indicators of its presence.
It was hoped by many that the disease might prove to be only one of
nursery stock and reproduction, but at several points in New England,
New York and Minnesota, it is attacking large trees. On Ribes the
disease was in 1916 generally prevalent throughout New England,
which means that the actual infection of pine is much more general
than is obvious at present. Inspection of nursery stock for blister
rust is largely futile since the rust often incubates in pine tissue for
many years before becoming apparent by distorting growth or fruiting.
According to Ravn this incubation period may be as long as twenty
years.
The control of the disease in America presents three separate
problems:
First. West of the Mississippi River. In this territory the disease
is not known to occur, but undoubtedly has been shipped in on nursery
stock of either pine or Ribes. If it has not been carried into this
territory on nursery stock already, there is little possibility of its ever
getting in by natural means. During the coming season an extensive
survey will be made of these states to determine whether the disease
_is or is not present. If the disease should once become established
under western forest conditions, its control would be hopeless. All
5-needle pines of this area, including the very valuable sugar pine and
330 BROOKLYN BOTANIC GARDEN MEMOIRS
western white pine, are subject to the disease, and wild Ribes of many
species are abundant. No species or variety of Ribes yet tested is
immune to the disease.
Second. From the Mississippi River to the Hudson River. There
is an area about 30 miles square in Minnesota and Wisconsin north-
west of St. Paul which is now known to be heavily infected. Probably
more infection will be found in Minnesota. In Michigan, Indiana,
Ohio, Pennsylvania, and New Jersey the disease has been found in a
few nurseries and plantations and is believed to have been eradicated
at these points. In New York west of the Hudson River it has been
found in both nurseries and plantations and largely eradicated, but
on account of the extensive planting of pine nursery stock in the
Adirondacks heavy infections are to be anticipated there. _In general,
the commercial currant-growing sections, such as the region from
Rochester to Buffalo, may be expected to soon show general infection.
In Canada the Niagara Peninsula is already generally infected, and
at least scattering infections occur elsewhere in Ontario. This infec-
tion is of course a serious menace to Michigan.
Third. East of the Hudson River. Here infection is so general
that the only hope of successful growing of white pine in the future
lies in the elimination of the alternate host of the disease; namely,
Ribes. Whether such elimination can be made at a sufficiently low
cost to be profitable remains to be seen. Probably in localities where
Ribes occur sparsely, as in Connecticut and Rhode Island, a great
deal can be accomplished. However, in many sections of rough
country, where wild Ribes are too prevalent to be profitably eradicated,
white pine growing may be expected to become impracticable.
Throughout any section where the blister rust becomes prevalent, the
effect is to make the white pine a cultivated plant; that is, it can not
be profitably grown, or perhaps not grown at all, unless the ground is
kept free from Ribes by artificial means.
There are certain difficulties which stand in the way of any general
campaign of disease control which involves wholesale eradication of
diseased and susceptible plants. It remains to be seen whether these
difficulties are or are not insurmountable.
1. There is in the United States no central authority to act in
any matter involving destruction of diseased plant material or pre-
cautionary destruction of that which is not diseased. Whatever
destruction is undertaken must be undertaken in each state under the
separate legal authority of that state. The state laws are not uni-
form. In some states they are adequate and well supported by public
sentiment, in others they are wholly inadequate and apply only to
special cases. In general there are few states in which the laws are
METCALF: WHITE PINE BLISTER RUST a5)
not so flexible but that a single unconvinced or cantankerous indi-
vidual can nullify the work of an entire community.
2. The laws governing plant eradication are administered in
different states by various officers, but in the majority of states by the
state nursery inspector, who is nearly always an entomologist. Some
of these entomologists are thoroughly trained in plant diseases and
fully appreciate their significance. Others have little knowledge of
them and less interest. All of these officers are overworked already.
In spite of the activities of plant pathologists, there is profound popular
ignorance as to the nature and significance of plant diseases and
especially of the dangerous qualities of newly imported diseases. The
general public is far better informed regarding “‘bugs”’ than regarding
fungi and as a matter of fact the average man considers that plant
diseases are caused by “bugs.”
3. In the case of the blister rust, there is no single interest or
centralized affiliation of interests whose securities are menaced by the
disease. The white pine industry is diffused over a wide territory
and in the hands of many separate individuals and organizations.
What is everybody’s business is likely to be nobody’s business. If
the white pine industry, like the redwood industry, for example, or
like the citrus industry, were in the hands of a few people or consti-
tuted the dominant business in certain areas, the control of the disease
would be much simpler.
4. Up to the present time there has been no adequate quarantine
against the disease, either state or national. This difficulty, however,
will shortly cease to exist as many states have recently declared
quarantines and on June 1, possibly earlier, an adequate national
quarantine will go into effect.
5. Finally, we have a very serious consideration which is applicable
to all undertakings at the present time. The nation is at war. The
young men who would ordinarily be employed in an eradication
campaign will soon be drawn away into military work or into the
various lines of industry which bear directly upon the conduct of the
war. The majority of the persons employed in this particular eradica-
tion campaign can only be employed from the middle of April to the
first of November. Necessarily, men will be loath to accept such
temporary employment when they can in other lines secure permanent
employment at an equally high or higher wage. In any case, the
work if successfully prosecuted or indeed if prosecuted at all will
involve much larger expenditures for wages than in normal times.
What then is the outlook for the control of the white pine blister
rust? It may be expected that the future course of the disease will
be much like that of the gypsy and browntail moths; that is, the dis-
—
352 BROOKLYN BOTANIC GARDEN MEMOIRS
ease can be controlled for a long period vf years in those localities
where infection is not) general but in areas of general infection the
control will be only local and the efficiency of this local control will
largely depend upon whether the white pine as a crop-is of sufficient
value to sustain the added expense of the eradication of Ribes. It
is to be hoped that any infections found west of the Mississippi River.
will be scattering and small, fer,-as has already been stated, if the
disease once becomes established under western forest conditions
its control even on a local basis -will be absolutely out of the question.
Two facts are always in favor of the control of this disease: (1) The
disease has two hosts and can not pass from pine to pine. (2) The
disease is a comparatively slow one; that is,.slow as compared with
such a disease as the chestnut blight. If at any future time in a
completely infected locality the increasing value of the white pine or a
change in industrial conditions makes local control profitable such
control can be undertaken regardless of the prevalence of the disease
at the time, since wherever Ribes can be thoroughly eradicated healthy
pine stock can be grown and will not take the disease from the already
diseased pines. :
The entire blister-rust problem is, however, but one phase of a
larger problem, which may be stated as follows: does free trade in
plant diseases and insect pests pay? Is it an economically sound
national policy? Is the entire importing nursery business worth as
much to the country as the damage which it has already caused?
Not a single plant disease or insect pest that has once become estab-
lished in this country has been eradicated or, in the present state of
knowledge, is ever likely to be. No matter how well controlled, it
remains in every case a permanent tax against our economic resources.
Even if we succeed in controlling the white pine blister rust we may
be absolutely certain that other.diseases and pests are being intro-
duced which will’ be just as serious, for we. know definitely that the
undesirable plant immigrants are not yet all here. It is much more
important to safeguard the country against further invasions of this
kind than to control this or any other disease or pest that has already
been carelessly permitted to establish itself.
It isa matter of common-knowledge, which I scarcely need to repeat
here, that the countries of Europe, and even ends of the earth like
Tasmania and South Africa, have long sincé protected themselves
against the importation of diseases and pests either by prohibition of
entry of nursery stock, or by exclusion of large classes of such stock.
The United States is far behind in this matter.
The future danger is far greater-than the present. The most
dangerous class of nursery stock is the ornamental trees “and shrubs,
VOLUME I, PLATE VI.
BROOKLYN BoTANiC GARDEN Memoirs.
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BROOKLYN BOTANIC GARDEN MEMOIRS. VoLuME J, PLATE VII.
METCALF: WHITE PINE BLISTER RUST
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METCALF: WHITE PINE BLISTER RUST 329
on account of the great number of species represented, and the widely
various parts of the earth from which such stock comes. No section
of the country is likely to suffer more from this source than Long
Island and the general vicinity of New York City. May I express
the hope that this Botanic Garden, already a leader in local and
national sentiment in horticultural and botanical affairs, will take a
position in this matter which will help to bring about the suppression
of this very dangerous traffic?
EXPLANATION OF PLATES VI AND VII
PLATE VI. A native white pine (Pinus strobus) in thick stand, completely girdled
by a blister rust canker. Kittery Point, Me. (Photograph by Mr. W. S. Carpenter,
of the New York State Conservation Commission.)
PLATE VII. A young native white pine (Pinus strobus), completely girdled and
showing several infections of blister rust on trunk and branches. Kittery Point, Me.
(Photograph by Mr. W. S. Carpenter, of the New York State Conservation Com-
mission. )
THE ROSY-SPORED AGARICS OF NORTH AMERICA
WILLIAM A. MURRILL
New York Botanical Garden
This subtribe of the gill-fungi is characterized by rosy or rosy-
ochraceous spores and is well represented by the common and widely
distributed species, Pluteus cervinus. The North American species
may be grouped in ten genera, distinguished by the following key:
Subtribe PLUTEANAE
Pileus irregular, dimidiate or resupinate. I. CLAUDOPUS.
Pileus regular, sometimes eccentric in Pleuropus.
Volva and annulus wanting.
Stipe cartilaginous.
Margin of pileus incurved when young.
-
Lamellae decurrent. 2. ECCILIA.
Lamellae adnate or adnexed. 3. LEPTONIELLA.
Margin of pileus straight and appressed when young;
lamellae free or adnexed. 4. NOLANEA.
Stipe fleshy.
Lamellae decurrent, rarely varying to adnate. 5. PLEUROPUS.
Lamellae sinuate or adnexed. :
Spores not angular, rosy-ochraceous in mass. 6. LEPISTA.
Spores angular, rose-colored in mass. 7. ENTOLOMA.
Lamellae free. 8. PLUTEUS.
Volva wanting, annulus present. g. CHAMAEOTA.
Volva present, annulus wanting. 10. VOLVARIOPSIS.
A few of these generic names may be unfamiliar to you, such as
Leptoniella for Leptonia, Pleuropus for Clitopilus, and Volvariopsis
for Volvaria, but these changes have been required by the rules of
nomenclature.
The time at my disposal will not permit more than a hasty sum-
mary of the North American species belonging to these genera. A
fuller treatment may be found in North American Flora, volume 10,
part 2, to be issued shortly.
1. CLAUDOPUS
Claudopus nidulans, the best known species, occurs throughout
Canada and the United States; C. avellaneus is known from Oregon
only; and there are no species reported from tropical North America.
334
MURRILL: ROSY-SPORED AGARICS 335
The other species of the genus occur in the eastern United States,
from the Atlantic seaboard to the Rocky Mountains.
2. XCCILIA
The best known species is F. atrides, of the eastern United States.
Three species are confined to the Pacific coast, and four to tropical
North America.
3. LEPTONIELLA
Leptoniella serrulata, characterized by the black, serrulate edges
of the lamellae, is the best known species. Four are confined to the
Pacific coast and seven to tropical North America. The rest occur in
the eastern United States.
4. NOLANEA
The two best known species are N. conica and N. mammosa, the
latter distributed throughout temperate North America but not
generally recognized by mycologists. One species is confined to the
Pacific coast and three to tropical North America.
5. PLEUROPUS
This genus contains many edible species, among them P. prunulus,
P. orcellus, and P. abortivus, the last readily distinguished by the
peculiar aborted hymenophores. Two species are confined to the
Pacific coast and one to tropical North America.
6. LEPISTA
One of the best edible species we have is L. personata, better known
as Tricholoma personatum. This species and L. tarda occur through-
out temperate North America. Two other species are confined to the
eastern United States, and there are none known from tropical North
America.
7. ENTOLOMA
This also is a temperate genus, the only species (E. Murraii)
reaching tropical North America being found in the high mountains
of Jamaica. Four species are confined to the Pacific coast. The
best known species in the eastern United States are: E. strictius, E.
Grayanum, E. sericeum, E. rhodopolium, E. Murrati, and E. salmoneum.
Because of the very poisonous European species, E. lividwm, the mem-
bers of this genus have been largely avoided by mycophagists.
23
336 BROOKLYN BOTANIC GARDEN MEMOIRS
8. PLUTEUS
The best known species are P. leoninus and P. cervinus, which are
widely distributed. Five species are confined to the Pacific coast and
fifteen to tropical North America.
9. CHAMAEOTA
Only one species, C. mammillata, has been known in this country
and this only from Michigan.
10. VOLVARIOPSIS
The species of this genus are apt to be widely distributed, as is
the case with many fungi which inhabit manure. The best known
species are probably V. bombycina, V. speciosa, V. volvacea, and V.
pusilla. Four species are confined to tropical North America.
GENERAL SUMMARY
Genera Oe eae need New Species Total
GIONGOP US ae ie 4 4 I 9
CCUAG enc eee eee I 15 9 25
IEA MN LTAN OS @ Oaee aid © AEC I 28 14 43
INOUDI CU erate ow ecrs 2. crcharefinete I 16 II . 2s
IPICULO PUSIas ee ns eee 2 22 5 30
UGE DUStCIL Seite ech ee eee 3 I fo) 4
ENLOLGING sre aces ck Gls hare 2 27 34 63
PIULEUS a ira eke Tame 2 25 30 57
Chamocolin rere ee fo) I I 2
WCU A OSU 5 5 4 bib bob NODE 6 13 2 21
LOG Nene setter ieee 22 153 107 282
THE CYTOLOGICAL STRUCTURE OF BOTRYORHIZA
HIPPOCRATEAE
EDGAR W. OLIVE
Brooklyn Botanic Garden
This species of rust, occurring on the host Hippocratea volubilis
L., was first described by the author, in collaboration with Professor
H. H. Whetzel, from material collected in Porto Rico in 1916.!_ It is
there recorded as a somewhat peculiar form, though somewhat like a
lepto-Uromyces, with only one spore form in its life-cycle. The fol-
lowing diagnosis is there published:? ““O. Pycnia wanting (probably
not formed). III. Telia mostly hypophyllous but sometimes amphi-
genous or caulicolous, generally from a localized mycelium, sometimes
from a systemic invasion affecting entire young shoots; localized sori
densely crowded in more or less orbicular or irregularly shaped,
somewhat hypertrophied pulvinate areas, I mm.—I cm. or more
across, the affected areas yellowish when young, when older becoming
whitish due to the germination of the spores; in older leaves often
killing affected spots, which turn brown, the resultant rounded,
swollen dead areas then bearing a striking resemblance to certain
insect galls.
“Telia pulverulent, erumpent, from a definite, superficial, ure-
dinoid hymenium which arises just under the epidermis, without
peridium; teliospores uninucleate, borne singly at the end of pedicels
which arise from a binucleate mycelium 13-14 by 18-24 y, thin-walled,
oval, with a rounded apical protuberance, germinating apically at
maturity to produce each a long, cross-septate basidium (promy-
celium) bearing 4 basidiospores (sporidia), similar in shape to the
teliospores and 8 by II-I2 yu.
“Vegetative mycelium composed of coarse intercellular hyphae,
made up of binucleated cells, some of which send large botryose, or
irregularly shaped, haustoria into adjacent cells.”’
The generic name, Botryorhiza, is, in fact, derived from the botryose
character of the haustoria, a striking feature which, so far as I am
aware, is possessed rarely if at all by other rusts. It is, however,
1 Endophyllum-like rusts of Porto Rico. Amer. Journ. Botany. 4: 44-52. fis.
I-3. 1917.
a Gr pr Ave
337
338 BROOKLYN BOTANIC GARDEN MEMOIRS
pointed out in the above mentioned paper, that certain smuts (as
Doassantia deformans, e. g.) also possess botryose haustoria.’
The 18 figures in Plate VIII show fairly clearly the salient features
of the cytological structure and development of Botryorhiza, with the
exception of the sexual fusions, which have not as yet been found.
Undoubtedly, however, these fusions and the consequent transition
from the uninucleate condition initiated in the germinating pro-
mycelium to the binucleate condition prevalent in the vegetative
mycelium, must take place early in the development of the latter.
Two hypophyllous sori are shown in Fig. 1, the one at the right
a very young one pushing through a stoma. The mycelium and
hymenial hyphae are seen to be composed of binucleate cells, as is
also the case of the young spores. Two of the peculiar botryose haus-
toria are shown at the lower portion of the figure, nearly filling the
host cells. Fig. 2 shows a portion of the coarse, branched, inter-
cellular mycelium, with some its binucleate cells. This mycelium
varies from about 5 to 7 «in diameter. One cell is shown with four
nuclei, evidently a result of a recent conjugate division. The con-
spicuous thickenings drawn along the edges of the hypha are colored
red in the preparation with Flemming’s triple stain; their mode of
origin and significance still remain to be solved.
Figs. 3-7 show five varying views of the large botryose haustoria.
The narrow isthmus connecting the enlarged haustorium with the
extra-cellular mycelium is clearly shown in each case, as is also the
interesting fact that the haustorium, even in those cases in which the
host cell is almost entirely filled by it, pushes in as it grows the plasma
membrane of the host protoplasm. Strictly speaking, therefore, in
no case is the haustorium really imside the host protoplasm. Fig. 3
is a section of a young haustorium showing this invagination of the
host cytoplasm. Four nuclei of the rust are also shown in this prepara-
tion. In Fig. 4 an older haustorium showing its peculiar botryose
swellings, has pushed up into an unusually dense mass of host cyto-
plasm, now shrunken away from the invaginated haustorium. Figs.
5, 6, and 7 show almost equally clearly this phenomenon of invagina-
tion of the host cytoplasm. In Fig. 5, particularly, the haustorium is
seen to almost fill the host cell. An idea of the large size of the haus-
toria may be gained from the fact that while the cells of the leaf of the
host Hippocratea measure from 15 to 20 uw in diameter, those of the
fungous haustoria range from about 10 to 14 mu in diameter.
Figs. 8 to 14 show various stages in the formation and germination
of the teliospores. Fig. 8 is of a young sporiferous hypha showing two
*Lutman, B. F. Some contributions to the life history and cytology of the
smuts. Trans. Wis. Acad. Sci. 16: 1191-1244. 1910. See his Figs. 44 and 45.
OLIVE: STRUCTURE OF BOTRYORHIZA HIPPOCRATEAE 339
pairs of conjugate nuclei, those of the stalk and also those of the young
spore, which is shown in process of abstriction. As in other cases of
cell-division among fungi and algae, a ring-formed constriction grows
in from the periphery, thus cutting off in this instance the binucleate
spore from the binucleate stalk cell. Fig. 9 is a drawing of such a
young spore, showing the two nuclei; in Fig. 10, the two nuclei have
fused, though it is evident from the presence of the two nucleoles that
this fusion has only recently taken place. Figs. 11 and 12 represent
mature spores, each borne on binucleate stalks, the latter figure
showing the apical protuberance so characteristic of most spores.
That this protuberance is the primordium of the apical germ-tube is
apparent from a perusal of Figs. 13 and 14. These spores, as is
stated above, germinate at once on maturity. In Fig. 14, the hetero-
typic nuclear division is proceeding; this, however, is so poorly stained
in the preparation, that it is not possible to make out the details of the
process. Figs. 15, 16, and 17 show the basidia, or promycelia, each
composed of four uninucleate cells, which result from the germination
of the teliospores. In Fig. 15 appears a type of germination which
apparently results from growth of the basidium in a very damp situa-
tion; the four cells of the basidia in such cases often break apart
and function independently. Fig. 16 shows the more usual type of
germination, in which each of the four cells sends out a branch, to
bear finally at the tip of each a single uninucleate basidiospore on a
sterigma. Two of these oval basidiospores are shown in Fig. 18.
GENERAL DISCUSSION
It will be seen from these figures and the accompanying description
that while Botryorhiza undoubtedly resembles a short-cycled lepto-
Uromyces in the one-celled character of its teliospores, it is sufficiently
distinct in other respects to justify its being placed in a new genus.
Some of these differences are as follows: the walls of the teliospores
in Botryorhiza are thin instead of thickened, as is usual in Uromyces;
they are colorless, instead of brown or otherwise colored, as in Uro-
myces; there are no germ-pores in the walls of the spores of Botryorhiza,
whereas the teliospores of Uromyces are characterized by one or more.
Finally, the possession of such strikingly large, botryose haustoria,
so characteristic of Botryorhiza, is, in my opinion, a very distinctive
feature.
Apparently the more usual type of rust haustorium, so far as our
few studies on the subject have revealed, is that of an irregular,
branching hypha. Atkinson’ has, however, figured the haustoria of
Uromyces caryphyllinus as somewhat irregular and botryose in form;
4 College Botany. Henry Holt & Co. P. 87.
340 BROOKLYN BOTANIC GARDEN MEMOIRS
and Pole Evans® in his careful investigation of the histology of nine
species of cereal rusts has shown that while the young haustoria of
various species may be small and sac-like, or even “hammer-headed”’
in others, the prevailing type of mature haustoria in these cereal rusts
seems to be the cylindrical or branched form. To
Attention was called in the earlier description of Botryorhiza to
the fact that Lutman had figured botryose haustoria in Doassantia
and it was there suggested that this Porto Rican rust might have some
other features in common with smuts. But the fact that the sporifer-
ous hyphae are sent out through a stoma or through the ruptured
epidermis before the spores themselves are cut off from their tips, and,
further, that there is produced in Botryorhiza (and apparently in rusts
in general) a definite, swperficial hymenial layer from which the spores
arise constitutes two essential points of difference from the smuts.
The latter, as Lutman has clearly emphasized,® have their spores pro-
duced either from a group of deeply imbedded multinucleate hyphae,
which break up directly into spores (in the Ustilaginaceae) or from
the tips of the side or main branches of the prevailingly binucleate
hyphae (as in the Tilletiaceae).
It has also been brought out in the description of the characteristics
of Botryorhiza that the mycelium is composed of a branching system
of very coarse hyphal threads. These hyphae measure from 5 to 7 u
in diameter. In my own work on various rusts, I had never before
met with such a coarse mycelium. Pole Evans,’ however, has called
attention to the fact that the mycelial threads of Puccinia glumarum
reach the relatively enormous size of I0 to 19 uw in diameter; also |
Dodge in his paper in this Memoir® has noted that Farlow® and
Wornle!® have found the hyphae in Gymnosporangium Ellisii to be
exceptionally large, being, according to the latter author, about 8 u
in diameter.
5 The cereal rusts. I. The development of their uredo mycelia. Annals of
Botany 21: 441-446. 1907.
STE Ge pasL2ls.
GDA 'S le
8 See p. 128.
®The gymnosporangia of the United States. Ann. Mem. Boston Soc. Nat.
Hist. 1880: 1-38.
10 Anatomische Untersuchung der durch Gymnosporangium-Arten hervor-
gerufenen Missbildungen. Forst. Nat. Zeits. 3: 68-84; 129-172. 1894.
BROOKLYN BoTANIC GARDEN MEMOIRS. Votume |, PLate VIII.
OLIVE: CYTOLOGICAL STRUCTURE OF BOTRYORHIZA
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OLIVE: STRUCTURE OF BOTRYORHIZA HIPPOCRATEAE 341
EXPLANATION OF PLATE VIII
All drawings have been made with the camera lucida, and with various combina-
tions of Zeiss apochromatic lenses. Except where otherwise noted, the magnification
has been 1,000 diameters.
Botryorhiza Hippocrateae Whetzel & Olive
Fic. 1. Two young sori, showing the hymenial layer, composed of binucleate
cells, and method of spore formation. At the right a young sorus pushing through a
stoma. Somewhat diagrammatic. > 500.
Fic. 2. Branching hypha, showing the binucleate cells, whose walls in some
places show peculiar thickenings. X 500.
Fic. 3. Young haustorium, containing 4 nuclei. Note the invagination of the
host cytoplasm.
Fic. 4. Another haustorium, pushing into a mass of dense, granular host
protoplasm.
Fic. 5. A fully mature haustorium, showing its botryose lobings.
Fics. 6 and 7. Haustoria in partial section; showing the pushing in of host
protoplasm.
Fic. 8. A young hypha from the hymenium, showing the constricting wall
cutting off stalk from spore. X 1,500.
Fic. 9. A young spore, with two nuclei.
Fic. 10. A young spore in which the two nuclei have just fused, as evidenced
by the presence of two nucleoles.
Fics. 11 and 12. Mature teliospores, showing binucleate stalks.
Fic. 13. A spore showing young germ-tube.
Fic. 14. Another spore, with growing basidium, or promycelium. The
nucleus is in metaphase of the heterotypic division; but the preparation is poorly
stained.
Fics. 15, 16 and 17. Three basidia, which have divided into the characteristic
4 cells. Fig. 16 shows the branches, each of which will bear ultimately a single
basidiospore.
Fic. 18. Two basidiospores, showing the uninucleate condition.
THE NUCLEUS AS A CENTER OF OXIDATION
We... V. OSTERHOUT
Harvard University
In 1897 Spitzer! reported that nucleoproteins extracted from
certain animal tissues possess the same oxidizing power as the tissues
themselves. The idea that the nucleus is a center of oxidation was
advocated by Loeb? who pointed out that it would explain why cells
deprived of their nuclei are unable to live for a long time or to regen-
erate missing parts.
R. Lillie* sought to obtain direct experimental evidence by applying
to the cell reagents which become colored on oxidation. The reagent
chiefly employed was a mixture of alpha naphthol and paraphenylene
diamine which yields upon oxidation a deep purple dye, indophenol.
The oxidation takes place slowly on exposure to air, but is greatly
accelerated in the presences of living cells or extracts of living tissues.
Lillie found that in certain tissues of the frog (especially liver, kidney
and leucocytes) the colored oxidation products were deposited in and
about the nucleus, especially at the surface of contact between nucleus
and cytoplasm. :
Wherry‘ applied methyl green to Amoeba and reported that it was
oxidized in the cytoplasm but not in the nucleus. Schultze® applied a
number of stains to plant and animal cells and found that they were
oxidized in the cytoplasm but not in the nucleus.
Unna’® has investigated a large number of cases by the use of leuco-
methylene blue and has reached the conclusion that the nucleus is a
center of oxidation. Unna’s theories have been criticized by Oppen-
heimer’ and by Schneider.’
Mathews? has come to the conclusion that the nucleus is directly
concerned in oxidation.
1 Pfliiger’s Archiv. 67: 615. 1897.
* Archiv. f. Entwickelungsmechanik der Organismen 8: 689. 1899.
§’ American Jour. of Physiology 7: 412. 1902.
‘Wherry, E. T. Science, N. S. 37: 908. 1913.
’ Schultze, W. H. Verhandl. d. deutsch. path. Ges. 16: 161. 1913.
®6Unna, P.G. Archiv. f. mikr. Anat. 78. 1911. Godoletz, P. und Unna. P.
jun. Berlin, klin. Wochenschrift 49: 1134. 1912. Unna, P. G. und Godoletz, L.
Oppenheimer’s Handb. d. Biochem. Erganzungsband. 1913. S. 327.
7 Oppenheimer, C. Die Fermente und ihre Wirkung 2: 790, 810. 1913.
8 Schneider, H. Zeit. wiss. Mikr. 31: 478. 1914.
° Mathews, A. P. Physiological Chemistry p. 180. 1915.
342
OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 343
Warburg’? found that it was possible to isolate the nuclei from
erythrocytes of birds (by freezing and thawing) and that such nuclei
consumed oxygen about as rapidly as the normal cells. While this
indicates that the nucleus is the principal agent in oxidation other
experiments of Warburg have been interpreted to indicate that oxida-
tion is practically confined to the surface of the cell.1! In these experi-
ments” it was found that NaOH greatly increased oxidation in the
sea-urchin egg but did not penetrate sufficiently to cause a change of
color in eggs stained with neutral red. In a later paper R. Lillie!
comes to the conclusion that rapid oxidation occurs at the surface
of the cell as well as at the surface of the nucleus. This conclusion
is based upon a study of the indophenol reaction in the corpuscles of
frog’s blood.
The use of the indophenol reaction may encounter an objection on
the ground that the result may depend to a considerable extent on the
manner in which the reagent penetrates. If the oxidizing substances
of the cell are largely concentrated in the nucleus those which are
present in the cytoplasm will first meet the reagent at the cell surface
and may produce at that point a deposit of granules of indophenol.
In the same manner the oxidizing substances which are retained
within the nucleus will first meet the reagent at the surface of the
nucleus and produce a deposit in that region.
If, therefore, the indophenol reaction shows a higher oxidative
activity in the nucleus it may doubtless be depended on, since its
error presumably lies in the opposite direction. But if it indicates a
marked oxidative activity at the surface of the cell (or at internal
surfaces, including that of the nucleus) we must be cautious in drawing
conclusions.
It would seem that more reliable evidence can be obtained by
investigating cases where it is not necessary that the reagent should
penetrate from without owing to the fact that the cell itself produces
substances which become colored on oxidation.
The writer has investigated a case of this kind. The plant chosen
was the Indian Pipe, Monotropa uniflora, which is extremely well suited
to such investigations because the colorless cells contain a’ chromo-
gen which oxidizes and darkens very rapidly upon injury. An addi-
10 Warburg, O. Zeit. f. physiol. chem. 70: 413. IgIO-II.
This interpretation is by no means necessary. Cf. Loeb and Wasteneys,
Jour. of Biochemistry 14: 459. 1913; 21: 153. 1915; also, Osterhout, ibid. 19: 335.
1914. Owing to the buffer action of protoplasm and to the presence of pigment
-the penetration of a small amount of alkali is not easily detected.
2 Warburg, O. Zeit. f. physiol. chem. 66: 305. tIg10. Biochem. Zeit. 29:
414. 1910.
18 Jour. of Biol. Chem. 15: 237. 1913.
344 BROOKLYN BOTANIC GARDEN MEMOIRS
tional advantage is that the leaves are so thin and transparent that
they may be placed under the microscope and the details of cell struc-
ture studied with care before the cells are injured or treated with
reagents.
In a typical leaf cell the cytoplasm is transparent and nearly color-
less, with a few granules, while the nucleus is only slightly less trans-
parent and as a rule shows a few granules and a nucleolus. When a
leaf is mounted in a drop of water under a cover glass the cells remain
unchanged in appearance for hours.
If an intact portion of the leaf is cut or crushed the cells in the
neighborhood of the injury soon change their appearance. In the
course of five or ten minutes the nuclei of the cells nearest the injury
assume a more granular (or vacuolated) appearance and soon begin to
darken. The darkening does not begin at the surface but appears to
take place almost simultaneously throughout the whole mass of the
nucleus. Not until the nucleus has become very dark (so as to stand
out very conspicuously when the preparation is viewed under the low
power of the microscope) does the cytoplasm begin to darken per-
ceptibly. It may be several hours after the nucleus has darkened
before a change of color can be perceived in the cytoplasm. (This
is also true where the thickness of the cytoplasm has been increased by
plasmolysis so as to be as great as that of the nucleus.) The darkening
of the cytoplasm does not seem to be more rapid at the surface than
elsewhere.
That the darkening is due to oxidation is shown by several facts.
Among these the following may be mentioned.
1. A microscope slide is smeared with vaseline, a leaf is laid upon
the vaseline and more vaseline is carefully placed upon the leaf. A
small splinter of glass (from a broken slide) is placed on the leaf and
another slide is gently pressed upon it, so as to spread the vaseline and
bring the glass splinter close to the leaf without injuring the latter.
Care should be taken that any air bubbles which may be included in
the vaseline are not in contact with the leaf in the neighborhood of
the splinter of glass.
The leaf is left over night in order that the oxygen present in the
intercellular spaces (or adhering to the surface of the leaf) may be
used up by respiration. On the following morning the upper slide is
pressed down with sufficient force to drive the splinter into the leaf
and crush it. It is then placed on the stage of the microscope and kept
under observation. It is found that while some darkening occurs it
is at first largely confined to the drops of juice forced out of the leaf
by the crushing (the juice seems to spread along the fibro-vascular
bundles ia some cases). The darkening of the nucleus and cytoplasm
is usually much slower than in air (especially with fresh leaves).
OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 345
The darkening which occurs is due in part to free oxygen left in the
leaf and in part to oxygen in compounds from which it can be split
off for the oxidation of the chromogen (analogous to anaérobic
respiration).
2. If leaves are torn in two or crushed at once, dropped into boiling
water, 0.1 M HCl, 0.1 M NaOH, 0.1 M KCN, or 3 percent H2Oz the
darkening does not occur. These agents are inhibitors of oxidation in
living tissues. Hydrogen peroxide may inhibit at high concentration,
but accelerate at low concentrations. In NaOH and KCN the leaf
becomes pale yellow: this seems to be due to the action of hydroxyl
ions.
3. The chromogen may be extracted by placing stems in 0.1 WM
NaOH in a bottle completely filled (so as to exclude air) and tightly
stoppered (with a glass stopper coated with vaseline). The solution
becomes pale yellow (or slightly reddish) and may be kept in this
condition for months. On opening the bottle and pouring out the
solution into a shallow dish it at once becomes red as the result of
oxidation. The behavior seems to be analogous to that of pyrogallol,
which is easily oxidized by the air in alkaline solution, but not in
neutral solution except under the influence of oxidases (from plants or
animals) or other catalyzers.
That the darkening of the nucleus is due to oxidation taking place
in the nucleus itself and not to the taking up by the nucleus of a stain
produced in the cytoplasm or vacuoles is shown by the following
experiment. Plants were ground in a mortar and allowed to stand
until they became black. The juice was squeezed out and centrifuged,
giving an inky fluid. In this were placed pieces of leaves which had
been treated with 0.1 KCN and afterward with water. The solution
was allowed to stand until it became concentrated by evaporation:
it then appeared black. It was found that where the nuclei had been
squeezed out of the cut cells by the knife they had taken up some stain
but not more than the cytoplasm. In cells which were merely cut
open there was little or no staining of the nucleus.
We must therefore conclude that oxidation occurs more rapidly in
the nucleus than elsewhere in the cell. The only way to escape this
conclusion would be by assuming that at the moment of injury there
is a sudden migration into the nucleus of some or all of the substances
necessary for the oxidation. This is not only very improbable from a
theoretical standpoint, but observation shows that it can not be the
case, for in this migration the substances would mingle and produce
the pigment either outside the nucleus or at its surface before any
pigment appeared in the interior of the nucleus. Observation of the
nucleus shows that the pigment appears as soon in the interior of the
nucleus as at its surface.
346 BROOKLYN BOTANIC GARDEN MEMOIRS
We may therefore conclude that the substances necessary for
oxidation do not suddenly migrate into the nucleus at the moment
of injury, but that they must exist there before the cell is injured.
We may ask why the nucleus does not become darkened in the
normal condition of the cell. The investigations of several workers
have made it probable that the pigments produced by oxidation under
normal conditions are at once reduced, giving up their oxygen to other
substances in the cell. When injury occurs the reduction is checked
more rapidly than the oxidation, with the result that the pigment
accumulates.
It is also possible that injury causes the admission of oxygen to
the cells.
In order to test the effect of the indophenol reaction on leaves of
Monotropa they were torn in two and placed in a mixture of equal
parts of aqueous I percent paraphenylene diamine and saturated
aqueous alpha naphthol. It was found that the result depends some-
what on the condition of the reagent. In the most favorable cases
the cells which were torn open became pale purple in color almost
at once, showing that the reagent readily penetrated them. Usually
the cell contents (cytoplasm, nucleus and vacuole) became at first
uniformly tinged with purple. After a while the nuclei would usually
assume a deeper purple than the remainder of the cell contents.
The cells lying a little further from the torn surface, which were
injured but not actually torn open, showed at first a pale yellowish
color which in some cases became deeper with. time and in other
cases gave way toa purplish tint. In most of these cells the nuclei
gradually became deeper in color than the other cell contents. Later
the cytoplasm became in some cases so deep in color as to obscure the
nuclei. Cells lying still further from the torn surface changed very
slowly (many remaining unchanged after some hours) so that it was
evident that the reagent penetrated from the torn surface and not
through the outer cell walls (which are normally in contact with the
ai!
In most cases the general result, after a few minutes, was a deep
purple band along the torn edge: inside the purple band was a yellow-
ish one of irregular outline, followed by nearly colorless intact cells
further away from the torn edge.
In I percent aqueous paraphenylene diamine the results were
similar but the purple color was replaced by a dirty brownish-red
(with more or less purplish tinge).
It should be pointed out that these results are most striking with
4 More rapid penetration from cut or torn surfaces is commonly observed in
the entrance of reagents into leaves, petals, etc.
OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 347
reagents which have stood long enough to take up oxygen, in conse-
quence of which the paraphenylene diamine becomes reddish in color,
while the alpha naphthol assumes a dirty grayish-purple.
When the reagents are freshly made up the action is very slow
unless hydrogen peroxide be added.. When a mixture is made up of
equal parts of each of the reagents previously mentioned and 0.3
percent hydrogen peroxide the results are similar to those just de-
scribed. But if stronger hydrogen peroxide be used a greater amount
of purple coloration is observed in the cells.
When 3 percent hydrogen peroxide is used (in place of 0.3 percent)
the following changes may be observed. A pronounced purple color
appears at once in the torn cells: this spreads rapidly to the adjacent
cells, which are still intact, and may extend through several rows of
intact cells. In these intact cells the first appearance of change is the
formation of purple granules of indophenol in the vacuole. The vacu-
ole becomes filled with these granules which show active Brownian
movement. Occasionally some of them come in contact with the
nucleus (or the film of cytoplasm which covers the nucleus) and stick
fast to it. At this time nucleus and cytoplasm are usually free from
granules or coloration. The purple color grows more intense until
the details of cell structure become obscured.
The general conclusion is that while the indophenol reaction indi-
cates that the nucleus is the center of oxidation it does not give as
definite information on this point as the formation of natural pigments
within the cell as the result of the oxidation of substances normally
present.
SUMMARY
Injury produces in the leaf-cells of the Indian Pipe (Monotropa
uniflora), a darkening which is due to oxidation. The oxidation is
much more rapid in the nucleus than in the cytoplasm and the facts
indicate that this is also the case with the oxidation of the uninjured
cell.
PHYSIOLOGICAL SPECIALIZATION OF PARASITIC
FUNGI
GEORGE M. REED
University of Missouri
One of the important developments in plant pathology in recent
years has been the discovery of races of well-defined morphological
species of parasitic fungi which are restricted to particular hosts.
These specialized races can be distinguished from each other only
by their ability to grow on some host plants and not on others. It is
now well established that species of parasitic fungi, identical in their
structural features as found on a more or less wide range of plants,
may consist of numerous races or strains which differ in their capacity
to infect the various hosts.
Apparently Schroeter (136), as early as 1879, was the first to call
attention to this phenomenon in connection with certain rusts on
Carex. It is, however, to Eriksson (34) that we are indebted for a
realization of the importance and significance of the host specialization
of fungous parasites and for the impetus to the numerous investiga-
tions devoted to this phase of plant pathology. Eriksson’s demon-
stration of races of Puccinia graminis, P. glumarum, P. dispersa and |
P. coronata, distinguishable from each other only on the basis of the
hosts that they are able to successfully attack, is the real starting point
for a general recognition of the phenomenon of host specialization of
fungous parasites.
Various terms have been introduced to apply to these races or
strains which show no anatomical differences, but are distinguishable
only by their physiological behavior in the choice of hosts. Schroeter
(138), in 1893, suggested the term sister species (Species sorores).
Klebahn (76), in 1892, described them as biologische Spezies. Rostrup
(121, 122), in 1894, suggested the term biologiske Arter, and, in 1896,
proposed another term biologische Rassen. In 1894 Hitchcock and
Carleton (63) proposed the term physiological species. Eriksson (34),
in 1894, introduced the generally used expression Specialisierte Formen
or formae speciales. Magnus (96), in 1894, employed the term
Gewohnheitsrassen or adapted races. Marchal (97) applied the term
races spéciali ées, while Ward (174), Salmon (123) and others have
used the expression biologic forms or biological forms. '
348
REED: SPECIALIZATION OF PARASITIC FUNGI 349
Whatever term has been applied, the underlying conception has
been the same, namely, that these races, strains, forms, etc., of distinct
morphological species of fungous parasites differ, not in discernible
structural features, but in their physiological behavior, as indicated
by their ability to infect some hosts and not others. They differ in
their ability to establish the parasitic relation with particular hosts and
thus secure the necessary food for their normal development. The
phenomenon is distinctly physiological and is doubtless quite com-
parable to the well-known behavior of saprophytic fungi on different
chemical substrata. Various saprophytes, structually similar, vary
in their ability to utilize different chemicals as sources of food, de-
pendent on their capacity to secrete the necessary enzymes. While
the strains of parasites may differ essentially in their ability to secure
food from a particular host, we must keep in mind the possibility of a
more complicated series of relations in which toxin and antitoxin
production are involved.
Many investigators of the phenomenon of host specialization have
made a large number of species on the basis of the results of their
inoculation tests. This is especially the case in the rusts where
Klebahn, Eriksson, Schneider, Fischer and others have raised many
forms to specific rank, although no distinct structural differences can
be observed. It may be noted that the races of Puccinia dispersa,
P. sessilis, P. Ribesti-Caricis, P. extensicola, Coleosporium Campanulae,
Melampsora populina, M. Tremulae and others, referred to below, are
regarded as good species by some students.
Fischer (45), in connection with the rusts, accepts as species the
following:
1. All rusts which are structurally distinct.
2. All rusts which have a different life-cycle; for example, forms
which are distinguished by the presence or absence of certain
spore-forms.
3. All forms which differ in their choice of hosts, in so far as the
hosts belong to different genera. In heteroecious rusts species
are recognized when the hosts of one generation, aecidial or
uredo and teleuto, belong to two different genera.
Fischer unites under one species as formae speciales or specialized
races all rusts which differ only physiologically and whose hosts are
species of a single genus. Whether a particular rust is a physiological
species or a specialized race is thus determined by the range of its
hosts.
It is doubtless true that many rusts, and other parasitic fungi
as well, which can be distinguished only by the hosts upon which they
grow, are just as distinct forms as others which are characterized by
24
350 BROOKLYN BOTANIC GARDEN MEMOIRS
minute structural differences. The poplar rusts of the group Melamp-
sora Tremulae, with their uredo and teleuto stages on Populus alba
and P. tremula and their aecidial stage on such widely separated hosts
as Larix decidua, Pinus silvestris, Mercurialis perennis and Chelidonium
major, certainly differ from each other in a fundamental way. The
difference between these is doubtless as significant as some of the
minute structural differences which distinguish other species.
It is, however, certainly important to recognize the fact that these
rusts, mildews, etc., referred to as specialized races, physiological
species, etc., can be distinguished only by cultural tests. This can
most easily be done by grouping them together on the basis of struc-
tural similarity. Klebahn (92) has done this recently in his scheme
for illustrating the relationships of the willow and poplar rusts, and
also in the case of the Ribes-Carex rusts. Arthur (10) adopts the same
plan in combining the Compositae-Carex rusts under Puccinia ex-
tensicola Plowr. In a similar way Tranzschel (155) combines the
various Centaurea-Carex rusts under the name Puccinia Centaureae-
Caricis.
At the present time, a large number of the parasitic fungi have
been investigated from the standpoint of specialization to particular
hosts and the phenomenon has been found to be of wide occurrence.
The present paper is an attempt to bring together the results of
numerous investigations bearing on this point.
THE Rusts—UREDINEAE
Puccinia graminis Pers. Extensive studies have been made in
both Europe and the United States on the specialization of the black
stem rust of the cereals and other grasses. Eriksson (34, 37, 38, 41)
in Sweden, Jaczewski (68) in Russia, Carleton (25, 26), Freeman and
Johnson (57), Arthur (2; 5, 6,7; 8} 10, 11), and Stakman and ea-
workers (143-149) in the United States, have reported the results of
_ their cultural experiments with this rust. Experiments have been
reported in which inoculation tests with both uredospores from the
various grass hosts and aecidiospores from the barberry have been
used. The general results of these experiments may best be sum-
marized as follows:
351.
SPECIALIZATION OF PARASITIC FUNGI
REED
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BROOKLYN BOTANIC GARDEN MEMOIRS
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355
SPECIALIZATION OF PARASITIC FUNGI
REED
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BROOKLYN BOTANIC GARDEN MEMOIRS
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REED: SPECIALIZATION OF PARASITIC FUNGI 357
According to the various workers the specialized race Avenae
occurs on a wide range of hosts. Eriksson (41) lists it on twenty
species belonging to fourteen genera; Jaczewski (68) lists it on seven
species belonging to six genera; Carleton (25) records it on nineteen
species belonging to fifteen genera; Stakman and Piemeisel (149)
record it on thirty-three species belonging to twenty-one genera. The
specialized race Tritici is reported by Carleton on seventeen species
belonging to seven genera; Stakman and Piemeisel list it on thirty-
three species belonging to nine genera; Jaczewski lists it on six species
belonging to five genera; Eriksson records it on the four cereals and
states that this race seems less sharply fixed in its host relations than
any of the others. Stakman and Piemeisel report another race,
Tritici compacti, from the Palouse wheat region of western United
States. This race is reported on twenty species belonging to six
genera. The specialized race Secalis also occurs on a number of hosts,
Eriksson reporting it on eleven species belonging to five genera,
Jaczewski reporting it on six species belonging to four genera and
Stakman and Piemeisel recording it on twenty-three species belonging
to nine genera. According to Eriksson, Jaczewski and Carleton the
specialized race Agrostis occurs only on species of Agrostis, while
Stakman and Piemeisel report it on thirteen species of grasses be-
longing to ten genera. The specialized race Arrhenathert occurs on
both Avena sativa and Arrhenatherum elatius. The races Airae, Poae,
Calamagrostis and A perae are limited to species of a single genus.
Eriksson and Henning (43) separated the timothy rust from the
black-stem rust and made it a distinct species, Puccinia Phlei-pratensis.
The separation was based largely on the fact that they were unable,
with one doubtful exception, to infect the barberry with teleutospores
from timothy. Other workers have made similar attempts but with-
out success. Stakman and Piemeisel consider it best to regard timothy
rust as a race of Puccinia graminis.
Eriksson (34) reports successful inoculations with uredospores from
timothy on Avena sativa, Festuca elatior and Secale cereale, negative
results being obtained with Triticum vulgare and Hordeum vulgare.
Johnson (69) transferred timothy. rust, using uredospores, to Avena
sativa, Arrhenatherum elatius, Dactylis glomerata, Festuca elatior, Poa
compressa and Secale cereale. Inoculations on thirteen other grasses,
including Triticum vulgare and Hordeum vulgare, gave negative results.
Stakman and Piemeisel (149), however, were able to infect eighteen
species of grasses belonging to thirteen genera, these including Avena
sativa, Hordeum vulgare and Secale cereale, but not Triticum vulgare.
They regard this race as being closely related to the race Avenae.
It may also be noted that Eriksson lists barley (Hordeum vulgare)
358 BROOKLYN BOTANIC GARDEN MEMOIRS
and rye (Secale cereale) as hosts for the same specialized race Secalis.
Carleton, on the other hand, places wheat and barley as common hosts
for the race Tritici, although he gives no information regarding the
relation of the rye rust to the other forms. Other variations in the
hosts for the different races, as reported by these three workers, also
occur. Eriksson lists Dactylis glomerata as a host for the race Avenae;
Jaczewski lists it for the race Secalis; and Carleton records it as a
host for both races Avenae and Tritict. Agropyron repens is a host for
the two races Secalis and Tritici according to Jaczewski. Carleton
records the race Avenae on Hordeum murinum, while Eriksson places
this host, along with the other species of Hordeum, as a host for the
race Secalis.
It is important to note that many grasses are listed by Stakman
and Piemeisel as common hosts for several races. They suggest that
the six races which they experimented with may be divided into two
groups on the basis of their parasitism. The races Tritici, Tritict
compacti and Secalis form one group; these vary in their capacity for
infecting certain hosts but all three vigorously infect A gropyron
cristatum, A. elongatum, A. smithi1, Bromus tectorum, Hordeum jubatum,
H. vulgare, Elymus canadensis and Hystrix patula. The other three
races, Agrostis, Avenae and Phleipratensis, also vary in their infecting
capacity but all vigorously attack Alopecurus geniculatus, A. pratensis,
Dactylis glomerata, Holcus lanatus and Koeleria cristata. Three hosts,
Bromus tectorum, Hordeum vulgare and Secale cereale, are infected by
all six races.
Stakman and Piemeisel do not regard the barley rust as a distinct
race. Barley is a very favorable host for races Secalis, Tritici, and
Tritict compacti, as well as being susceptible to the other three races.
In the field barley seems especially to harbor the race Tvitici.
Freeman and Johnson (57) have confined their work almost ex-
clusively to the cereal hosts of Puccinia graminis. They conclude
that their experiments indicate the existence of four specialized races:
Tritict on wheat, Hordei on barley, Secalis on rye, and Avenae on oats.
The rust on wheat can be transferred to barley and rye, but not to
oats; the rust on barley can be transferred to the other three cereals,
rye, oats and wheat; the rust on rye can be transferred to barley, but
not to wheat nor oats; the rust on oats can be transferred to barley.
These workers also report that Hordeum jubatum can be infected with
the rust of both wheat and barley; Agropyron repens with the rust of
wheat; and Dactylis glomerata with the rust of oats. So far as the
cereals are concerned it appears that the different grains may be
hosts for more than one specialized race of the black-stem rust. Stak-
man and Piemeisel, however, do not regard Hordei as distinct from
Tritict.
REED: SPECIALIZATION OF PARASITIC FUNGI 359
Stakman’s (143) results with the cereal rusts, in general, confirm
those of Freeman and Johnson. He found that uredospores from
barley infected rye, barley and wheat, but not oats; uredospores from
oats infected wheat, rye and oats, but not barley; uredospores from
rye infected rye, but not wheat nor barley; uredospores from wheat
infected barley, rye and wheat, but not oats. Uredospores from
Agropyron repens gave the following results: on wheat, on oats, on
barley and on rye.
Stakman (144) briefly mentions some other results with this rust.
He states that uredospores from Agropyron repens, A. tenerum, A.
caninum, A. smithii and Hordeum jubatum readily infect barley and
rye, very slightly wheat and practically fail to infect oats. Uredo-
spores from Dactylis glomerata and Poa nemoralis infect oats, but not
the other cereals. Practically no success was attained in trying to
infect any of the cereals with uredospores from Agrostis alba, A.
stolonifera, Anthoxanthum odoratum, Calamagrostis canadensis, Poa
pratensis and P. compressa.
Bolley and Pritchard (20) state that uredospores from barley
(Hordeum jubatum) and Avena fatua were able to infect wheat and
uredospores from wheat produced infection on barley and Hordeum
jubatum.
Pritchard (110), in North Dakota, suggests that distinct races
occur on wheat and barley, although he gives no experimental evi-
dence. Another race occurs on rye, oats, Avena fatua, Agropyron
repens, A. tenerum and Hordeum jubatum, as indicated by inoculations
with uredospores from the gramineous hosts and aecidiospores from
the barberry.
Gassner (58) has reported brief results on this rust in South America.
He was able to infect wheat with uredospores taken from barley, oats,
rye, Lolium temulentum and Dactylis glomerata. Barley was also
infected by uredospores from wheat.
As noted before, the infecting capacity of aecidiospores from bar-
berry has been tested, as well as that of the uredospores from various
gramineous hosts. Eriksson (41) has infected the barberry with
teleutospores from more than fifty different grasses. In some cases
the aecidiospores thus experimentally produced were used to inoculate
various grasses. In general, the aecidiospores from the barberry are
restricted in their ability to infect grasses in the same way as uredo-
spores from the grasses which were used as a source for the teleuto-
spores for inoculating the barberry. It should be noted, however,
-that aecidiospores from the barberry, produced by inoculation with
teleutospores from Bromus madritensis, Briza maxima, Festuca myurus
and Phalaris canariensis, recorded as hosts for the specialized race
360 BROOKLYN BOTANIC GARDEN MEMOIRS
Avenae, infected Secale cereale as well as Avena sativa. Eriksson,
however, gives no information regarding the infecting capacity of
uredospores from the four grasses mentioned.
Jaczewski (68) made fairly complete tests with aecidiospores from
the barberry, obtained by inoculation with teleutospores from various
gramineous hosts. He records exact correspondence between the
results obtained with the aecidiospores and uredospores from the
grasses used as a source for the teleutosporic infections of the barberry.
Pritchard (110) found that aecidiospores from barberry in the open
infected Avena sativa, A. fatua, Agropyron tenerum, A. repens, Hordeum
jubatum and Secale cereale, but not Hordeum vulgare nor Triticum vul-
gare. They thus correspond to the infecting capacity of uredospores
from the gramineous hosts.
Stakman’s (143) results are in harmony with those mentioned. In
one trial, the barberry was infected with teleutospores from wheat,
although standing in the open. The aecidiospores, when inoculated
onto various plants,*infected the following: wheat, barley, oats, rye
and Triticum monococcum. In another test, aecidiospores from the
barberry, produced by inoculation with teleutospores from A gropyron
repens, infected wheat, barley, and rye, but not oats, while a similar
series, starting with teleutospores from wheat, infected wheat, barley,
rye, but not oats. These results are in close correspondence to those
previously mentioned for uredospores.
Arthur (7, 8, 10) has infected the barberry with teleutospores from
Agrostis alba; the aecidiospores produced infected Hordeum vulgare
and Triticum vulgare but not Avena sativa. Teleutospores from
Elymus canadensis also infected the barberry but the aecidiospores
developed failed to infect Secale cereale and Triticum vulgare. Other
results of Arthur are the following: aecidiospores from barberry,
arising from inoculation with teleutospores from Agropyron tenerum,
infected Avena sativa and aecidiospores, arising from inoculation with
teleutospores from Sitanion longifolium, infected Triticum vulgare.
Freeman and Johnson (57) studied the variations in the size and
shape of the uredospores of the different races. While the uredospores
of the same race varied considerably in these points, yet they found
well-defined differences in the uredospores of the various races.
Stakman and Piemeisel have made similar studies and confirm the
conclusions of Freeman and Johnson.
Puccinia coronata Corda. Next to Puccinia graminis the crown
rust of grasses has been the most extensively investigated rust from
the standpoint of its heteroecism and its restriction to hosts. De-
Bary (17) first established the fact that the aecidial stage of a crown
rust on grasses occurred on Rhamnus. Plowright (108) seems to have
REED: SPECIALIZATION OF PARASITIC FUNGI 361
been the first to suggest that the aecidial stage of the rust on Rhamnus
cathartica and Rh. Frangula belonged to two different groups of
gramineous hosts. This supposition was later confirmed by Eriksson
(34, 37, 42), Klebahn (77-81, 90) and Miihlethaler (101, 102). These
same workers have further discovered the existence of races of the rust
on the groups of grasses which have their aecidial hosts on different
species of Rhamnus. ‘The specialization of Puccinia coronata Corda
has been found to be as follows:
I. SPECIALIZED RACES WITH THEIR AECIDIAL STAGE ON Rhamnus cathartica, Rh.
utilis, Rh. dahurica, Rh. saxatilis AND Rh. Imeretina (Puccinia coronifera Kleb.)
1. Avenae on Avena sativa and A. brevis.
2. Alopecurt on Alopecurus pratensis, A. arundinaceus and, to some extent, on
Avena sativa.
3. Festucae on Festuca arundinacea, F. elatior, F. gigantea, F. varia and F. alpina.
4. Lolit on Lolium remotum var. aristatum, L. perenne, L. rigidum, L. temulentum,
L. italicum, Festuca elatior and Holcus lanatus.
5. Glyceriae on Glyceria aquatica.
6. Agropyrt on Agropyron repens.
7. Epigaei on Calamagrostis epigeios and to some extent on Avena sativa.
8. Holct on Holcus lanatus and Lolium perenne.
9g. Bromi on Bromus erectus, B. erectus var. condensatus, B. inermis, B. secalinus, B.
sterilis, B. tectorum, B. commutatus and probably B. asper.
10. Arrhenatheri on Arrhenatherum elatius.
Il. SPECIALIZED RACES WITH THEIR AECIDIAL STAGE ON Rhamnus Frangula, Rh.
Purshiana, Rh. Alaternus, Rh. californica AND Rh. Imeretina
(Puccinia coronata (Corda) Kleb.)
1. Calamagrostis on Calamagrostis arundinacea, C. lanceolata, C. phragmitoides, C.
calybea and Phalaris arundinacea.
. Phalaridis on Phalaris arundinacea, Calamagrostis arundinacea and C. lanceolata.
. Agrostis on Agrostis vulgaris and A. stolontfera.
. Holct on Hoicus lanatus and H. mollis.
. Agropyri on Agropyron repens.
nb &® dN
Il1, Race wiTH ITs AECIDIAL STAGE ON Rhamnus alpina, Rh. pumila, Rh. Imeretina
AND Rh. Purshiana (Puccinia alpinae-coronata Miihlethaler)
Calamagrostis on Calamagrostis varia and C. tenella.
IV. RACE WITH ITs AECIDIAL STAGE ON Rhamnus dahurica
(Puccinia himalensis (Barclay) Diet.)
Brachypodii on Brachypodium silvaticum.
V. RACE WHOSE AECIDIAL STAGE IS UNKNOWN
Melicae on Melica nutans.
362 BROOKLYN BOTANIC GARDEN MEMOIRS
It will be noted that the specialized races of groups I, II, and III
have Rhamnus Imeretina as a common aecidial host. Rhamnus
Purshiana also occurs as an aecidial host for races of groups I and III.
It also appears that specialized races on Holcus and Agropyron occur
in both groups I and II. In connection with the gramineous hosts
certain specialized races also overlap. For example, Avena sativa is a
host for specialized races Avenae, Alopecuri and Epigaei (Group 1);
Festuca elatior is a host for specialized races Lolii and Festucae (Group
1); Calamagrostis arundinacea, C. lanceolata and Phalaris arundinacea
are hosts for specialized races Calamagrostis and Phalaridis (Group I).
Carleton (26), in this country, has tested the host relations of the
crown rust on oats. He finds that the rust on Avena sativa can be
transferred to Avena sativa patula, A. sativa orientalis, A. sativa nuda,
A. fatua, A. pratensis, Alopecurus alpestris, Phleum pratense, Ph.
asperum, Dactylis glomerata, Aira caespitosa, Holcus mollis, Eatonia
sp. indet., Koeleria cristata, Anthoxanthum odoratum, Festuca sp. indet.,
Phalaris arundinacea, Polypogon monspeliensis, Trisetum subspicatum,
Brizopyron siculum and Poa annua. Carleton also transferred the
rust from Phalaris caroliniana to Avena sativa and Dactylis glomerata.
The rust on Arrhenatherum elatius was also transferred to Avena sativa.
Aecidiospores from Rhamnus lanceolata readily infected Phalaris
caroliniana and Avena sativa.
Arthur (6, 11, 14) reports the successful infection of Avena sativa
with ,aecidiospores from Rhamnus lanceolata, Rh. caroliniana and Rh.
cathartica. He also succeeded in infecting Rhamnus alnifolia with
teleutospores from Calamagrostis canadensis. Teleutospores from
Holcus lanatus and Scolochloa festucacea failed to infect Rhamnus
cathartica.
Treboux (159, 160) reports the results of experiments with the
crown rust carried out in southern Russia which are quite at variance
with those obtained in Europe. He finds that aecidiospores from
Rhamnus cathartica obtained from a common source infected fifty-
one species of grasses belonging to the genera Alopecurus, Agropyron,
Agrostis, Aira, Arrhenatherum, Avena, Brachypodium, Briza, Bromus,
Calamagrostis, Dactylis, Eatonia, Festuca, Glyceria, Hierchloa, Holcus,
Hordeum, Koeleria, Lolium, Melica, Phalaris, Poa, Phleum, Poly-
pogon, Sclerochloa, Secale, Sesleria and Triticum. In these experi-
ments with Puccinia coronifera Kleb., Treboux obtained positive
results with three hosts of P. coronata (Corda) Kleb.—A grostis stolonzf-
era, Calamagrostis arundinacea and Phalaris arundinacea. Treboux
has also used the aecidiospores from Rhamnus Frangula to successfully
infect nine species of grasses, among them Avena sativa, a host belong-
ing to P. coronifera Kleb. Treboux’s tests with uredospores show the
REED: SPECIALIZATION OF PARASITIC FUNGI 363
same lack of specialization to particular hosts as Carleton reports for
the crown rust in the United States.
Puccinia glumarum (Schm.) Eriks. and. Henn. Eriksson (34) is
the only investigator to report on the host relations of this rust. He
claims the existence of five specialized races:
Tritict on Triticum vulgare.
. Secalis on Secale cereale and Triticum vulgare.
. Elymi on Elymus arenarius.
. Agropyri on Agropyron repens.
. Hordet on Hordeum vulgare.
Puccinia dispersa Eriks. and Henn. Ericksson (34, 40) first
separated the hosts of this rust into five distinct groups each sup-
porting a distinct specialized race. Later each race was raised to
specific rank, based on the host relations and life history.
1. Secalis on Secale cereale. ‘This race has its aecidial stage on different
species of Anchusa.
2. Agropyri on Agropyron repens (Puccinia agropyrina Eriks.).
Aecidial host unknown.
3. Bromi on Bromus species (Puccinia bromina Eriks.). According to
Miiller (99), this race has its aecidial stage on Pulmonaria mon-
tana and Symphytum officinalis.
4. Triticti on Triticum vulgare (Puccinia triticina Eriks.). Aecidial
host unknown.
5. Holct on Holcus lanatus and H. mollis (Puccinia holcina Eriks.).
Aecidial host unknown.
6. Triseti on Trisetum flavescens (Puccinia Triseti Eriks.). Aecidial
host unknown.
Miiller, Ward, and Freeman have studied the rust of the bromes.
Miller (103) found that aecidiospores from Pulmonaria montana
infected Bromus arvensis, B. brachystachys, B. erectus, B. mollis and
B. secalinus. Aecidiospores from Symphytum officinalis also infected
these species of Bromus and, in addition, B. brizaeformis. Miiller
obtained the following results with uredospores: (1) uredospores from
Bromus erectus infected B. arvensis, B. brachystachys, B. erectus, B.
macrostachys and B. mollis; (2) uredospores from B. arvensis infected
B. arvensis, B. brachystachys, B. inermis and B. mollis; (3) uredospores
from B. mollis infected B. brachystachys, B. macrostachys and B. mollis.
Ward (171, 172) has carried out a large series of experiments with
this rust. In his inoculation tests he used species of Bromus belonging
to each of the five recognized subdivisions of the genus. Uredospores
from eleven different species, belonging to three different sections of
the genus, were used. Ward found marked differences in the sus-
ceptibility of the bromes to the uredospores from different hosts. In
ap ® NH
364 BROOKLYN BOTANIC GARDEN MEMOIRS
general, he concludes that species of the same section of the genus as
the one serving as a source of uredospores were more fully infected
than the species of other sections. The evidence for Ward’s con-
clusion is not very striking except in the case of the two hosts Bromus
mollis and B. sterilis. Some of Ward’s data may be indicated as
follows:
Source of Uredospores
Host Inoculated Bromus) Bromus Bromus\ Bromus
arven- | brizae-| PB*ORUS | secali- \arduen-| Promtus
sis | formis mollis nus | nensis| Sterttis
Serrafalcus: |
BONUS GLUCNSES.< on. oc oe «ea 12/13°| 10/9 33/97 6/8
Bromus brachystachys.......... rai Pra ra 77
Bromus brizaeformis........... Fi 7 DSi) eLA25 ll Bis
IBY OTTUSIST USC URE EE eae eter ly 2n/e7 14/59
Bromus macrostachys...........| 14/15 16/17 5/19 5/5
Bromus moliiformis............ | | | 2/6 1/25
Bromusimolise een or eee 8/15 | 21/26) 119/154 | 3/8 1/8 T/T,
Bromus pendulinus............ 8/6 | 43/40) 30/50 | |» 7/65
Bromus) SCCOMNUS!.. 22> ciesne eee } 14/14) 14/15| 31/61 16/16| 8/8
Bromus vestitus.........0.05-- 3/4 1/4
Libertia: |
Bromus arduennensts..........-- | 13/14 8/7
Stenobromus: | :
BrOMmUus ZUSSONY. woes ess es - 10/26| 6/53 37/60
Bromus madritensts............ 11/13 | 1/13 43/68
Bromus Maximus. ... 060.000 eee eresi el 2/82
BROMUS SIC IIS) case ene eee | 4/148 126/146
Freeman (56) has made a further study of the brome rust, using
uredospores from Bromus mollis and B. sterilis. He states that twenty-
two different species of Bromus remained free from infection following
inoculation with rust from both hosts. Eleven species were infected
with uredospores from B. mollis but not with uredospores from B.
sterilis. Only one host, B. sterilis, was infected by uredospores from
B. sterilis, and not by uredospores from B. mollis. Five species were
infected with spores from both grasses.
Puccinia Stipina Tranzschel. Under this name, Tranzschel (156)
groups the North American rust Puccinia Stipae Arth., with uredo
and teleuto on Stipa spartea and aecidial stage on Aster ericoides, A.
multiflorus, A. Novae-angliae and Solidago canadensis, and the Euro-
pean rust, Puccinia Stipae Bubak, with uredo and teleuto stages on
Stipa capillata and aecidial stage on Thymus and Salvia. Klebahn
(87, 91) finds evidence for two specialized races in the European rust:
5 The denominator indicates the number of leaves inoculated and the numerator
the number infected.
REED: SPECIALIZATION OF PARASITIC FUNGI 365
1. Thymi-Stipae Kleb.; aecidial stage on Thymus serpyphyllum, T.
angustifolius and T. vulgaris.
2. Salviae-Stipae Kleb.; aecidial stage on Salvia silvestris and S.
pratensis.
Puccinia sessilis Schneid. This rust, which has its uredo and
teleuto stages on Phalaris arundinacea and its aecidial stage on various
plants of the Liliaceae, Amaryllidaceae, Araceae and Orchidaceae, has
been broken up into a number of distinct species, based largely upon
the choice of the aecidial host. Klebahn (77-83, 86, 87, 89-91) has
carried out the cultural tests with the rust and recognizes the following
relations:
1. Puccinia Smilacearum-Digraphidis (Sopp.) Kleb.
a. Smilacearum-Digraphidis typica Kleb.; aecidial stage on Con-
vallaria majalis, Maianthemum Obifolium, Polygonatum
multiflorum, P. officinale, P. verticillatum and Paris quadri-
folia.
b. Convallariae-Digraphidis (Sopp.) Kleb.; aecidial stage on
Convallaria majalis.
c. Paridi-Digraphidis (Plowr.) Kleb.; aecidial stage on Paris
quadrifolia.
. Puccinia Allu-Phalaridis Kleb.; aecidial stage on Allium ursinum.
3. Puccinia Orchidearum-Phalaridis Kleb.; aecidial stage on Gymna-
denia conopea, Listera ovata, Orchis maculata, O. latifolia,
Platanthera bifolia and P. Chlorantha.
4. Puccinia Ari-Phalaridis (Plowr.) Kleb.; aecidial stage on Arum
maculatum.
5. Puccinia Schmidtiana Diet.; aecidial stage on Leucojum aestivum
and L. vernum.
It may be especially noted that, according to Klebahn, Puccinia
Smilacearum-Digraphidis includes a race which occurs on a number of
hosts and two races which occur on only one host. The hosts for the
second two races are also hosts for the first race. It is further to be
noted that the species recognized are also to be distinguished on the
basis of host relations.
Puccinia Caricis (Schum.) Rebent. Klebahn (91) indicates the
following specialization in the Urtica-Carex rust, the races being indi-
cated by the choice of uredo and teleuto host:
1. Urticae-acutae on Carex acuta, C. Goodenoughi, to a less extent C.
stricta.
2. Urticae-hirtae on Carex hirta.
3. Urticae-acutiformis on Carex acutiformis, C. Kochiana, to a less
extent on C. pseudocyperus.
4. Urticae-vesicariae on Carex vesicaria.
25
iS)
366 BROOKLYN BOTANIC GARDEN MEMOIRS
Tranzschel (155) suggests the possibility of two other races, one on
Carex pallescens and a second on Carex vaginata.
Puccinia Centaureae-Caricis Tranz. Tranzschel (155) suggests
that the various Centaurea-Carex rusts may best be grouped under the
above name. The following rusts are thus included:
1. Puccinia Caricis-montanae Ed. Fischer. Bandi (15) claims to have
found evidence for the occurrence of specialization in this
heteroecious rust which forms its uredospores and teleuto-
spores on Carex montana and its aecidiospores on species of
Centaurea. He mentions two specialized races based on
the choice of aecidial host:
a. On Centaurea scabiosa.
b. On Centaurea montana.
. Puccinia tenuistipes Rostrup; aecidial stage on Centaurea Jacea;
uredo and teleuto stages on Carex muricata.
3. Puccinia arenarticola Plowr.; aecidial stage on Centaurea nigra;
uredo and teleuto stages on Carex arenaria. :
4. Puccinia Jaceae-leporinae Tranz.; aecidial stage on Centaurea
Jacea; uredo and teleuto stages on Carex leporina.
5. Puccinia Jacea-capillaris Tranz.; aecidial stage on Centaurea Jacea;
uredo and teleuto stages on Carex capillaris.
6. An unnamed rust with aecidial stage on Centaurea orientalis; uredo
and teleuto stages on Carex gynobasis.
Puccinia extensicola Plowr. This name is applied to a group of
rusts which have their aecidial stage on Compositae and their uredo
and teleuto stages on species of Carex. Arthur (2, 3, 5, 6, 8, 9, 10,
II, 13) has listed a number of forms under distinct names, but later
suggests that they are merely specialized races of Puccinia extensicola.
The following rusts are regarded as belonging here:
1. Puccinia Caricis-Erigerontis Arth.; aecidial stage on FErigeron
annuus, E. canadensis and E. philadelphicus; uredo and teleuto
stages on Carex festucaceda.
2. Puccinia Caricis-Asteris Arth.; aecidial stage on Aster acuminatus,
A. adscendens, A. cordifolius, A. paniculatus and Solidago
graminifolia; uredo and teleuto stages on Carex festiva, C. foenea
C. retrorsa, C. rosea, C. scoparia and C. trisperma.
3. Puccinia Caricis-Solidaginis Arth.; aecidial stage on Solidago caesia,
S. canadensis, S. graminifolia, S. rigida, S. serotina and S.
ulmifolia; uredo and teleuto stages on Carex Jamesti, C.
scoparia, C. sparganoides and C. stipata.
It is suggested that a distinct race may occur on Carex scoparia
and Solidago graminifolia.
Puccinia silvatica Schroet. This rust is reported as having its
Ny
REED: SPECIALIZATION OF PARASITIC FUNGI 367
uredo and teleuto stages on various species of Carex, while the aecidial
stage occurs on Taraxacum officinalis, Crepis biennis, Lappa officinalis
and three species of Senecio. Schroeter (137) proved that the aecidium
on Taraxacum was connected with the rust on Carex brizoides and C.
praecox. Later he connected the aecidium on Senecio nemorensis
with the rust on Carex brizoides. Dietel (27) connected the aecidium
on Lappa with the Carex rust and Juel (72) and Bubak (24) established
the connection between the aecidium on Crepis and the Puccinia on
Carex. Wagner (167, 169), however, claims that a particular collec-
tion of teleutospores from Carex would not infect Taraxacum, Lappa
and Senecio but only one of these aecidial hosts. Some collections of
teleutospores infect one aecidial host while other collections infect a
still different aecidial host. There is an indication, then, of a special-
ization to particular hosts in this rust.
Puccinia Ribesii-Caricis Kleb. This rust has its aecidial stage on
species of Ribes and its uredo and teleuto stages on various species of
Carex. Arthur (2, 4, 5, 6, 7, 8, 11, 13, 14), in this country, has carried
out inoculation tests with the Ribes-Carex rust for a number of years.
He finds that the aecidial hosts include Ribes aureum, R. Cynosbatt,
R. gracile, R. prostratum, R. rotundifolium and R. uva-crispa; other
species not infected are R. floridum, R. oxyacanthoides and R. rubrum.
Teleutospores were used from Carex arctata, C. crinita, C. debtlis, C.
gracillima, C. intumescens, C. pallescens, C. pubescens, C. squarrosa,
C. tenuis, and C. tetanica. Fraser (55) reports the successful infection
of Ribes oxyacanthoides with teleutospores from Carex arctata and
Carex crinita. There appears to be no indication of the existence of
specialized races.
In Europe, Klebahn (87, 91) has extensively studied the Ribes-
Carex rust and finds evidence for the existence of five specialized races.
The specialization occurs very largely in the choice of the uredo and
teleuto host, as all the races pass over to practically the same species
of Ribes as aecidial hosts. Klebahn distinguishes the following on the
basis of cultural tests:
1. Puccinia Pringsheimiana Kleb.; uredo and teleuto tages on Carex
acuta, C. caespitosa, C. Goodenough, and C. stricta; aecidial
stage on Ribes alpinum, R. aureum, R. Grossularia, R. rubrum
and R. sanguineum.
2. Puccinia Ribesti-Pseudocypert Kleb.; uredo and teleuto stages on
Carex pseudocyperus; aecidial stage on Ribes alpinum, R.
aureum, R. Grossularia, R. nigrum, R. rubrum and R. san-
guineum.
3. Puccinia Ribis nigri-Paniculatae Kleb.; uredo and teleuto stages on
Carex paniculata and C. paradoxa; aecidial stage on R. alpinum,
R. aureum, R. nigrum, R. rubrum and R. sanguineum.
368 BROOKLYN BOTANIC GARDEN MEMOIRS
4. Puccinia Magnusti Kleb.; uredo and teleuto stages on Carex acutt-
formis and C. riparia; aecidial stage on Ribes alpinum, R.
aureum, R. nigrum and R. sanguineum.
5. Puccinia Ribis nigri-acutae Kleb.; uredo and teleuto stages on
Carex acuta and C. stricta; aecidial stage on Ribes alpinum, R.
aureum, R. nigrum and R. sanguineum.
Puccinia Bistortae (Str.) DC. This rust has its uredo and teleuto
stages on species of Polygonum and its aecidial stage on various
umbellifers. It is broken up into distinct races based upon the choice
of the aecidial host. By some, these races are regarded as true species.
According to Klebahn (91) they are as follows:
1. Puccinia Angelicae-Bistortae Kleb. with the aecidial stage on
Angelica silvestris and Carum carvt.
2. Puccinia Conopodii-Bistortae Kleb. with the aecidial stage on
Conopodium denudatum.
Puccinia mammillata Schroet. This rust, also with its uredo and
teleuto stages on Polygonum, as a result of the work of Bubak (21)
and Semadeni (140), is separable into two races:
1. Puccinia Angelicae-mammillata Kleb. with the aecidial stage on
Angelica silvestris (Aecidium Bubakianum Juel).
2. Puccinia Mei-mammillata Semadeni with the aecidial stage on
Meum mutellina.
Puccinia Polygoni-amphibii Pers. Several workers claim the
existence of at least two rusts on the various species of Polygonum
on the basis of minor morphological characteristics. Puccinia Poly-
goni-amphibii Pers. is recorded on Polygonum amphibium and P.
Polygoni-Convolvuli DC. on Polygonum convolvulus. P. and H.
Sydow (151) in their monograph of the rusts, however, claim that the
differences are not sufficient to distinguish the species and consequently
list the various Polygonums as hosts for the one rust, Puccinia Poly-
goni-amphibu.
Tranzschel (152, 153) first demonstrated the heteroecism of this
Polygonum rust, connecting the uredo and teleuto stages on Polygonum
amphibium with the aecidial stage on Geranium palustre and G.
pratense. He further found that the uredo and teleuto stages on
Polygonum convolvulus was connected with the aecidial stage on
Geranium pusillum. A number of other workers have confirmed the
connection between the uredo and teleuto stages on Polygonum and
the aecidial stage on Geranium.
Jacob (66, 67) has carried out the most extensive series of cultural
experiments with the Polygonum rust, using teleutospores from Poly-
gonum amphibium, P. convolvulus and P. dumetorum. The aecidio-
spores produced experimentally on the various species of Geranium,
REED: SPECIALIZATION OF PARASITIC FUNGI 369
and also uredospores produced by aecidiosporic inoculations were
also used. As the result of the experiments, Jacob concludes that the
rusts on Polygonum amphibium and on P. convolvulus and P. dume-
torum are distinct. The relationship of these two forms may be
indicated as follows:
1. Puccinia Polygoni-amphibi Pers.; aecidial stage on Geranium
albanum, G. columbinum, G. dissectum, G. lucidum, G. molle,
G. nodosum, G. pratense, G. pusillum, G. pyrenaicum, G. rivulare,
G. rotundifolium and G. sanguineum; uredo and teleuto stages
on Polygonum amphibium.
2. Puccinia Polygoni Alb. et Schw. (P. Polygoni-Convolvuli DC.);
aecidial stage on Geranium columbinum, G. dissectum, G. molle,
G. pusillum, and G. rotundifolium; uredo and teleuto stages on
Polygonum convolvulus and P. dumetorum.
It may be noted that the same species of Geranium occur as hosts
for both races, although the host list for Puccinia Polygoni-amphibu
includes more species than the other.
Puccinia absinthii DC. Klebahn (91) suggests the probability of
specialized races in this rust although no cultural experiments have
been carried out. He reports, however, minute differences in the spore
measurements of the different forms. The following races are indicated:
1. Absinth on Artemisia absinthium.
2. Artemisiae on Artemisia vulgaris.
3. Abrotani on Artemisia abrotanum.
Puccinia bullata (Pers.) Winter. This rust, according to Semadeni
(135, 136), has a race specialized to Silaus pratensis and one to Thys-
selinum palustre.
Puccinia carduorum Jacky. Probst (112) recognizes three special-
ized races in this rust.
I. Crispi on Carduus crispus and C. personata.
2. Deflorati on Carduus defloratus.
3. A third race probably occurs on Carduus nutans.
Puccinia Centaurea Mart. Jacky (64) and Hasler (62) have both
studied the specialization of this rust. Jacky suggested the occurrence
of two specialized races: (a) Jaceae on Centaurea Jacea and (b) Ner-
vosae on C. nervosa. Hasler separates out three rusts, making them
species and finds additional host specialization in one of these. His
arrangement is as follows:
1. Puccinia Centaureae-vallesiacae Hasler on Centaurea vallesiaca, C.
alba, C. rhenana and, to a less extent, on C. cyanus.
2. Puccinia Jaceae Otth on Centaurea Jacea, C. rhenana and, to a less
extent, on Centaurea austriaca, C. Jacea var. longifolia, C.
phrygia and C. transalpina.
370 BROOKLYN BOTANIC GARDEN MEMOIRS
3. Puccinia Centaureae (Mart.) Hasler.
a. Scabtosae on Centaurea scabiosa.
b. Nigrae on Centaurea nigra.
c. Nervosae on Centaurea nervosa.
d. Transalpinae on Centaurea transalpina, C. alba, C. austriaca,
C. Jacea var. longifolia, C. nervosa, C. nigrescens and C.
phrygia.
Puccinia chaerophylli Purt. Semadeni (139, 140) claims that a
race of this rust occurs on Anthriscus silvestris and another on Chaero-
phyllum aureum.
Puccinia Epilobii-tetragoni (DC.) Winter. Dietel (29) was unable
to infect Epilobium hirsutum with aecidiospores from E. tetragonum.
This is the only indication of specialization in this rust.
Puccinia Geranii-silvatici Karst. This rust has been reported on a
few species of Geranium in widely separated localities. It occurs
commonly on Geranium silvaticum in Europe. Its restriction to widely
separated regions has led to the suggestion that the rust consists of
geographically specialized races. Jacob (66, 67) has shown by cul-
tural experiments that there is no specialization in Europe, for both
Geranium silvaticum and G. rotundifolium are readily infected by
teleutospores from the former.
Puccinia Helianthi Schw. Arthur, Kellerman, and Jacky have
made inoculation tests using teleutospores of this rust. Jacky (65),
using teleutospores from Helianthus annuus, infected’H. annuus, H.
cucumertfolius, and H. californicus, but failed to infect H. maximiliani,
HA. multiflorus, H. rigidus, H. scaberrimus and H. tuberosus.
Kellerman (74, 75) obtained negative results with teleutospores
from H. annuus on eighteen species of Helianthus. His results with
teleutospores from H. ambiguus were negative on twelve species and
also negative with teleutospores from H. decapetalus on eight species.
He found, however, that teleutospores from H. mollis infected H.
annuus and H. mollis but not fourteen other species; teleutospores
from HH. grosse-serratus infected H. annuus, H. decapetalus, H. giganteus,
H. grosse-serratus, H. Kellermani, H. orygalus and H. tracheifolius,
but not H. maximiliani nor H. mollis; teleutospores from H. tuberosus
infected only H. annuus.
Arthur (3, 4, 5, 6) secured the following results: (1) Teleutospores
from Helianthus mollis infected H. annuus, H. hirsutus, H. mollis,
H. occidentalis, H. strumosus and H. tomentosus, but not H. grosse-
serratus, H. Kellermani, H. laetiflorus, H. longifolius, H. orygalus and
HI. tuberosus; (2) teleutospores from H. grosse-serratus infected H.
annuus, EH. grosse-serratus, H. maximiliani and H. tomentosus but not
H. decapetalus, H. hirsutus, H. laetiflorus, H. mollis, H. occidentalis,
REED: SPECIALIZATION OF PARASITIC FUNGI 371
H. orygalus, H. strumosus nor H. tuberosus; (3) teleutospores from
H. laetiflorus infected H. annuus, H. divaricatus, H. Kellermani, H.
laetiflorus, H. mollis, H. occidentalis and H. tomentosus, but not H.
grosse-serratus, H. hirsutus, H. orygalus, H. strumosus, nor H. tuberosus.
About the only conclusion that one can draw is that H. annuus
and H. tomentosus are readily infected with teleutospores from a variety
of hosts. The evidence for distinct specialized races is not very clear.
Tranzschel (158) states that he was able to infect Helianthus annuus
with teleutospores from Xanthium strumarium; the aecidiospores
produced on Helianthus annuus infected both H. annuus and Xan-
thium strumarium.
Puccinia Hieracii (Schum.) Mart. Jacky (64) first investigated
this rust and found some evidence for specialization, distinguishing
one race on Mieracium villosum and suggesting the possibility of several
others. Probst (113), more recently, has studied the rust and has
come to the conclusion that there are two subspecies, each with several
specialized races. He arranges the forms as follows:
1. Puccinia Piloselloidarum Probst.
. Hoppeani on Meracium hoppeanum.
. Peleteriant on Hieracium peleterianum.
. Pilosellae (a) on Mieracium pilosella.
. Pilosellae (6) on Hieracium pilosella.
Velutini on Mieracium pilosella velutinum.
Auriculae on Hieracium auricula and H. peleterianum.
Floretint on Hieracium florentinum var. obscurum.
h. Ziziant on Hieracium bauhini, H. florentinum var. alethes,
H. pratense and H. zizianum. :
2. Puccinia Mieracit.
a. Silvatict on Hieracium glaucum, H. humile, H. ochroleucum,
HI. pictum, H. silvaticum and H. trebevicianum.
Silvatict pleiotricht on Hieracium silvaticum var. pleiotrichum.
Silvatict gentili on Hieracium folcanum and H. intybaceum.
d. Schmidt on Hieracium Schmidtii, H. humile and H. ochro-
leucum.
e. Cinerascentis on Hieracium cinerascens and H. ochroleucum.
Puccinia Leontodontis Jacky. Probst (112) states that three
specialized races occur in this rust:
1. Hispidi on Leontodon hispidus.
2. Autumnalis on Leontodon autumnalis.
3. Pyrenaict on Leontodon pyrenaicus.
Puccinia petroselini (DC.) Lindr. Semadeni (139, 140) dis-
tinguishes two specialized races of this rust, one on Aethusa cynapium
and the other on Petroselinum sativum.
va Sh O Qaee
ao So
372 BROOKLYN BOTANIC GARDEN MEMOIRS
Puccinia Pulsatillae Kalchbr. Bubak (22) distinguishes the fol-
lowing races of this rust, basing the separation on the distribution and
character of the sorus, not on cultural experiments:
. Concortica on Pulsatilla alpina and P. sulphurea.
. Atragenicola on Atragene alpina.
Genuina on Anemone silvestris and Pulsatila patens.
Pulsatillarum on Pulsatilla pratensis and P. vulgaris.
Puccinia Ribis DC. Eriksson (39) distinguishes a specialized race
Rubri, for he found that teleutospores from Ribes rubrum would infect
this species but not R. nigrum nor R. Grossulania.
Uromyces alchimillae (Pers.) Lév. Fischer (53) has carried out a
few experiments with this rust which tend to show the existence of a
host specialization. The results reported, however, do not indicate
any close correspondence between the plants infected with a particular
collection of uredospores and the systematic grouping of the host
plants within the genus Alchimilla. Fischer found that uredospores
from hosts belonging to the section Vulgares infected plants belonging
to sections Pubescentes and Splendentes.
Uromyces caryophyllinus (Schrank) Winter. Fischer (49-51)
gives experimental evidence for the occurrence of specialized races in
the carnation rust. The rust is heteroecious, the aecidial stage oc-
curring on Euphorbia Gerardiana and the uredo and teleuto stages on
various Caryophyllaceae. Fischer finds that aecidiospores from
Euphorbia Gerardiana in one locality are able to infect only Tunica
prolifera, while aecidiospores collected on the same host in another
region are able to infect only Saponaria ocymoides. In still other
localities, however, a race of rust is found that is able to infect both
Tunica and Saponaria.
Uromyces Dactylidis Otth and U.Poae Rabh. These rusts,
probably indistinguishable by any well-defined structural character-
istics, are further alike in having their aecidial stage on species of
Ranunculus. The former develops its uredo and teleuto stages on
Dactylis glomerata, while the latter has the corresponding stages on
several species of Poa, Agrostis alba also being listed asa host. A large
number of workers have contributed to our knowledge concerning the
heteroecism of these rusts.
Krieg (93, 94) and Klebahn (89, 91) have worked with Uromyces
Dactylidis from the standpoint of specialization and the following
races are indicated:
1. Aecidial stage on Ranunculus bulbosus and R. repens.
2. Aecidial stage on Ranunculus lanuginosus, to a slight extent on
R. bulbosus (Uromyces lanuginosi-dactylidis Kleb.).
3. Aecidial stage on Ranunculus aconitifolius, R. alpestris, R. glacialis,
and R. platanifolius (Uromyces platanifolii-dactylidis Krieg.).
Oo N
REED: SPECIALIZATION OF PARASITIC FUNGI 373
4. Aecidial stage on Ranunculus silvaticus (Uromyces silvatici-dactylidis
Krieg.).
5. Aecidial stage on Ranunculus acer and R. polyanthemos.
Juel (73) distinguishes several forms in Uromyces Poae:
1. Ficariae-nemoralis on Ranunculus Ficaria and Poa nemoralis.
2. Ficariae-trivialis on Ranunculus Ficaria and Poa nemoralis and
P. palustris.
3. Ficariae-pratensis on Ranunculus Ficaria and Poa pratensis.
4. Repentis-nemoralis on Ranunculus repens and R. bulbosus and Poa
nemoralis.
5. Repentis-trivialis on Ranunculus repens and Poa trivialis and P.
annua.
6. Auricomi-pratensis on Ranunculus auricomus and Poa pratensis and
P. nemoralis.
Cassubici-pratensis on Ranunculus cassubicus and Poa pratensis.
Repentis-pratensis on Ranunculus repens and Poa pratensis.
9. Bullati-bulbosae on Ranunculus bullatus and Poa bulbosa.
Uromyces Fabae (Pers.) de Bary. Jordi (70, 71) has distinguished
the following specialized races on this autoecious rust which is recorded
as occurring on species of Lathyrus, Lens, Pisum, and Vicia:
1. On Lathyrus vernus and probably on Pisum sativum.
2. On Vicia Faba and Pisum sativum.
3. On Vicia cracca, V. hirsuta and Pisum sativum.
It should be noted that Pisum sativum is listed as a host for all
three specialized races.
Uromyces Geranii (DC.) Otth and Wartm., and U. Kabatianus
Bubak. The first mentioned species is reported as occurring on a
large number of different species of Geranium, while the second men-
tioned has been reported only on Geranium pyrenaicum. Bubak (23)
has called attention to minor morphological differences between the
two rusts.
Bock (19) carried out tests with uredospores of Uromyces geranii
from Geranium silvaticum and was able to infect sixteen species while
thirteen gave negative results. Jacob (66, 67) also conducted experi-
ments with the rust, using teleutospores from Geranium silvaticum and
aecidiospores produced from the teleutosporic inoculation. A con-
siderable number of species of Geranium were successfully infected.
A few gave negative results. Jacob also used teleutospores of Uro-
myces Kabatianus from Geranium pyrenaicum and found that this
rust could also be transferred to a considerable number of other species
of Geranium. The host range of the two rusts is very similar and the
differences between the two depend upon minor structural differences
and not on host relations.
coon]
374 BROOKLYN BOTANIC GARDEN MEMOIRS
Uromyces Pisi (Pers.) Winter. This rust is heteroecious forming
its aecidial stage on various species of Euphorbia and its uredo and
teleuto stages on species of Lathyrus and Pisum. The aecidial my-
celium is perennial in the Euphorbia host. It may also be noted that
aecidial stages on Euphorbia, especially E. cyparissias, have been
connected with a number of different species of Uromyces on legumes.
The aecidia belonging to these different rusts are quite indistinguish-
able by structural features. Jordi (70, 71) suggests that specialized
races of U. Pisi occur on Lathyrus pratensis and Vicia cracca; the
latter race is by some recognized as a species—Uromyces Fischeri-
Eduardi.
Uromyces proeminens (DC.) Lév. P. and H. Sydow (151) list
twenty-eight species of Uromyces on the species of Euphorbia, basing
their separations largely on the studies of Tranzschel (157) in this
group of rusts. Nine species are recorded as autoecious and have all
four spore-forms present, the others either being short-cycled or the
life history incompletely known. The structural differences between
many of these species are very slight and the characters used as a basis
for separation are, in many cases, quite variable. In fact Arthur (12)
has grouped several of the full-cycled forms under the one species
Nigredo proeminens, suggesting the occurrence of specialized races.
Arthur (1, 2, 3) has obtained the following results with inoculation
experiments: aecidiospores from Euphorbia nutans infected E. nutans
but not EF. maculata, E. marginata nor E. humistrata; aecidiospores
from, E. humistrata iffected E. humistrata and E. nutans but not
E. maculata; uredospores from E. dentata infected E. dentata but not
E. humistrata, E. nutans nor E. marginata; uredospores from E. nutans
infected EL. nutans but not E. maculata.
Arthur (12) suggests that a race is restricted to the section Poin-
seltia, a second race to the section Dichrophyllum, a third race to the
prostrate species, and the fourth to the more upright species of the
section Chamaesyce of the genus Euphorbia.
Uromyces Scirpi (Cast.) Burr. This rust has its uredo and teleuto
stages on Scirpus maritimus and its aecidial stage on Glaux maritima,
IMppuris vulgaris, Berula angustifolia, Daucus carota, Oenanthes
aquatica, O. crocata, Pastinaca sativa and Sium latifolium in Europe.
The similar rust in this country forms its uredo and teleuto stages on
Scirpus americanus, S. campestris, S. fluviatilis and S. robustus, and
its aecidial stage on Cicuta bulbifera, C. maculata, Glaux maritima,
Oenanthes Californica and Sium cicutaefolium. A good deal of experi-
mentation has been carried on by Klebahn and others to determine the
host relations of the rust and various races have been separated out
as distinct species. Klebahn (89, 91) distinguishes the following:
REED: SPECIALIZATION OF PARASITIC FUNGI a75
1. Uromyces Pastinacae-scirpi Kleb.; aecidial stage on Pastinaca
sativa.
2. Uromyces Berulae-scirpi Kleb.; aecidial stage on Berula angusti-
folia. a
Dietel (28) was able to infect with teleutospores from a single host
Sium latifolium and Hippurus vulgaris.
Gymnosporangium tremelloides Hartig. There is but little indi-
cation in the literature of the existence of specialized races in Gymno-
sporangium. Most of the species of this genus are quite restricted in
host range, often occurring on only a few species of a single genus of
host. Klebahn (89) gives some evidence for a host specialization of
Gymnosporangium tremelloides, the teleutospores of which are produced
on Juniperus communis. On the basis of cultural tests he suggests the
following:
1. Gymnosporangium Ariae-tremelloides Kleb. Aecidial stage on
Sorbus aria and S. torminalis.
2. Gymnosporangium Mali-tremelloides Kleb. Aecidial stage on Pyrus
malus.
Ochrospora Ariae (Fuckel.) Syd. Fischer (45) stated that he was
unable to infect Aruncus silvestris with uredospores of Ochrospora ariae
from Sorbus aucuparia. Tranzschel (152, 153), Fischer (45) and
Klebahn (89) showed that Sorbus aria, S. scandica and Pyrus malus
were all hosts of the same race, but Aruncus silvestris could not be
infected with spores from the same source. Later, however, Fischer
(48) was able to infect Sorbus aucuparia and Aruncus silvestris with
the same aecidiosporic material from Anemone nemorosa.
Pucciniastrum Abieti-Chamaenerii Kleb. and P. Epilobii (Pers.)
Otth. These two rusts, according to Klebahn (83, 87), are dis-
tinguished almost solely on the basis of their life cycle. The former
is heteroecious with its aecidial stage on Abies balsamea and A. pectinata
and its uredo and teleuto stages on Epilobium angustifolium, E.
Dodonacum and E. latifolium. The latter has its uredo and teleuto
stages on a number of other species of Epilobium and its aecidial host
is as yet not known.
Phragmidium disciflorum (Tode.) James. Bandi (15) suggests
that a host specialization occurs in this rust. He reports one race on
Rosa cinnamomea, R. pimpinellifolia and R. rubrifolia and a second
race on R. canina and R. centifolia.
Coleosporium Campanulae (Pers.) Lév. This rust has its aecidial
stage on Pinus montana, P. rigida and P. sylvestris, and its uredo and
teleuto stages on a considerable number of different species and general
of the Campanulaceae. Wagner (170) and Klebahn (87, 91) have
made extensive studies with reference to the specialization of the rust
on different hosts. Klebahn makes the following races:
376 BROOKLYN BOTANIC GARDEN MEMOIRS
1. Campanulae-rapunculoides Kleb. on Campanula rapunculoides and,
in favorable cultures, on C. bononiensis, C. glomerata, C.
glomerata var. dahurica, C. lamiifolia, C. latifolia, C. nobilis,
Phyteuma orbiculare and Ph. spicatum; not on Campanula
rotundifolia, nor C. trachelium.
. Campanulae-tracheliit Kleb. on Campanula trachelium and, in favor-
able cultures, on C. bononiensis, C. glomerata, C. glomerata var.
dahurica, C. latifolia var. macrantha, C. nobilis, C. rapunculoides
and Wahlenbergia hederacea; not on Campanula pusilla, C.
rotundifolia nor C. turbinata.
3. Campanulae-rotundifoliae Kleb. on Campanula rotundifolia and, in
favorable cultures, on C. bonontensis, C. glomerata var. dahurica,
C. pusilla, C. turbinata, Phyteuma orbiculare, Ph. spicatum and
Wahlenbergia hederacea; not on Campanula rapunculoides nor
C. trachelium.
Wagner indicated the following additional races:
4. Campanulae-Phyteumatis Wagner on Phyteuma spicatum.
Campanulae-macranthae Wagner on Campanula macrantha (C.
latifolia var. macrantha).
6. Campanulae-patulae Wagner on Campanula rotundifolia and C.
patula.
Klebahn, however, does not believe that these can be distinguished
from the first three. It may also be noted that a number of plants
occur as hosts for more than one race.
Coleosporium Senecionis Fr. Wagner (168) first gave evidence of
host specialization in this rust. Fischer (54) has also investigated the
rust and the following races are indicated:
1. Senecionts silvatict on Senecio silvaticus, S. viscosus and S. vulgaris
(Senecionis I of Wagner).
. Senecionis Fuchsti on Senecio Fuchsii and S. nemorensis (Senecionis
II of Wagner).
3. Senecionis subalpini on Senecio subalpinus (Coleosporium subalpinum
Wagner).
4. Senecionis doronici on Senecio doronicum.
Melampsora Euphorbiae (Schub.) Cast. This rust, completing its
life cycle on species of Euphorbia, has been studied by W. Miller
(105, 106) who claims the existence of the following races:
1. Euphorbiae-cyparissiae W. Miiller on Euphorbia cyparissias.
2. Euphorbiae-exiguae W. Miiller on Euphorbia exigua.
3. Euphorbiae-pepli W. Miiller on Euphorbia peplus.
Melampsora Euphorbiae dulcis Otth. This autoecious rust on
Euphorbia is very similar to Melampsora Euphorbiae. As a result of
the experimental work of W. Miiller (105, 106) and Klebahn (gt) two
races are indicated:
NO
on
i)
REED: SPECIALIZATION OF PARASITIC FUNGI 37T.
1. Euphorbiae-dulcis s. str. on Euphorbia dulcis.
2. Euphorbiae-strictae W. Miiller on Euphorbia stricta and E. platy-
phyllos.
Melampsora populina Lév. and Melampsora Tremulae Tul. A
number of different species of Melampsora are recorded as having their
uredo and teleuto stages on the various kinds of poplars. These rusts,
however, constitute a group of closely related forms and differ but
little in their structural characteristics. The main differences appear
to be in the choice of hosts, especially in the aecidial stage.
The relationships between the poplar rusts may best be indicated
by grouping them under the above names. Melampsora populina
is distinguished from M. Tremulae by the fact that the teleutospores
are subcuticular while in the case of M. Tremulae they are subepidermal.
The uredo and teleuto hosts are also different species of Populus and
serve further as a means of distinguishing between the two. Klebahn
(87, 91) has made a special study of these rusts.
Melampsora populina includes two rusts which can be distinguished
only by the choice of the aecidial host. These are:
1. Melampsora Allii-populina Kleb.; aecidial stage on Allium cepa
and A. ursinum; uredo and teleuto stages on Populus balsamif-
era and P. nigra.
2. Melampsora Larici-populina Kleb.; aecidial stage on Larix decidua;
uredo and teleuto stages on Populus balsamifera and P. nigra.
Melampsora Tremulae includes four or five rusts whose uredo and
teleuto stages occur on Populus alba and P. tremula, rarely on other
species; the aecidial stage is found on widely separated host plants.
The following belong in this group:
1. Melampsora Larici-Tremulae Kleb.; aecidial stage on Larix decidua.
2. Melampsora pinitorqua Rostr.; aecidial stage on Pinus silvestris.
3. Melampsora Rostrupii Wagner; aecidial stage on Mercurialis
perennis.
4. Melampsora Magnusiana Wagner; aecidial stage on Chelidonium
major.
5. Melampsora Klebahni Bubak; aecidial stage on Corydalis cava, C.
digitata, C. fabacea, C. laxa and C. solida. This may not be
distinct from the preceding one.
Melampsoras of Salix. The Melampsoras on different species of
Salix constitute a complex group of interrelated rusts. The structural
differences between the large number of commonly recognized species
are comparatively insignificant. In order to segregate the different
species it is necessary to rely, to a large extent, upon differences in
the choice of host. The willow rusts also afford many parallels among
the poplar rusts and in some cases it is not possible to distinguish the
rusts on these two genera except by the choice of host.
378 BROOKLYN BOTANIC GARDEN MEMOIRS
Klebahn (87, 91) and Schneider (133-135) have been the principal
investigators of the willow rusts from the standpoint of host special-
ization. While many forms have been segregated out, Klebahn (92)
has recently indicated the value of grouping these on the basis of
structural features. The relationship of some of these different willow |
rusts, based upon structural characteristics and physiological special-
ization, may be indicated.
Melampsora Larici epitea Kleb. The aecidial stage occurs on
Larix decidua and the uredo and teleuto stages on various species of
Salix. Klebahn (87, 91) and Schneider (133-135) have carried out a
number of inoculation tests and have distinguished the following
races:
a. Larici-epitea typica Kleb. on Salix aurita, S. cinerea, S. caprea,
S. bppophaéfolia and S. viminalis.
b. Larici-daphnoides Kleb. on Salix daphnoides.
‘c. Larici-retusae Ed. Fischer on Salix retusa and S. herbacea.
d. Larici-nigricantis Schneider on Salix nigricans, S. glabra and S.
hegetschweilert.
e. Larici-purpureae Schneider on Salix purpurea.
f. Larict-reticulatae Schneider on Salix reticulata and S. hastata.
Klebahn states that minute structural differences can be observed
in. @, 6, and. c.
Melampsora Ribesii-purpureae Kleb. The aecidial stage occurs
on species of Ribes and the uredo and teleuto stages on various species
of Salix. Specialized races occur, according to Klebahn (87, 91) and
Schneider (133-135), as follows:
a. Ribesvi-purpureae Kleb. on Salix purpurea and S. purpurea X S.
viminalis; to a less extent on S. daphnoides.
b. Ribesti-auritae Kleb. on Salix aurita and possibly on S. Caprea
and S. cinerea.
c. Ribesit-grandifoliae Schneider on Salix grandifolia and S. aurita,
and possibly on S. arbuscula.
Klebahn places (a) and (b) under a separate species Melampsora
Ribesti-epitea Kleb.
Melampsora Evonymi-Capraearum Kleb. The aecidial stage
appears on species of Evonymus and the uredo and teleuto stages on
various species of Salix. Schneider (133-135) distinguishes two spe-
cialized races:
a. Evonymi-capraearum typica Schneider on Salix aurita, S. Caprea,
and S. cinerea.
b. Evonymi-incanae Schneider on Salix incana and S. caprea, not on
S. aurita nor S. cinerea.
Melampsora Larici-pentandrae Kleb. and M. Salicis-albae Kleb.
REED? SPECIALIZATION OF PARASITIC FUNGI awe
According to Klebahn (84, 87, 91), Melampsora Larici-pentandrae
Kleb., with its aecidial stage on Larix decidua and uredo and teleuto
stages on Salix fragilis and S. pentandra, is hardly distinguishable
from Melampsora Salicis-albae Kleb., with its aecidial stage on species
of Allium and its uredo and teleuto stages on Salix alba, except on the
basis of host relations.
Melampsora Allii-fragilis Kleb. and M. Galanthi-fragilis Kleb.
According to Klebahn (84, 85, 87, 91), a similar relation exists between
Melampsora Allii-fragilis Kleb., with its aecidial stage on species of
Allium and uredo and teleuto stages on Salix fragilis and S. pentandra,
and Melampsora Galanthi-fragilis Kleb., with its aecidial stage on
Galanthus niwalis and uredo and teleuto stages on the same two species
of Salix.
Melampsorella Caryophyllacearum (DC.) Schroet. Bubak (24),
Klebahn (87), and Fischer (45) have given some evidence for special-
ization in this rust. Fischer, using the same collection of aecidio-
spores, infected Stellaria media, S. graminea, Arenaria serphyllifolia.
Malachium aquaticum and Cerastitum sp. indet., while Cerastiwm
arvense, Moehringia trinervia and M. mucosa were not infected. Kle-
bahn obtained somewhat similar results.
Melampsoridium betulinum (Tul.) Kleb. Klebahn (87, 91) sug-
gests the occurrence of at least two specialized races of this rust which
has its aecidial stage on Larix decidua and its uredo and teleuto stages
on species of Betula. He found that aecidiospores from Larix, pro-
duced by inoculation with teleutospores from Betula pubescens, in-
fected B. pubescens and B. nana very abundantly and B. verrucosa
sparingly; aecidiospores from Larix, produced by inoculation with
teleutospores from Betula verrucosa, infected B. verrucosa abundantly,
B. nana sparingly, and B. pubescens not at all. Accordingly the fol-
lowing races are indicated:
1. Betulae-verrucosae Kleb. on Betula verrucosa and B. nana.
2. Betulae-pubescentis Kleb. on Betula pubescens, B. nana and, to a
slight extent, on B. verrucosa.
The above review lists the occurrence of host specialization in
more than fifty different rusts. The nature of the specialization of
these rusts may be indicated by arranging them in the following
groups:
1. Heteroecious rusts in which the aecidial host (or hosts) is common
to several specialized races which occur on the uredo and teleuto
hosts: Coleosporium Campanulae (Pers.) Lév., C. Senecionis
Fr., Melampsora Evonymi-Caprearum Kleb., M. Larici-epitea
Kleb., M. Ribesti-purpureae Kleb., Melampsorella Caryo-
phyllacearum Schroet., Melampsoridium betulinum (Tul.) Kleb.,
380 BROOKLYN BOTANIC GARDEN MEMOIRS
Ochropsora Ariae (Fuck.) Syd., Puccinia graminis Pers., P.
Caricis (Schum.) Rebent., P. Polygoni-amphibu Pers., Uro-
myces caryophyllinus (Schrank) Winter, and Uromyces Pisi
(Pers.) Winter.
Certain other rusts might also be looked for in this group as
Puccinia coronata (Corda) Kleb., P. loli Niels. (P. coronifera Kleb.)
and P. bromina Eriks.
2. Heteroecious rusts in which the uredo and teleuto host (or hosts)
is common to several races which occur on the aecidial hosts:
Gymnosporangium tremelloides Hartig, Melampsora populina
Lév., M. Tremulae Tul., Puccinia Bistortae (Str.) DC., P.
mammullata Schroet., P. sessilis Schneid., P. silvatica Schroet.,
Uromyces Dactylidis Otth, U. Scirpi (Cast.) Burr.
3. Heteroecious rusts in which the specialized races are recognized by
their selection of both aecidial and uredo and teleuto hosts:
Puccinia Centaureae-Caricis Tranz., P. coronata Corda, P.
dispersa Eriks. and Henn., P. extensicola Plowr., P. glumarum
(Schm.) Eriks. and Henn., P. Ribesii-Caricis Kleb., P. Stipina
Tranz., Uromyces Poae Rebent.
4. Autoecious rusts and those whose life history is incompletely
known. The following belong in this group: Melampsora
Euphorbiae (Schub.) Cast., M. Euphorbiae-dulcis Otth, Phrag-
midium disciflorum (Tode.) James, Puccinia Absinthi DC..,
P. bullata (Pers.) Winter., P. Carduorum Jacky, P. Centaureae
Mart., P. Chaerophyllt Purt., P. Epilobt-tetragoni (DC.)
Winter., P. Heltantht Schw., P. Hiteracu (Schum.) Mart.,
P. Leontodontis Jacky, P. Petroselint (DC.) Lindr., P. Pulsatillae
Kalchbr., P. Ribis DC., Uromyces Fabae (Pers.) de Bary, U.
proeminens (DC.) Lév.
POWDERY MILDEWS—ERYSIPHACEAE
The first work carried on to determine whether a host special-
ization occurs among the species of the powdery mildews was that
of Neger (107), the results of which were published in 1902. Since
then a number of workers have contributed to the evidence for special-
ized races in this well-defined group of parasites. In fact, at the pres-
ent time, one or more species of five of the six genera of the Ery-
siphaceae have been tested. In most cases, however, the data are
very meager and it is not possible to draw any definite conclusions.
In a few cases, notably for Erysiphe graminis and E. cichoracearum,
the facts are better established.
Erysiphe cichoracearum DC. The host relations of this species
was first reported upon by Neger (107) who obtained the following
REED: SPECIALIZATION OF PARASITIC FUNGI 381
results: (1) conidia from Artemisia vulgaris, Lactuca muralis and
Lithospermum arvense infected plants of the same species but did not
infect each other nor some other plants tested; (2) conidia from
Hieracium murorum infected H. murorum and Leontodon taraxacum,
the latter very slightly; (3) conidia from Senecio vulgaris infected
S. vulgaris and Lactuca muralis; (4) conidia from Lappa major,
Plantago major, Pulmonaria officinalis and Verbascum thapsiforme
failed to infect any host inoculated.
Salmon (128) reports the successful infection of Plantago major
and P. media, using conidia from the former; negative results were
obtained with Plantago lanceolata, Eupatorium cannabinum and
Galium A parine.
The writer (115, 116) has carried on a very extensive series of
experiments with this mildew, particularly with the cucurbit hosts.
In the main, the mildew as found on the Hubbard squash (Cucurbita
maxima) was used for inoculating the various plants. It was found
that this mildew readily infected Cucurbita maxima (seven varieties),
_C. moschata (three varieties), C. pepo (seventeen varieties), C. foeti-
dissima, Cucumis dipsaceus, C. melo (nine varieties), C. sativus (eight
varieties), Cyclanthera explodens, Echinocystis lobata, Lagenaria vulgaris
(six varieties), Momordica charantia and Sicyos angulatus. Partial
infection of the following was obtained: Citrullus vulgaris (seven
varieties), Cucumis anguria (two varieties), Ecballium elaterium,
Melothria scabra and Momordica balsamina. Coccinea cordifolia,
Luffa acutangula and L. Aegyptiaca proved to be entirely resistant.
To a slight extent it was possible to transfer the cucurbit mildew to
the sunflower (Helianthus annuus) and plantain (Plantago rugellit).
Efforts to transfer it to Aster cordifolius, A. laevis and Solidago caesia
failed. It was also found that a race of mildew occurred on Aster
cordifolius, A. laevis and A. sagittifolius, not passing over to Cucurbita
maxima nor Solidago caesia. Another race occurs on Solidago caesia,
infecting this species but not passing over to asters or cucurbits.
Erysiphe graminis DC. Marchal (97) was the first to demonstrate
host specialization in this powdery mildew. Asa result of his tests he
concluded that the following races may be distinguished, although he
gives us no details of his evidence:
1. Tritici upon Triticum vulgare, T. Spelta, T. polonicum and T.
turgidum; not on T. durum, T. monococcum nor T. dicoccum.
2. Hordei upon Hordeum hexastichon, H. vulgare, H. trifurcatum, H.
nudum, H. jubatum and H. murinum; not on H. maritimum,
H. secalinum nor H. bulbosum.
3. Secalis upon Secale cereale and S. anatolicum.
4. Avenae upon Avena sativa, A. fatua, A. orientalis and Arrhenatherum
elatius.
26
382 BROOKLYN BOTANIC GARDEN MEMOIRS
5. Poae upon Poa annua, P. trivialis, P. pratensis, P. caesia, P.
mutalensis, P. nemoralis and P. serotina.
6. Agropyri upon Agropyron.
7. Bromi upon various species of Bromus.
Salmon (123, 128) has infected Avena nuda, A. brevis and A. sativa
with conidia from A. nuda; conidia from A. sterilis infected A. pratense
and A. sativa; and conidia from A. sativa infected A. sativa, A. brevis,
A. nuda, A. orientalis, A. sterilis and A. strigosa. Attempts to infect
twelve other grasses, belonging to different genera, with the oat mildew
failed.
The writer (120) has carried out an extensive series of experiments
with the powdery mildew on Avena sativa. Infection occurred on the
following: A. barbata, A. brevis, A. fatua, A. fatua var. glabrata, A.
ludoviciana, A. nuda, A. nuda var. chinensis, A. nuda var. elegantissima
A. planiculmis, A. pratensis, A. pubescens, A. purpurea, A. sativa
(sixteen varieties), A. sativa orientalis (six varieties), A. sterilis, A.
strigosa and A. sulcata. In most cases, in a large number of trials,
one hundred percent of infection was secured. The oat mildew also
infected Arrhenatherum avenaceum. Negative results were obtained
with Avena bromoides and A. sempervirens, as well as grasses belonging
to other genera. This race, then, extends over a wide range of species
and varieties of Avena, but, with the exception of Arrhenatherum
avenaceum, is restricted to this genus.
Salmon (123, 132) has reported only a few results with the powdery
mildew of wheat. He successfully infected Triticum vulgare and T.
Spelta with conidia from the former. He also states that young
seedlings of Hordeum silvaticum could be infected with the same
mildew.
Vavilov (164) has tested out, under field conditions, seven hundred
and fifty-five ‘‘pure-lines’’ belonging to the different species and
varieties of Triticum. He used pure lines belonging to thirty varieties
of Triticum vulgare, seven of T. compactum, ten of T. turgidum, nine
of T. Spelta, fifteen of 7. durum, three of T. polonicum, five of T.
dicoccum and four of T. monococcum. In general the pure lines be-
longing to the different varieties of 7. vulgare, T. compactum and
T. Spelia are extremely susceptible to the mildew, while the pure
lines of the varieties of T. durum, T. turgidum, T. polonicum and
IT’. monococcum proved, in the main, to be quite resistant. The pure
lines of some varieties of 7. dicoccum proved to be highly susceptible,
while the pure lines of other varieties were markedly resistant. Dif-
ferences in the susceptibility of the pure lines were noted and certain
varieties, notably T. vulgare var. fuliginosum and T. dicoccum var.
picnurum stood out as distinctly immune. However, the pure lines
REED: SPECIALIZATION OF PARASITIC FUNGI 383
belonging to the various varieties of all the species of Triticum, with
the exception of certain ones, as those just noted, proved quite sus-
ceptible when they were tested under greenhouse conditions. Vavilov
suggests that the greenhouse conditions are much more favorable to
the development of the powdery mildew and thus more or less immune
races may be successfully attacked by the fungus.
The writer (117, 118, 120) has carried on a very extensive series
of tests with the powdery mildew of the wheat. One hundred and
sixty-one varieties belonging to the eight recognized types or species
of Triticum have been tested under greenhouse conditions, many tests
having been made with nearly all of these. The results are summarized
in the following table:
VARIETIES OF SPECIES OR TYPES OF Tyviticum IN RELATION TO THE WHEAT
MILDEW
: Aa fe 100% infec- 90-99% 50-89% | 10-49% o-9%
Species of type Varieties tion infection infection infection infection
Compacium .. 2s. 625s: 6 2 I 2 I fo)
MDGCOCCUINE | ow ice ss os 24 8 3 6 B 4
DET OY en 45 36 fe) 6 2 I
Monococcum......... 6 (0) I 3 I I
POLONICUM s 0 . 33/33/34/34 38/47| 0/28 27/27 | 0/13| 9/22! | 10/16
3. Bromus commutatus... . 0/7 | o/9 |21/21| 0/7 | 4/8%| 0/7 | 6/7 6/7
4. Bromus arvensis....... 0/9 8/8
5. Bromus tectorum....... | 8/8 | 7/8
_ From his data, Salmon concluded that four, or possibly even five,
specialized races exist within this genus. In looking over his data,
however, there seems to be very little, if any, difference between some
of them. The races on B. interruptus and B. hordeaceus differ only
in their capacity for infecting B. commuiatus, the mildew on B. horde-
aceus infecting this host, while that on B. interruptus does not. The
race on B. commutatus differs from that on B. hordeaceus in not in-
fecting B. mollis and B. interruptus. The race on B. arvensis infects
this same species but not B. mollis. Finally the race on B. tectorum
differs from that on B. hordeaceus in being able to infect B. sterilis.
It is evident that these races are not distinctly marked off from one
another.
But little work has been done using ascospores from various hosts.
Marchal (98) mentions the following results: (1) ascospores from
Hordeum vulgare infected H. vulgare, H. distichon, H. trifurcatum and
H. Zeocriton, but not Avena sativa, Secale cereale nor Triticum vulgare;
(2) ascospores from Secale cereale infected S. cereale, but not Hordeum
vulgare nor Triticum vulgare; (3) ascospores from Triticum vulgare
infected T. vulgare but not Agropyron caninum, Avena sativa, Hordeum
vulgare nor Secale cereale. Salmon (124, 132) found that ascospores
from Hordeum vulgare infected H. vulgare, H. trifurcatum and H.
Zeocriton, but not H. bulbosum, H. jubatum, H. maritimum, H. secali-
num, Avena sativa, Secale cereale nor Triticum vulgare; ascospores from
Bromus commutatus infected B. commutatus and B. hordeaceus, but not
‘0'The denominator indicates the number of leaves inoculated, the numerator
indicates the number infected.
1 Subinfection.
386 BROOKLYN BOTANIC GARDEN MEMOIRS
B. racemosus. These results correspond exactly with the infecting
capacity of conidia from the same hosts.
The evidence is quite conclusive for the existence of highly special-
ized races in the grass mildew. For the most part these races are
definitely restricted to the species of a single genus of host plants.
Erysiphe Galeopsidis DC. Neger (107) tested the infecting
capacity of the mildew on Galeopsis tetrahit and found that this host
was infected, while negative results were obtained on Calamintha
acinos, Glechoma hederacea and Stachys recta. Salmon (128) obtained
positive results with the mildew from Ballota nigra on this same host
but failed to infect Salvia verticillata and Leonurus cardiaca.
Erysiphe Polygoni DC. Neger (107) used the mildew from the
following hosts: Galium silvaticum, Heracleum spondylium, Hypericum
perforatum, Ranunculus repens and Trifolium incarnatum. Positive
results were obtained when the mildew was sown on plants of the same
host from which it was obtained and negative results on all other hosts
tested. In one case he noted a slight infection of Galium silvaticum
with conidia from Ranunculus repens but this was probably a foreign
infection. .
Salmon (123) successfully infected Pisum arvense with conidia from
P. sativum. Other legumes gave negative results. Conidia from
‘Trifolium pratense infected this host but gave negative results on
seven other species of this genus as well as on species of other genera
tested.
Microsphaera Astragali (DC.) Trev. The only results recorded
for species of this genus are those of Neger (107). He infected A séra-
galus glycyphyllus and A. cicer with conidia from the former. Three
other hosts gave negative results.
Uncinula aceris (DC.) Sacc. and U. salicis (DC.) Winter. Neger
(107) used conidia of the former species from Acer pseudoplatanus to
successfully infect A. pseudoplatanus and A. campestre. Conidia of
the second species from Salix purpurea infected S. purpurea and S.
caprea.
Phyllactinia corylea (Pers.) Karst. Neger (107) reports one test
with conidia from Corylus avellana, these failing to infect the same
host. Voglino (166), using conidia from Corylus, infected Corylus
but not Carpinus, and conidia from Carpinus infected Carpinus but
not Corylus. He further found that ascospores from Carpinus infected
Carpinus but not Fagus, while ascospores from Fagus infected Fagus
but not Carpinus.
Sphaerotheca Humuli (DC.) Burr. Salmon (128) used conidia of
this mildew from Potentilla reptans to infect P. reptans; no infection
occurred on Agrimonia Eupatoria, Alchemilla arvensis, A. vulgaris,
Fragaria (cult. sp.) Poterium officinale nor Spiraea ulmanria.
‘
REED: SPECIALIZATION OF PARASITIC FUNGI 387
Steiner (150) found that the mildew on Alchemilla was confined
to the species of this genus. He also claimed to be able to distinguish
specialized races within this genus of host plants. He found that
conidia from Alchemilla pastoralis and A. flexicaulis were alike in
infecting power except that conidia from the former host would not
infect A. Alpigena and only slightly A. pubescens, while conidia from
A. flexicaulis infected A. Alpigena slightly and A. pubescens not at all.
Conidia from A. impexa would not infect A. Alpina vera nor A. nitida,
while conidia from A. pastoralis at least partially infected these hosts.
Steiner also found that conidia from the Vulgares section of the host
genus would not produce full infection on species of the Alpinae sec-
tion, although conidia from species of the latter section vigorously
infected species of the former. Steiner noted marked differences in
the relation of the species of Alchemilla to the mildew, dividing them
into the following groups: (1) Susceptible species, as A. impexa;
(2) immune species, as A. conjuncta; (3) species susceptible to the
mildew from some hosts, while immune to that from others, as A.
micans.
Sphaerotheca Humuli (DC.) Burr. var. fuliginea (Schlecht) Sal-
mon. Salmon (128) has made a few tests with conidia of this mildew
from Plantago lanceolata and Taraxacum officinale. Conidia from the
former infected the same host but not Plantago major nor Taraxacum
officinale; conidia from Taraxacum officinale infected T. officinale but
not Fragaria (cult. sp.), Plantago media nor P. lanceolata.
Oidium on Euonymus japonicus. Salmon (131) reports the fol-
lowing results with this mildew whose identity was not fully deter-
mined; conidia from Euonymus japonicus infected E. japonicus var.
aureus, var. albomarginatus, var. ovatus aureus, var. microphyllus, var.
President Gunter, E. radicans var. microphyllus and var. Silver Gem,
but not E. nanus, E. americanus var. angustifolius, E. chinensis, E.
europaeus, E. radicans var. carrierei, Celastrus scandens, C. articulatus,
C. orixa nor Prunus laurocerasus var. latifolia.
ADDITIONAL FUNGI
Physiological specialization has also been investigated in a few
other groups of fungi by a number of different workers, but, outside
of the rusts and powdery mildews, no extensive studies have been
made.
Synchytrium taraxaci de B. and Wor. In the Chytridiaceae,
Liidi (95) has tested the infecting capacity of swarmspores of Syn-
chytrium taraxaci from Taraxacum officinale. He tried to infect nine-
teen species of Compositae which do not belong to the subdivision
Cichoraceae, but with negative results in every case. He also used
388 BROOKLYN BOTANIC GARDEN MEMOIRS
twenty-one species which belong to genera of this subgroup, but was
able to infect only four species of the genus Taraxacum: T. officinale,
T. ceratophorum, T. palustre and T. erythrospermum. ‘Three other
species of this genus tested remained free from the fungus. In many
of his experiments, Liidi kept control plants of 7. officinale and these
were readily infected by the swarmspores.
Albugo candida (Pers.) Roussel. Eberhardt (31, 32) has made
inoculation tests with this parasite. His results are as follows:
1. Conidia from Capsella Bursa-pastoris infected C. Bursa-pastoris,
Arabis alpina, Iberis amara and Lepidium sativum.
2. Conidia from Capsella Heegeri infected C. Bursa-pastoris and
Lepidium sativum.
3. Conidia and oospores from Lepidium sativum infected L. sativum
and Capsella Bursa-pastoris.
4. Conidia from Arabis alpina infected A. alpina, A. Halleri, A.
hirsuta, A. turrita, Capsella Bursa-pastoris, Cardamine pratensis,
Iberis amara, Lepidium sativum, Senebiera coronopus, but not
Brassica napus, B. nigra, B. oleracea, Raphanus sativus nor
Sinapis arvensis.
5. Conidia from Brassica Rapa infected B. Rapa, B. nigra, B. oleracea
(var. botrytis, capitata, congyloides), Diplotaxis tenuifolia and
Sinapis arvensis, but not Capsella Bursa-pastoris, Iberis amara
nor Lepidium sativum. On the basis of these experiments, ~
Eberhardt concludes that there are two specialized races of
the parasite:
1. On Arabis-Capsella-Lepidium.
2. On Brassica-Diplotaxis-Sinapis.
Melhus (99) has also tested the infecting capacity of Albugo candida
on the radish (Raphanus sativus). He was able to infect this host,
twenty-two varieties being equally susceptible, also Raphanus caudatus,
Brassica alba (white mustard) and Brassica oleracea (cabbage, fifteen
varieties). In the case of the latter plants infection was less certain
than for the radish. The following plants gave negative results:
Brassica rapa (turnip, ten varieties), B. nigra (black mustard), B.
campestris (rutabaga, three varieties), Capsella Bursa-pastoris (shep-
herd’s purse), Lepidium sativum (garden cress), L. virginicum (wild
pepper grass), Sisymbrium officinale, S. altissimum (hedge mustard),
Iberis umbellata (candytuft), Nasturtium officinale (water cress) and
Chetranthus Cheiri (wall flower).
Peronospora parasitica (Pers.) de Bary. Gaumann (59) has
carried out a few experiments with this fungus and finds a high degree
of host specialization. The fungus that occurs on Capsella cannot
infect other Cruciferae. The same is true of the race on Brassica.
REED: SPECIALIZATION OF PARASITIC FUNGI 389
In some cases the parasite seems to be restricted to a single species,
for the fungus on Sisymbrium officinale does not infect S. sophia. On
the other hand the same race occurs on Brassica oleracea and B. rapa.
Taphrina aurea (Pers.) Fr. Giesenhagen (60), as a result of his
work on the Exoasceae, suggests that Taphrina aurea, which infects
three species of Populus, is becoming specialized into races, each of
which is adapted to a single species of Populus.
Claviceps purpurea (Fr.) Tul. Stager (142) has found five special-
ized races in the ergot of rye, Claviceps purpurea. One race occurs on
rye and also on seventeen other species of grasses; a second race occurs
only on Glyceria fluitans; a third is confined to species of Lolium; a
fourth to Poa annua; while the fifth is found on Brachypodium sil-
vaticum and Milium effusum. Both conidia and ascospores, where
tested, are limited in the same fashion.
Staiger did not find any such specialization in Claviceps micro-
cephala. ‘This ergot is reported on only three grasses.
Plowrightia morbosa (Schw.) Sacc. Gilbert (61) reports that
Plowrightia morbosa (Schw.) Sacc. is specialized on the choke cherry
(Prunus virginiana) and wild plum (Prunus americana). Ascospores,
conidia, and pycnospores from the former host were inoculated into
the wild plum without giving any evidence of infection. On the other
hand, ascospores and conidia under like conditions, readily infected
the choke cherry, giving rise to normal knots. A study of the distri-
bution of the fungus on the two hosts lends confirmatory evidence as
to the specialization, for in one locality the disease may be prevalent
on one host while absent from the other.
Rhytisma acerinum (Pers.) Fr. Miiller (104) has made a study of
this parasite on various maples and concludes that it consists of
several specialized races. One race Platanoides is found on Acer
platanoides infecting only slightly Acer campestris and A. pseudo-
platanus; a second race Campestris occurs on Acer campestris, to a
slight extent on A. platanoides, but does not occur on A. pseudoplata-
nus; a third race, which is given specific rank as Rhytisma pseudopla-
tant, occurs only on Acer pseudoplatanus. Tubeuf (162) inoculated
Acer pseudoplatanus, A. platanoides, A. campestris and A. negundo
with ascospores from the first named host, infection occurring only
on this one maple.
Colletotrichum lindemuthianum (Sacc. and Magn.) Briosi and
Cavara. Barrus (16) has studied the relation of bean varieties to the
common anthracnose, Colletotrichum lindemuthianum. He tested the
susceptibility of one hundred sixty-one varieties to a culture of this
organism and found that, while most of the varieties were susceptible
in varying degrees, a few seemed to be immune. When, however,
390 BROOKLYN BOTANIC GARDEN MEMOIRS
other cultures of the organism were used to inoculate the varieties a
different arrangement of susceptibility became evident. Varieties
quite immune to the one strain were severely attacked by another
strain. All the varieties proved quite susceptible to at least one of
the strains used. This indicates the existence of distinct races of this
parasite with fairly definite host limitations.
Edgerton and Moreland (33) have made similar studies with cul-
tures of this same fungus. Their results also indicate differences in
the infecting capacity of strains of the fungus isolated from different
varieties of beans. Some beans, like the snap beans, appear to be
quite susceptible to a number of different strains. Other varieties,
while quite susceptible to certain strains, are resistant to strains from a
different source. .
Edgerton and Moreland have also studied cultures of Glomerella
gossypii (South.) Edg., the cotton anthracnose fungus. They do not
find evidence for the existence of specialized strains in this fungus,
for all the cultures isolated from different sources were able to infect a
large number of cotton varieties.
GENERAL DISCUSSION—BRIDGING Hosts
The above review of the investigations with reference to host
specialization of parasitic fungi indicates that the phenomenon is of
general occurrence. The work done is particularly extensive in con-
nection with the rusts and the powdery mildews, but sufficient has
been accomplished in other groups to make clear the presence of
specialized races.
It is highly probable that the same phenomenon is of wide occur-
rence among other groups of plant parasites. A large number of
so-called species of the Imperfect Fungi, as Cercospora, Phyllosticta,
and Septoria, may really be only specialized races of a relatively small
number of forms distinct on structural grounds. In several genera of
the Imperfect Fungi, as those mentioned, a very large number of
species have been recorded; in fact many of them are, mainly at present
at least, identified by the host upon which they grow. Cultural ex-
periments may result in grouping many of these together, at the same
time making clear the physiological host relations.
We are not, however, to assume that host specialization is of
universal occurrence. There are several cases on record ‘where the
fungus shows no evidence of the specialization of its hosts. A striking
case of this sort is that of Puccinia subnitens Diet: Arthur (5) has
been able to infect ten hosts, belonging to the families Chenopodiaceae,
Cruciferae, and Capparidaceae, with teleutospores from Distichlis
spicata. Bethel (18), using teleutospores from the same grass, recently
REED: SPECIALIZATION OF PARASITIC FUNGI 391
has succeeded in producing aecidia on twenty-two species, belonging
to fifteen genera, distributed among six different families, as follows:
1. Polygonaceae: Polygonum aviculare, P. erectum and P. ramosissi-
mum.
2. Chenopodiaceae: Salsola pestifer, Chenopodium album, C. glaucum,
C. lanceolatum, C. pagonum, Monolepis nuttalliana and Kochia
scoparia.
Amaranthaceae: Amaranthus retroflecus and A. blitoides.
Nyctaginaceae: Abronia fragrans.
. Cruciferae: Capsella Bursa-pastoris, Lepidium densiflorum, L.
medium, Erysimum asperum, Sophia pinnata, Roripa palustris,
Thaspi arvense and Sisymbrium altissimum.
6. Capparidaceae: Cleome serrulata.
It is also suggested that species of Papaveraceae may be aecidial
hosts for the same rust.
Bock (19) has carried out rather extensive cultural tests with
Puccinia Gentianae (Str.) Link. He reports no evidence for host
specialization, finding that a large number of species of Gentiana were
readily infected with rust from two different species.
Another illustration of the same condition is reported by Camilla
Popta (109) who has been able to infect a number of umbellifers with
the same race of Protomyces macrosporus. The following plants were
infected with the fungus from Aegopodium podograria: Cicuta virosa,
Seseli montanum, Libanotis vulgaris, Palimba chalrau, Bubon gemmif-
erum, Pachypleurum alpinum, Bunium virescens, Ferula thyrsiflora,
Trinia vulgaris and Athamanta cretensis.
If now we compare the degree of specialization routed in the
different parasitic fungi we find the greatest divergences. Ina general
way the specialized races may be grouped on the basis of their host
gl as follows:
. Specialized races restricted to certain species of a genus of
ee. Here belong the specialized races of Erysiphe graminis within
the genera Bromus and Hordeum; of Puccinia dispersa within the
genus Bromus; the specialized races of Phragmidium disciflorum,
Puccinia Centaureae, P. Epilobii-tetragoni, P. Helianthi, P. Hieraci,
P. Pulsatillae, P. Ribis, P. Ribesti-Caricis, P. Caricis-montanae,
Uromyces Poae, U. proeminens and Melampsora Euphorbiae.
2. Specialized races restricted to a particular genus of host plants.
A number of specialized races of this sort are known. Among the
Erysiphaceae we find the specialized races of Erysiphe graminis on
Agropyron, Dactylis, Poa and Secale and of Erysiphe cichoracearum
on Aster and Solidago. Among the rusts we find the races of Puccinia
graminis on Aira, Poa, Calamagrostis and Apera; of P. coronata on
Abe
g0L BROOKLYN BOTANIC GARDEN MEMOIRS
Glyceria, Agropyron and Bromus; most of the races of P. glumarum
and P. dispersa.
3. Specialized races occurring on two or more genera but belonging
to the same family. As examples, we may mention the specialized
races Avenae, Secalis and Tritict of Puccinia graminis: Loli, Calama-
grostis and Phalaridis of P. coronata; Secalis of P. glumarum; Orchide-
arum-phalaridis of P. sessilis; the races of P. Ribesti-Caricis, P.
extensicola, P. silvatica, P. Bistortae, P. mammillata, Albugo candida
and Claviceps purpurea.
4. Races occurring on hosts belonging to different families as
Puccinia subnitens, Uromyces Scirpi and Erysiphe cichoracearum.
When we compare the specialization of a parasite in relation to a
particular host we also find the greatest differences. For example
Puccinia graminis avenae occurs, according to Eriksson (41), on twenty
species of grasses belonging to fourteen genera, Carleton (25) recording
it on nineteen species belonging to fifteen genera, Jaczewski (68) on
seven species belonging to six genera and Stakman and Piemeisel (149)
on thirty-three species belonging to twenty-one genera. All agree
that this rust occurs on a wide range of more or less unrelated hosts.
On the other hand, Puccinia coronata avenae is restricted to species
of Avena and possibly Arrhenatherum. The powdery mildew of oats,
Erysiphe graminis avenae, is also sharply restricted to Avena, infecting
to some extent Arrhenatherum. The crown rust and powdery mildew
are similar in infecting a large number of species and varieties of
Avena.
A similar condition is found in the case of the parasites occurring
on Secale cereale: Claviceps purpurea secalis occurs on eighteen species
belonging to eleven genera; Puccinia graminis secalis, according to
Eriksson (41) on eleven species belonging to five genera and according
to Stakman and Piemeisel (149) on twenty-three species belonging to
nine genera; Puccinia glumarum secalis on Secale cereale and Triticum
vulgare; Erystphe gramints secalis on two species of Secale; and Puc-
cima dispersa secalis on Secale cereale.
Many of the specialized races, while in the main restricted to certain
hosts, yet are able to infect to a greater or less extent a number of
other plants. Generally these races are distinguished by their ability
to fully infect certain hosts while their development on others is
weak and limited. Very good illustrations of such races are found
among several of the rusts. The various races of Coleosporium
Campanulae are not sharply limited to definite hosts, but the hosts of
one race may also be attacked by other races. The same is true of
the races of Puccinia graminis, P. coronata, P. Ribesii-Caricis, Melamp-
sora Larici-epitea and Uromyces Poae.
REED: SPECIALIZATION OF PARASITIC FUNGI 393
It has repeatedly been suggested by many investigators that special-
ized races of parasitic fungi may extend their normal host range by
passing through certain so-called “bridging hosts.’”’ There are many
cases known where a particular host plant can be infected by two or
more races of a parasite. Such hosts may enable a specialized race
to infect a wider range of plants.
Ward (172) apparently was the first to emphasize this possibility
as a result of his studies of the behavior of Puccinia dispersa on various
bromes. Ward supposed that Bromus arduennensis, which is readily
infected with uredospores of Puccinia dispersa from Bromus mollis
of the section Serrafalcus, as well as by uredospores from B. arduen-
nensis of the section Libertia, served as a means for the rust on bromes
_ of the section Serrafalcus to pass over on to bromes of the section
Libertia. The following data indicate Ward’s results as bearing on
this point.
Uredospores Uredospores
from Bromus from Bromus
Section Libertia: arduennensis mollis
LE HE ETO UTA TEINOSO OD OE I OO NU DE 8/72 13/14
ibromus arduennensts var. villosus...2... 05.2 +e sees 10/10 1/14
Section Serrafalcus:
LER IST ALLE ROO OD AOE AAA OE: OT oe OTE 1/8 119/154
ESO LUSUSCCOLLILUUS ENA Pt isd dey ase Go Ao cake a oak Sys a 8/8 31/61
Stenobromus:
HSROPEUSEHIDMITLUS eA tacts iste Se ne sae were. 0/6 1/74
TVAR TIS NEL OSSL-Sa hs CORig CHE o TEE ORR 0/8 4/148
An examination of the above data leads one to conclude that the
rust on the two hosts, Bromus arduennensis and B. mollis, are practi-
cally identical in their capacity for infecting other bromes. The
rust on Bromus arduennensis does not have any wider host range than
the rust on B. mollis. Both grasses appear rather to be hosts for the
same strain of rust.
Ward’s evidence that Bromus Krausei and B. pendulinus may serve
as bridging hosts is perhaps stronger. The following data indicate
the relation of these bromes to the rust on Bromus sterilis and B. mollis.
Uredospores Uredospores
from Bromus from Bromus
Serrafalcus: sterilis mollis
PS OULIESNISS LULSCL EL Aae es Sine Are AR tea ie Crees 14/298 27/27
Mirgnius PENAIINUS,. ):. wi ese ew AG eS. es 17/65 50/50
ESPOITAZES IL OULU ONITIAS. seduce teteme es ye raiclls ome Peake eho esas 1/25 2/26
BIEL PSU DO SADOE SOL ne Se noe L/i37 119/154
PSR ILS UCSULLULS Ore Oe ee ee ee hee 1/4 3/4.
Stenobromus:
IY CATALSPSICLELI Siar tomes Fas NS Wee Aco ey Sbesbautis eet? 126/146 4/148
TESA OTUILS’ OUSSONU eae Pests GN oe ia ae tere to oars 37/60 6/53
1 The denominator of the fraction indicates the number of leaves inoculated
and the numerator the number infected.
18The denominator of the fraction indicates the number of leaves inoculated
and the numerator the number infected.
394 BROOKLYN BOTANIC GARDEN MEMOIRS
From this data it is evident that Bromus Krausei and B. pendulinus
are susceptible to the rust on both B. mollis and B. sterilis. From the
standpoint of bridging hosts, however, what we need to know is the
infecting capacity of uredospores from B. ‘Krauset and B. pendulinus,
produced by inoculation with uredospores from B. mollis and B. sterilis.
It may well be that B. Krausei and B. pendulinus are merely hosts for
the rust on both B. mollis and B. sterilis. At least the evidence is
not complete for proving that they are bridging hosts.
Freeman and Johnson (57) conclude that barley is a bridging host
enabling the specialized races of Puccinia graminis on wheat and rye
to infect oats. They find that the rust on wheat will not infect oats,
but will infect barley; the same is true of the rust on rye. When,
however, the rust on barley, produced by inoculation from either
wheat or rye is sown on oats, infection occurs to a very slight extent.
The data upon which this conclusion is based are as follows: (1) uredo-
spores from wheat to barley (26/31),'4 to barley (28/42), to barley
(16/16), to oats (2/54); (2) uredospores from rye to barley (23/31),
to oats (1/22). As noted before, the barely rust is able to infect all
four cereals, but rye and oats less completely than wheat and barley.
The indications are that wheat rust and rye rust, as a consequence of
growing on barley for one or more generations, are able to infect oats.
It is evident, however, that the data obtained are rather meager and
very much more extensive series of inoculations should be carried out.
Evans (44) has crossed a wheat (Bob’s Rust Proof) resistant to
rust (Puccinia graminis) with another wheat (Wol Koren) which is
highly susceptible. Evans found that the hybrid, although of more
vigorous growth than either parent, was much more severely attacked
by rust than the susceptible parent. It was also noted that in pot
cultures in the greenhouse the hybrid produced an abundance of
teleutospores, which rarely occurred on either parent under the same
conditions.
Evans next tried to determine the infecting capacities of the rust
after it had developed on the hybrid. He found that the rust from
the hybrid infected the susceptible parent much more severely than
the rust originally found on it. Not only that but the rust on the
hybrid readily attacked the resistant parent. Accordingly it is
suggested that hybrid plants may play an important part in the trans-
mission of parasites from susceptible to resistant varieties by increasing
the virulence of the parasite.
Stakman and Piemeisel (149) record many grasses as hosts for
more than one race of Puccinia graminis. In fact Bromus tectorum,
144The denominator of the fraction indicates the number of leaves inoculated
and the numerator the number infected.
REED: SPECIALIZATION OF PARASITIC FUNGI 395
Hordeum vulgare, and Secale cereale are infected by all six races that
they worked with. They insist, however, that these races are all
distinct and that bridging hosts are not present. The grasses which
harbor more than one race of rust are, of course, important in the
spread of these races, even though they do not enable them to increase
their usual host range.
Johnson (69) reports that certain grasses enable the timothy rust
to extend its normal range. He found that this rust would not directly
infect Hordeum vulgare nor Triticum vulgare. However, when the
timothy rust was transferred to Avena sativa, the uredospores produced
on this host infected Hordeum vulgare. Further uredospores pro-
duced on Festuca elatior by inoculation from timothy infected both
Hordeum vulgare and Triticum vulgare. It was also found that uredo-
spores from Dactylis glomerata, produced by inoculation from timothy,
infected Triticum vulgare.
Stakman and Jensen (145), however, find no evidence for bridging
hosts in the timothy rust. Neither Avena sativa nor Dactylis glomerata
increased the host range. They also report that Hordeum vulgare is a
host for the timothy rust. Stakman and Piemeisel have further
extended the host range of this rust and emphasized its relation to
the race Avenae.
Arthur (5) suggests that Helianthus annuus may be a bridging
host for various races of the sunflower rust, Puccinia Heliantht, special-
ized to a narrow range of species of Helianthus. H. annuus seems to
be readily infected by means of teleutospores from other sunflower
hosts. Arthur (3, 4) and Kellerman (74, 75) report successful infec-
tions with teleutospores from H. mollis and H. grosse-serratus; Arthur
(5) further reports successful infection with teleutospores from H.
laetiflorus and Kellerman (75) with teleutospores from H. tuberosus.
The teleutospores from these hosts vary in their ability to infect other
Helianthus species and, according to results reported, are not able to
infect each other except that, according to Arthur, infection of H.
mollis occurred when teleutospores from H. laetiflorus were used.
Neither Arthur nor Kellerman have reported positive tests with the
sunflower rust found in nature on H. annuus or produced on it experi-
mentally by using spores from other species. Jacky (65) in Europe
reports a few tests with teleutospores from H. annuus; these were
able to infect only three out of eight species tested. As yet no one
has clearly shown that the rust on H. annuus has a wider range of
hosts than the rust on H. mollis, H. grosse-serratus, etc. In fact, the
evidence is much stronger that H. annuus is a very susceptible host
to the various races of rust occurring on other species of Helianthus,
if such races really exist, than that H. annuus is a bridging host.
396 BROOKLYN BOTANIC GARDEN MEMOIRS
In various heteroecious rusts it has been suggested that the aecidial
host may act as a bridge for races occurring on the uredo and teleuto
hosts to pass over on to other species normally beyond their range of
infection. In the case of Puccinia graminis the aecidial host, Berberis
vulgaris, is common to all the races specialized on different kinds of
grasses. In the aecidial stage the difference between the races on the
gramineous hosts might disappear and the aecidiospores produced on
the barberry might have a much wider range of infection.
Several investigators have published data bearing on this point.
Eriksson (41) has infected the barberry using teleutospores from more
than fifty different grasses. In some of these cases the aecidiospores
produced were used to inoculate various gramineous hosts. Some of
Eriksson’s results may be indicated in the following summary:
Aecidiospores Sown on
ul | a
N 98/9 See cate Soe siete ae +) 8 S als *
a a a a
AVENAE |
Aveng SaUva..... 00. v2.58 aes 2/215 1/1 0/2\o/2 0/2
Briza maxima............ I/I \o/1/1/1 o/I
Bromus arvensis .......... 1/1 |o/I o/I
Bromus brachystachys..... rey |Keyfas o/1,0/1 o/I
Bromus madritensis....... 1/1 1/1 0/1\1/I 0/1
Dactylis glomerata ........ 1/1
Festuc@ MYUPUS.. 5 + ashe 1/I \0/1\1/1 o/I
Festuca tenuiflora......... 1/1 10/1 0/1 o/I
Koeleria setacea........... 1/1 o/1| \o/I 0/1/0/1
Milium effusum .......... 2B ea| 0/1 o/I o/I
Phalaris canariensis.......| 1/1 | G/II/1 o/I
Phleum asperum.......... yi (0/1/0/1 o/I
Vulpia bromoides......... 1/1 | 0/1 0/1 o/I
SECALIS |
Agropyron repens......... 0/1 1/1 1/1)1/t o/I
BROMUS SECHLINUS «ote. oe. O/ | 5.) 8/x
Elymus sibiricus.......... 1/t
Hordeum vulgare.......... 0/2 2/2\r/t 1/2
SICCOLCNGEV CLIC ina en See 0/2 2/2\2/2 0/2
AIRAE |
AWK DOWUNICD. 4». 5... a2 ee | 1/1
POAE |
PGUSGUCSTO RI a sils Rieke teak | | o/1|1/1
Poa compressa ........... | 0/3: ate | jo/r|x/t 1/1,0/1|0/1
TRITICI beret not
Triticum vulgare.......... 1/3 | |2/3|1/2 6/7
In general, the aecidiospores from the barberry are restricted in
16 The denominator of the fraction indicates the number of tests and the numer-
ator the number that were successful.
REED: SPECIALIZATION OF PARASITIC FUNGI 397
their capacity for infection in the same way as uredospores from the
same gramineous hosts used as a source of the teleutospores for in-
fecting the barberry. The only marked variation from this is in the
case of aecidiospores from the barberry produced by inoculation with
teleutospores from Bromus madritensis, Briza maxima, Festuca myurus,
and Phalaris canariensis, all hosts for the race Avenae, which infected
not only Avena sativa but also Secale cereale, a host of race Secalis of
the rust.
Jaczewski (68) found close correspondence in the infecting capacity
of aecidiospores from the barberry arising from teleutospore inocu-
lations from definite plants and that of the uredospores from the same
gramineous hosts. In fact his establishment of the nine specialized
races of the black stem rust in Russia is based as much on aecidiospore
inoculations as on uredospore inoculations.
Stakman (143) also found no essential differences in the infecting
capacity of uredospores from wheat and Agropyron repens and that of
aecidiospores from the barberry arising as a result of inoculation with
teleutospores from these same hosts. Pritchard (110) also found a
correspondence between the infecting capacity of uredospores and
aecidiospores.
Bolley and Pritchard (20) assert that aecidiospores from a single
barberry hedge have been used to infect wheat, oats, barley, Hordeum
jubatum, Agropyron tenerum and A. repens. The origin of the infec-
tion of the barberry was not known but probably was due to teleuto-
spores from Hordeum jubatum. In 1905 aecidiospores from barberry
readily infected barley and Hordeum jubatum, less readily wheat, and
oats hardly at all.
Arthur (10), however, has come to the conclusion that the “ bar-
berry acts as a bridging host between each and every other gramineous
host.’’ The evidence that he gives in support of his statement may
be indicated: (1) aecidiospores from barberry, produced by inocula-
tion with teleutospores from Agrostis alba, infected wheat and barley,
but not oats; (2) aecidiospores from barberry produced by inocula-
tion with teleutospores from Agropyron tenerum, infected oats; (3)
aecidiospores from the barberry, produced by inoculation with teleuto-
spores from Sitanion longifolium, infected wheat; (4) aecidiospores
from the barberry, produced by inoculation with teleutospores from
Elymus canadensis, failed to infect wheat and rye. No one has
recorded any data on the infecting capacity of uredospores developed
on Sitanion longifolium; accordingly we have no information whether
this rust is a distinct race or whether S. longifolium is merely a host
for specialized race Tritici. Further, Agropyron tenerum is a host for
specialized race Avenae; hence the results recorded are just what one
27
398 BROOKLYN BOTANIC GARDEN MEMOIRS
would expect. Agrostis alba, however, seems to harbor a distinct race
of rust and so the result recorded is not in line.
Taking the results of aecidiospore inoculations as a whole, there
seems to be no good reason for assuming that aecidiospores from the
barberry, produced by teleutospores from a known grass, have any
greater range of hosts than uredospores from the same grass. It
appears that the racial strains of the black stem rust are not so sharply
fixed in their host restrictions in either the uredo or aecidial stage.
Further, the nature of the specialization is different in Europe from
what it is in America. It is not surprising, then, that these races are
able to grow on other hosts. There is, however, no clear indication :
that the barberry acts in any way as a bridging host and that it
enables the races on different grasses to increase their range.
The possibility of the aecidial host serving as a means for extending
the host range of specialized races is quite apparent in Puccinia
coronata Corda. Miihlethaler (102) records Rhamnus Imeretina as an
aecidial host for specialized races of three of the main subgroups of
the crown rust: Puccinia coronata (Corda )Kleb., P. coronifera Kleb.
and P. alpinae coronata Mihlethaler. Rhamnus Purshiana is likewise
an aecidial host for the races on two subgroups—P. coronata (Corda)
Kleb. and P. alpinae coronata Miihlethaler. There is, however, no
evidence at hand to indicate that, as a matter of fact, these species do,
in any way, act as bridging hosts.
There are many other cases where a particular species of plant is
a host for two or more specialized races of a parasite and it might be
possible for these to enable the different races to extend their host
range. A few cases of this sort may be mentioned. According to
Jaczewski (68) Agropyron repens and A. caninum are hosts for the
races Tritict and Secalis of Puccinia graminis and this might serve to
enable one race to pass over on to the hosts of the other. There is,
however, no experimental proof in support of the suggestion. Stak-
man and Piemeisel (149) record a number of hosts as common to several
or all of the six races they studied. However, no bridging occurs,
each race being distinct. According to Miihlethaler (102), Festuca
elatior is a host for races Festucae and Lolit of Puccinia coronata.
Phalaris arundinacea is the only uredo and teleuto host for the special-
ized races of Puccinia sessilis with their aecidial stages on Liliaceae,
Orchidaceae, Amaryllidaceae and Araceae. Several species of Ribes
are common aecidial hosts for the specialized races of Puccinia Ribesti-
Caricis. Similar conditions are found among a large number of other
forms—Uromyces Dactylidis, U. Poae, U. Fabae, U. Scirpi, Coleo-
sporium Campanulae, Melampsora Larici-epitea, M. populina, M.
Tremulae, etc.
REED: SPECIALIZATION OF PARASITIC FUNGI 399
The occurrence of bridging hosts has been suggested in other
groups of parasites as well as in the rusts. Salmon (126, 127), in
connection with his work on the powdery mildew of the bromes, has
suggested the possibility that Bromus hordeaceus may act as a bridging
host for the mildew on Bromus racemosus and B. commutatus. He
found that the mildew on B. racemosus failed to infect B. commutatus
(0/12),'° while it readily infected B. hordeaceus (34/34). Furthermore
conidia from B. commutatus failed to infect B. racemosus (0/36), while
infecting B. hordeaceus (40/49). Salmon supposes that B. hordeaceus
may act as a bridge for the mildew on B. racemosus to pass over to
B. commutatus and also the reverse. In one case, Salmon infected
B. hordeaceus with conidia from B. racemosus. The conidia produced
on the former were then used to infect B. commutatus. Salmon, how-
ever, did not test the infecting capacity of the conidia thus produced
on B. commutatus.
Steiner (150), in his work with the mildew (Sphaerotheca Humult
(DC.) Burrill) of Alchemilla, reports the occurrence of bridging hosts.
He states that conidia from Alchemilla connivens and A. pubescens
readily infected A. pastoralis and A. impexa but gave negative results
when sown on A. micans. On the other hand, A. micans is readily
infected with conidia from A. pastoralis and A. impexa. Accordingly
Steiner supposes that A. pastoralis and A. impexa may carry the mil-
dew over to A. micans from A. connivens and A. pubescens. Some
evidence is also given to indicate that A. impexa is a bridging host
between A. nitida and A. fallax. Steiner’s conclusions, however, are
based on only a few tests.
In the Erysiphaceae the question has been raised whether the
ascospores and conidia from a particular plant possess the same in-
fecting capacities. Marchal (98), Salmon (124, 132), and Voglino
(166) have, in a few cases, used ascospores for inoculation tests and
they report that the capacity of ascospores for infection is identical
with that of conidia from the same host.
It is, however, somewhat surprising that the evidence that various
races of parasites may increase their range by means of bridging hosts
is so very meager, if such really occurs. In those cases where the
suggestion of bridging has been most emphasized one is not impressed
with the data supplied. In fact, in all such cases the races of the
parasite are not sharply limited in their host range. They may infect
some hosts more readily and more vigorously than others, but the
virulence of the parasite does not seem to be increased or decreased
by developing on congenial or uncongenial hosts.
16The denominator of the fraction indicates the number of leaves inoculated
and the numerator the number infected.
400 BROOKLYN BOTANIC GARDEN MEMOIRS
The notion of bridging hosts, of course, implies that the fungus
undergoes a physiological change in consequence of its new habitat
and thus becomes able to attack other hosts. The change is certainly
closely associated with variation in virulence so well known in the
case of pathogenic bacteria. It is quite likely that fungous parasites
vary in virulence or can be made to do so by suitable experimental
methods. At present, however, we have no good evidence that this
has been done in any particular case. The results of Evans (44),
referred to above, point in this direction. In this case, however, the
data are not at all extensive. The facts might be explained by an
increase in susceptibility of both resistant and susceptible parent in
response to a change in the environment; or external factors may
have favored a more successful invasion on the part of the rust parasite.
Passing reference may be made to the work of Salmon (129, 130),
Ward (173), Stakman (143) and Spinks (141) which clearly indicates
that a plant may be rendered more or less susceptible to fungous
invasion by means of certain agencies. The work of these investi-
gators shows that mineral starvation, excess of nutrients, mechanical
injuries, anaesthetics, etc., modify the relations of a plant to fungous
invasion.
It has been pointed out by Eriksson (35), Ward (172, 174), Vavilov
(164, 165) and others that a specialized race tends to occur on more or
less closely related hosts. There are, however, great differences among
the specialized races in this respect. As pointed out above, the host
range of these races may be narrow or wide. Within a single species
of parasite we may have a race occurring on many hosts belonging to
different genera and another race restricted to a single genus or even
certain species of a genus. Puccinia graminis, as well as other fungi,
includes races of such wide differences in host range.
Attempts have been made to utilize the infective capacity of a
parasite to determine the genetic relationship of hosts. Eriksson (35)
applied this test in determining the possible relation of a rye-wheat
hybrid to the two parents. Ward (171, 172) reports a fairly close
correspondence between the hosts of the more or less well-defined
races of Puccinia dispersa bromi and the grouping of the bromes on
other grounds. Vavilov (154, 155) has used Puccinia dispersa tritici
and Erysiphe graminis tritici as a test in determining the relationship
of types and varieties of Triticum. He also used Puccinia graminis
avenae and P. coronata avenae as a similar test in connection with species
and varieties of Avena. It is interesting to note that the parasites on
wheat gave practically the same results and these are both quite
narrowly specialized races. The rusts on oats, however, did not give
corresponding results, Puccinia graminis avenae infecting a wider
REED: SPECIALIZATION OF PARASITIC FUNGI 401
range of varieties than P. coronata avenae. The former, as noted above,
occurs on a number of species of grasses belonging to different genera,
whereas the latter is closely restricted to the genera Avena and Ar-
rhenatherum.
It has been suggested that there is a connection between stability
of host species and the occurrence of specialized races. Edgerton and
Moreland (33) suppose that the explanation of the difference between
Bean Anthracnose and Cotton Anthracnose is due to the fact that
bean varieties are quite distinct and well marked, with little or no
crossing and, accordingly, no intergrading forms. On the other hand,
cotton varieties readily cross and thus a series of intergrading forms
occurs onto which the anthracnose fungus may spread. This explana-
tion, however, cannot have any very general explanation. The evi-
dence indicates that the willows hybridize quite readily and yet a
number of specialized races of willow rusts are recorded.
Magnus (96) suggested that the existence of these specialized races
may indicate an adaptation on the part of the parasite to live on
particular hosts. He makes a distinction between adaptive races and
biologic forms. The former term is applied to strains or races of a
parasite which tend to infect certain hosts more readily than others.
On the other hand, the biologic form is sharply restricted to its hosts.
Dietel (30) points out that a parasite may first have attacked a wide
range of hosts, gradually becoming broken up into races adapted to
certain hosts and finally limited to them. It is possible that the rela-
tive abundance of the hosts may have been a factor in this process.
The presence or absence of hosts in a given locality may also have
played a part.
The question has often been raised as to whether these specialized
races of fungi are constant or fixed. Montemartini (100) has recently
raised the question again as a result of his studies with certain para-
sites. He inclines to believe that these races are not fixed and definite
and so capable of being carried from one region to another but rather
that they are local adaptive forms, perhaps dependent on the vari-
ability or distribution of the host plants. He suggests that they are
not permanent but temporary, owing their origin to the various nutri-
tive conditions afforded by different hosts.
I have already called attention to the fact that some specialized
races are distinguished by their ability to infect some hosts more fully
than others. The races of Puccinia Ribesii-Caricis, P. Hieracii, P.
graminis, Uromyces Dactylidis and Coleosporium Campanulae afford
illustrations of this condition. The infection or non-infection of
certain hosts is dependent to a great extent upon particularly favorable
experimental conditions. It is well known that various factors do
402 BROOKLYN BOTANIC GARDEN MEMOIRS
influence in a striking way the capacity of a parasite for infecting
hosts.
The extent of the stability of specialized races must vary greatly
in different cases. Some of the forms above considered are doubtless
as fixed and constant as many parasites which show structural differ-
ences. Mention may be made of the races of Melampsora Tremulae
as illustrating a good case of physiological species. In studying the
races of Erysiphe graminis one also gets a strong impression of their
constancy and definiteness and they seem as real as though separable
by structural features.
In other cases, however, this is not true. One is not impressed
with the definiteness of races in Puccinia Helianthi, P. Hieracii and
others. These are not characterized by well-defined host limitations.
Perhaps one is justified in distinguishing different types of specializa-
tion as physiological species, races, strains, etc.
But little data are available for comparing the specialization of
the same fungus in widely separated localities. The specialization of
Puccinia graminis is apparently the same in Sweden and Russia but
it has taken a quite different course in North America. Treboux
(159, 160) finds a different condition in Puccinia coronata in southern
Russia from what Eriksson (37, 42) finds in Sweden and Miihlethaler
(102) finds in Switzerland. Carleton’s (26) results in the United
States also diverge widely from those of Eriksson. Arthur (11, 13)
does not find evidence for specialization in Puccinia Ribesi-Caricts
in this country, while Klebahn (87, 91) reports several fairly well-
defined races of this rust in Europe. On the other hand, there seems
to be no essential difference in the specialization of Erysiphe graminis
in Europe and North America. The same sharp host limitation seems
to occur in both countries. Further data are necessary before we are
able to determine the relation between the specialization of parasites
in different regions.
In a few cases the races are also characterized by minor structural
differences. Freeman and Johnson (57) and later Stakman and
Piemeisel (149) have noted variations in the size and shape of the
uredospores which distinguish the races of Puccinia, graminis. Kle-
bahn (87, 91) has noted similar differences in the spores of the various
races of Puccinia Absinthi, Melampsora Larici-epitea and others.
Fischer (46, 47), as well as others, has suggested that physiological
specialization is a starting point for the origination of forms distinct
on structural grounds. The suggestion is plausible, for it is possible
to arrange a series of forms ranging from races differing in host rela-
tions through all stages to others showing constant structural differ-
ences.
REED: SPECIALIZATION OF PARASITIC FUNGI 403
The general occurrence of specialized races of parasitic fungi makes
their study particularly important. It is especially desirable td
know the exact host relations of the different races as well as to deter-
mine whether the races are stable and constant or whether they are
capable of a change of virulence and consequently able to extend their
host range.
20.
21.
22.
ae
24.
4 25.
26.
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Beitrage zur Kenntnis der Coleosporien und der Blasenroste den Kiefern
(Pinus silvestris L. und P. montana Mill.). Zeitschr. f. Pflanzenkr. 6: 9-13.
1896.
Beitrage zur Kenntniss der Puccinia silvatica Schroeter und der Puccinia
sessilis Schneider. Ber. d. d. Bot. Gesell. 14: 212-215. 1896.
Beitrage zur Kenntniss der Coleosporien und der Blasenroste der
Kiefern (Pinus silvestris L. and Pinus montana Mill.) III. Zeitschr. f.
Pflanzenkr. 8: 257-262. 1898.
. Ward, H. M. On the Relations between Host and Parasite in the Bromes
and their Brown Rust, Puccinia dispersa Erikss. Ann. Bot. 16: 233-315.
1902.
—— Further Observations on the Brown Rust of the Bromes, Puccinia
dispersa (Erikss.) and its Adaptive Parasitism. Annales Mycol. 1: 132-151.
1903.
Experiments on the Effect of Mineral Starvation on the Parasitism of
the Uredine Fungus Puccinia dispersa on Species of Bromus. Proc. Roy.
Soc. (London) 71: 138-151. 1902.
—— Recent Researches on the Parasitism of Fungi. Ann. Bot. 19: I-54.
1905.
RELATION OF MARL PONDS AND PEAT BOGS
W. W. ROWLEE
Cornell University
The filled-in lakes and ponds of western New York are of two
distinct types, the bogs often called peat or cranberry bogs, and the
marl ponds. These ponds are alike in that both occupy depressions in
the terrain and both are filled with water from springs at their bottom
or near their shores. They also resemble each other in that both are
subject to filling in with material produced by organic life in and
around them. ‘They differ from each other however in the character
of the water they contain, in the flora which inhabits the water and
the adjacent shore and the method by which they are filled in.
The glaciation of the country left a terrain with potholes and other
depressions particularly favorable to peat-bog formation.
The material with which peat bogs are filled consists mainly of
sphagnum and heath-like plants always much disintegrated and
accumulated principally around the shores. Peat bogs have long
been a subject of interest. Mitchill in 1798 studied them and set
forth their general structural characteristics in a paragraph of his
report as commissioner for the Agricultural Society of New York,
as follows: ‘‘As the peat is formed, layer over layer, in the course of
successive vegetations, it can be easily explained how trunks of trees,
fossil wood, and bodies and bones of animals came to be buried so
deep below the present surface; because at the same time when the
trees fell, and animals died, in the places where they are now found,
they were upon the top, and, by the perpetual growth of the plants
around, they have in many places, become covered to a great depth.”
He was particularly impressed with the bones of extinct species of
animals found in bogs in Orange County and other parts of eastern
New York. Dachnowski in 1912 gives a very comprehensive dis-
cussion and classification of the distribution of species in the several
areas on the surface of a bog.
The flora now found on the peat bogs corresponds to the flora of
colder climates. The flora of the marl ponds corresponds more closely
to that of the seashore in the same or more southerly latitudes. Marl
ponds are filled in not only near the shores but in all parts of the pond
where the water is not too deep. The water in the marl ponds is
410
ROWLEE: RELATION OF MARL PONDS AND PEAT BOGS 4ll
decidedly hard, that is, it is impregnated with lime while the water of
peat bogs is soft, that is, it is not alkaline in its reaction.
In a series of careful studies Davis has shown that marl is com-
posed mainly of the remains of the alga Chara. Chara thrives in
hard water and its cell walls are impregnated with calcium carbonate.
In many marl ponds a complete transition from the living Chara to
characteristic marl can be seen. Since Chara grows submerged and
the principal bog plants grow emerged it is evident why there is filling
at the shore in the one and in all parts of the pond in the other.
In sounding many peat bogs in western New York, the writer was
somewhat surprised to find many of them underlayed with marl.
Fic. 1. Marl bog, northeast side of Lowry’s Pond, West Junius, Seneca Co.,
N. Y. Species of sedges the dominant vegetation form.
The assumption had been that the alkaline or nonalkaline character
of the water originally filling the depressions determined whether bog
vegetation or marl pond vegetation would develop in it. Is it possible
that a pond might be alkaline during one stage of its existence and
then become non-alkaline in a later stage? An affirmative conclusion
seems inevitable.
It is Dachnowski’s view that there are changes in the vertical or
historical succession in the bogs, for he says: ‘‘While working on the
eeology of ravines near Ann Arbor, Michigan, I became convinced
that the reactions of plants on their habitat were equally as great and
profound, in some cases, as the influence of edaphic and climatic
412 BROOKLYN BOTANIC GARDEN MEMOIRS
factors. In various places the decomposed remains of an earlier
vegetation led to mechanical and chemical changes in the soil, the
extent of which was more effective toward breaking up the flora into a
heterogeneous formation, accompanied by a frequent replacement of
one dominating group by another.”
There are some bogs without evidence of marl at the bottom.
Davis reports all the many peat bogs examined by him in Maine as
resting on sand, clay or rock bottom, none on marl. Most of the
peat bogs in the Adirondack region of New York have no marl at the
bottom. These presumably were from the first supplied by springs
of non-alkaline water. Some of the peat bogs of central and western
Fic. 2. Peat bog near McLean, Tompkins Co., N. Y. Chamaedaphne, Andro-
meda, Ledum and other heaths together with Sphagnum form the dominant vegeta-
tion.
New York have great masses of marl under them. Such a one is near
Peterboro in Madison County and another on Gorham Creek in On-
tario County. Here it is apparent that some agency changed the
composition of the water to such an extent that oxylophytes found
conditions congenial.
The most extensive marl ponds in the region are in the vicinity
of the limestone belt of central New York. They occur at West
Junius in Seneca County and southwest of Rochester, especially at
Bergen in Genessee County. Isolated and much smaller marl ponds
ROWLEE: RELATION OF MARL PONDS AND PEAT BOGS 413
occur at Tully and near Cortland. The limestone outcrop in these
regions accounts for a continuous supply of water impregnated with
lime. The peat bogs with the greatest thickness of marl under them
occur not far from this same limestone belt, while the peat bogs with
little or no marl are usually farthest from the limestone outcrop.
It does not seem at first thought as though Chara could be the
agency causing the radical change in the history of the vegetation of
the pond and the accumulations in these depressions. In so far as
vegetation is concerned calcium carbonate is relatively insoluble.
Fic. 3. Transition bog near Cortland, N. Y. A thick bed of marl is over-
laid with about four feet of fibrous peat. Marl was excavated from the hole in the
foreground. Sedges are prominent in the vegetation now covering the surface of
the bog.
Not so, however, is the lime in spring water. The water with available
(more or less) free lime is what Chara takes in and in its life processes
converts into calcium carbonates secreted in its walls. That an im-
mense amount of lime is converted is shown by the bulk of marl in the
ponds. Where the amount of the lime in the original soil was not
large, rain water constantly tended to wash it out and in the course of
time the lime content of the water would be decreased. The ponds
were artesian pools fed by these springs and as the character of the
water changed there was, if our theory is correct, a corresponding
28
414 ~ BROOKLYN BOTANIC GARDEN MEMOIRS
change in the flora. There would be then two phases in the life of
our bogs.
rst. The marl ponds in which lime-loving plants predominated
and especially Chara and which are filled with and often surrounded
by beds of marl or “‘bog lime.”
2nd. Peat bogs which from the bottom up are composed of non-
alkine peat which in all their history have been inhabited by oxylo-
phytes.
Between these two phases gradations of all degrees occur in the
peat bogs of western New York. The succession might, of course,
be in the other direction if the calciferous phase should become pre-
dominant over a previous carboniferous one. But such a case is
unknown in western New York. ;
PATHOLOGICAL PROBLEMS IN THE DISTRIBUTION
OF PERISHABLE PLANT PRODUCTS!
@.. E SHEAR
Bureau of Plant Industry, U. S. Department of Agriculture
INTRODUCTION
Under our present social and economic conditions, public interest
is being aroused and directed to questions and problems which have
been largely overlooked or neglected in less strenuous times. The
conservation of our natural resources, especially our food products,
greatly to our discredit as a nation, has, until very recently, been too
largely neglected. Max O’Rell is said to have made the statement
some years ago that Europe could be fed on what America wastes.
This statement is probably somewhat exaggerated, but unfortunately
has too many facts to support it.
At present, these questions, so far as food products are concerned,
are of vital importance, not merely figuratively but literally. We
wish to call attention here to loss and waste occurring in connection
with the distribution of fruits and vegetables.
A large proportion of the fruit and truck crops grown never reach
the consumer. Part of this loss occurs on the farm and in the orchard,
and part in transit and distribution. Adams? in his recent work on
marketing perishable farm products, asserts that at least 25 percent
of the perishables which arrive at the wholesale markets is hauled to
the dump pile because it is unfit for human consumption. This
statement we fear is not based upon sufficient data to be accepted.
It is, however, the opinion of one writer on the subject.
In 1910, according to Dr. Pennington,* the New York Board of
Health condemned and destroyed over twelve million pounds of fruit
and over seven million pounds of vegetables. This presumably does
not represent the total loss, as a considerable amount probably escaped
detection. It is easy to see that such great destruction of food
! Published by permission of the Secretary of Agriculture.
* Adams, Arthur B. Marketing perishable farm products. Studies in history,
economics, and public law, Columbia University. 72%: 25. New York. 1916.
8 Pennington, Mary E. Proper handling of foodstuffs from farm to market.
In Report of the Mayor’s Market Commission of New York City, p. 257, New York,
1913.
415
416 BROOKLYN BOTANIC GARDEN MEMOIRS
products must be a serious drain on our food supply and must add
materially to the cost of living. No adequate estimate can, however,
be made of the enormous economic loss represented in such cases.
Think of the time, labor, and investment involved in the planting,
cultivating, harvesting, and hauling of these products, and of the
freight or express, refrigeration and delivery charges paid! This
is one of the most expensive forms of economic loss imaginable.
CAUSES OF LOSSES
The important problems which confront investigators, producers,
carriers, and consumers, are the causes and means of prevention of
these enormous losses. The producers and transportation companies
have heretofore been too much inclined to look upon their share of
these losses as among the natural hazards of their business. The
carriers, however, are now realizing more fully than ever before,
the great reduction in their income due to the payment of claims from
shippers for loss resulting from decay and spoilage of products in
transit. According to the report of the American Association of
Refrigeration,* the total amount of claims paid by 180 railroads in
1914 for loss of perishable freight was $4,977,383.09; of this amount
over one half, or $2,687,393.36 was for fruit and vegetables. This, of
course, does not represent all the railroads of the country nor all the
losses on the roads represented.
In order to devise means of reducing or preventing this enormous
destruction of food products, it is, of course, first necessary to deter-
mine the causes and their relations and importance. The deteriora-
tion of fruits and vegetables in transit is due chiefly to the action of
parasitic or saprophytic fungi. Natural ripening processes and changes
in the cell contents caused by the accumulation of respiration products
or smothering, may also render the articles unfit for food. These
changes are usually hastened by high temperature and lack of ventila-
tion. Each kind of fruit has, of course, its own natural keeping qualities.
Some kinds, like strawberries, raspberries, blackberries, and figs, soon
become spoiled under optimum conditions, while others, like apples,
may be kept in good condition for relatively long periods. The
structure and composition of the ordinary perishable plant products
and their relations to the keeping and carrying qualities of such
products are fairly well known and no discussion of them will be
attempted here.
There are many other factors, however, involved in determining
the keeping and carrying qualities of fruits and vegetables, such as
* Bulletin No. 2. Issued by Commission on Railway and Steamship Refrigera-
tion of the American Association of Refrigeration, p. 82, June, 1916.
SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 417
soil and climatic conditions under which they are grown, methods of
cultivation and fertilization, nature of the variety, condition as to
maturity at time of harvesting, methods and care in harvesting,
grading, packing, and handling previous to shipment, and methods of
loading, stowing, and bracing in the cars. Any or all these factors
may be and frequently are involved in the final decay due to parasitic
or saprophytic fungi occurring in the field or in transit. Hence it
is of the utmost importance to obtain as complete knowledge as
possible of the various organisms which attack the particular product,
their life histories, the time, mode, and conditions of infection and
development and also their relations to methods of handling and their
temperature, moisture, and host relations. These problems are
primarily pathological.
Growers and shippers long ago discovered that storing fruits and
vegetables at low temperature prolongs their keeping. This observa-
tion finally led to the development of commercial cold storage and
refrigeration methods and practices. These methods and practices
have developed thus far largely along empirical lines. It happens
that growth in most of the organisms which destroy perishable plant
products is inhibited at from 33° to 36° F. Therefore, if fruit or
vegetables, though infected with fungi, are placed under such tempera-
ture conditions before development of these organisms is too far
advanced, growth of the fungi will be temporarily suspended. In
some cases, therefore, refrigeration may simply delay the destruction
of the product and shift or render uncertain the responsibility for its
loss which may occur before it reaches the consumer.
It will appear evident, therefore, that in order to devise methods
of preventing or avoiding such losses, all the factors involved in any
particular case must be accurately determined as well as their relations
and relative importance. Because it is known that certain fungi
destroy certain fruits and vegetables and that these fungi occur in
the orchard or on the farm, it has been inferred by some that the
presence of such organisms on decayed products at destination is
sufficient evidence that the responsibility for the loss rests with the
grower. This may be true in the case of some particular product
affected with some particular disease when shipped without refrigera-
tion. In the case of refrigerated products, however, our experience
and that of others has shown that in order to determine the real cause
or causes and the responsibility for loss in any specific case, the whole
history of picking, packing, handling and treatment of the product
must be known, or at least its history from the field to destination.
This has been very strikingly brought out in the investigations of
418 BROOKLYN BOTANIC GARDEN MEMOIRS
citrus fruits and also in recent investigations of raspberry, strawberry,
and cranberry losses.
The practice of Pap eratien of fruits and vegetables in transit
is for two purposes, viz.: to retard the natural ripening processes which
continue after the crop is harvested and to prevent the development
of destructive fungi which are assumed to be present and are likely
to develop unless a constant low temperature is maintained. In
many cases it is practically impossible to eliminate the organisms
which cause decay and all the handling in such cases must be with the
presumption of their presence and the possibility of their rapid develop-
ment under favorable conditions. .
Each product and each fungus has its own peculiarities and reac-
tions under various conditions and treatment. This may be illustrated
by citing a few specific cases.
STEM-END RoT AND ANTHRACNOSE OF WATERMELON
Meier® has given an account of a decay of watermelons in the
field and in transit, caused by a species of Diplodia. It has been
found that this organism is a wound parasite, and infection takes
place through the stem end of a melon after it has been cut from the
vine. A practical method of preventing this infection by the appli-
cation of a fungicide before shipment has been found to be the simplest
means of preventing decay from this cause. The anthracnose of
watermelons can also be largely controlled by proper field treatment.
LEAK OF POTATOES
This trouble, which is most prevalent on the Pacific coast, has
-.been found by Hawkins® to be due chiefly to Pythium debaryanum.
It has been shown that infection occurs in the field and through wounds
only; hence, the most practical means of prevention is to avoid as
much as possible injury in digging and handling and to sort out all
wounded potatoes before shipping. This is a case in which trans-
portation methods and facilities are not the controlling factor in deter-
mining the condition of the product upon its arrival in the market, but
are of minor importance.
Potatoes, watermelons, and similar products which are not usually
shipped under refrigeration can fortunately be more or less satis-
factorily insured against loss by proper treatment previous to ship-
ment. The temperature, ventilation and handling of such products
> Meier, F. C. Watermelon stem-end rot. Journal of Agricultural Research
6: 149-152. Ap. 24, 1916.
®° Hawkins, Lon A. The disease of potatoes known as “Leak. " On Jour. Agri.
Res. 6: 627-640. 1 fig. pl. XV. 1916.
SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 419
en route, however, have considerable influence upon their condition
at destination, even when all practical field treatments and precautions
have been taken.
CITRUS FRUITS
In the case of citrus fruits, it has been found that where the chief
cause of decay is Penicillium, one of the important factors in its
control is to avoid, as far as possible, all injury to the fruit in picking
and packing, as the fungus enters only through wounds. It is also
necessary to ship this fruit under proper refrigeration in order to
insure its arrival in good condition in distant markets. While proper
care in picking, packing, and handling are of primary importance in
determining the keeping qualities of these fruits, proper refrigeration
and prompt delivery are also essential to prevent loss from this and
other organisms.
’ CRANBERRY ROTS
Cranberries under proper conditions of cultivation and handling
possess excellent shipping and keeping qualities and as they are mostly
distributed during cool weather do not require refrigeration. Spray-
ing to prevent fungous diseases in the field, careful picking and hand-
ling with temporary storage in cool ventilated houses and packing
in proper packages will ordinarily insure their reaching market in
good condition with the usual means of transportation. Most of
the losses occur before shipment and much loss of fruit held for late
shipment is due to the natural ripening processes of the fruit, the
action of the respiration products and smothering. Proper venti-
lation would prevent the latter.
RASPBERRY Rots
Ramsey’ reports the results of studies of shipments of raspberries
from the Pacific coast under various conditions. The decay was
attributed to Botrytis and Penicillium. He found that care in handling
and prompt cooling were among the most important factors in suc-
cessful shipment, but that maintaining a uniform low temperature in
transit was also essential.
STRAWBERRY LEAK
In the cases of strawberries which have been investigated by
Stevens and Wilcox,® of the Bureau of Plant Industry, for the past
7Ramsey, H. J. Factors governing the successful shipment of red raspberries
from the Puyallup Valley. U.S. D. A. Bul. 274: 1915.
8 Stevens, N. E., and Wilcox, R. RB. Rhizopus rot of strawberries in transit.
U.S. DY A. Bull. 531: 4-7. 1917.
420 BROOKLYN BOTANIC GARDEN MEMOIRS
two years, it has been found that “leak,” a decay caused by Rhizopus,
which is the most rapid-growing destructive organism attacking this
fruit, can be controlled by proper methods of picking, handling, and
shipping. Rhizopus spores seem to be practically omnipresent and
it is impossible to eliminate them. All handling of strawberries must,
therefore, be based upon the assumption of their presence. The
fungus, however, cannot gain entrance through the uninjured tissues
and does not develop seriously at a temperature below 45-50° F.
Southern-grown strawberries having fair natural shipping qualities,
carefully picked and handled, and not subjected to too high tempera-
tures before shipment, can with proper refrigeration and transportation
be delivered in northern markets in good condition. If a carload of
strawberries shows much “‘leak”’ at destination, it may be due to
delay or rough handling of cars in transit, and faulty refrigeration;
or it may be due to improper treatment by the grower or shipper.
Only a full knowledge of all the facts in any particular case can deter-
mine the exact cause or causes and responsibility. The presence of
the fungus on the fruit at destination is not sufficient to throw the
blame on the grower or shipper.
Brown Rot oF PEACHES
The brown-rot fungus, Sclerotinia cinerea, which is one of the most
serious causes of decay of peaches, is very common and widely distrib-
uted and is found in practically all peach orchards in humid regions.
It is doubtful whether a shipment of peaches grown in such a region
could be found which did not contain spores of this organism. Not-
withstanding the general presence of this fungus on peaches, if the
fruit is picked at the proper stage of development, and properly
handled, packed and refrigerated in transit, such fruit may, and
usually does, reach distant markets and the consumer in good condi-
tion. The fact that a carload of peaches arrives at destination in a
decayed condition and the brown-rot fungus is present, does not
necessarily indicate that the grower is to blame for the loss.
Mr. J. A. Ruddick,? Canadian Dairy and Cold Storage Commis-
sioner, states that Canadian peaches from the Niagara district
are successfully shipped from Canada to Liverpool and London, the
time in transit to London being twelve days, and also that in 1910
twenty-three thousand cases were shipped from Cape Town, South
Africa, to London, arriving in good condition. Seventeen days was
the minimum time in transit. Other shipments from the same place
® Ruddick, J. A. Cold storage for apples and other fruit. Evidence of Mr. J.
A. Ruddick before the Select Standing Committee on Agriculture and Colonization,
IQIO-II, pp. 106-109. Ottawa, IQII.
SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 421
were made to Canada by way of New York, also arriving in good con-
dition and selling at one shilling each. This indicates some of the
possibilities of shipping such a perishable fruit as the peach when
properly handled and treated, even though ‘inherently liable to
deterioration and decay.”’
Stevens,!° citing the fact that peaches become infected with the
brown-rot fungus only in the orchard or before shipment, says: ‘‘We
may be sure that if infected at destination, shipment was also infected
at the starting point. It appears clear to the writer that in both these
cases (Sclerotinia libertiana on lettuce being the other case mentioned)
the responsibility rests with the shipper just as much as it would if a
consignment of horses infected with glanders but not yet showing the
disease was placed upon the cars.”’
We have been unable to discover facts or data to support the
statement that in such fruit responsibility for losses in transit rests
entirely with the shipper. Investigations have shown that the brown-
rot fungus makes little or no growth at a temperature of 32-35° F.
(o-2° C.),!! and that if the fruit is kept at this temperature brown
rot does not develop. Of course, all practical field treatment to reduce
infection should be practiced.
As a result of the above quoted and similar statements some
railroad representatives have taken the position that the presence
in a shipment of spoiled fruit or vegetables at destination of destructive
fungi known to originate in the field, is sufficient to justify the con-
clusion that the carrier is free from any responsibility for the loss.
Such a general conclusion as this is not in accord with the facts and
is fraught with great possibilities of injustice. Every effort should
be made to correct this mistake. In the past the transportation
companies have undoubtedly paid many unjust claims. Now there
seems to be danger of the pendulum swinging to the other extreme,
resulting in the rejection of just claims.
It is possible for transportation companies to prevent any just
claims for losses due to destructive fungi by furnishing proper cars
and refrigeration service and delivering the products on schedule
time. With the recent improvement of refrigerator cars a sufficiently
uniform low temperature throughout the load can be maintained to
avoid the trouble which so frequently occurs of having variations
in temperature, of 20 degrees or more, between the top and bottom
of the car, as reported by Ramsey.” Under such conditions decay
10 Stevens, F. L. Some problems of plant pathology in reference to transporta-
tion. Phytopathology 5: 108. Ap. I9I5.
11 Brooks, Chas. and Cooley, J. S. Temperature relations of apple-rot fungi.
Jour. Agr. Res. 8: p. 163, Jan. 1917.
2 Ramsey, H. J., 1. c.
422 BROOKLYN BOTANIC GARDEN MEMOIRS
and loss in some products are sure to occur, no matter how great
care has been given them before shipment.
The grower and the shipper, however, are subject to many hazards
and conditions, some of which are beyond their control. Chief
among these are climatic factors. There are many things, however,
which can be done by the producer to prevent or reduce losses of this
kind. All practical means of prevention should be utilized and as
soon as the cause of the trouble in any specific case is determined,
every reasonable and practicable effort should be made by the pro-
ducer or shipper to remove such cause. On the other hand, the carrier
should modify and improve his equipment and methods when neces-
sary to insure the delivery of perishable products in a sound condition.
The various cases described above show something of the variety
and complexity of the problems involved and the need of thorough
investigations to discover the causes and remedies in each case.
The numerous factors involved in the case of any two products or
diseases are frequently not the same and when they happen to be the
same are not of equal importance. Most of them are primarily
pathological or have very direct pathological bearings. Where fungi
are concerned, as in most cases, full knowledge must be obtained of
their host relations, time, mode and conditions of infection, tempera-
ture and moisture relations and the effect on their development of
various methods of treatment of the fruit or vegetable during its
production, harvesting, packing, handling and transportation.
The present agencies interested in and at present studying these
problems are the pathologists, pomologists, horticulturists, refrigera-
tion engineers, the specialists in markets and marketing, the railway
freight claims associations, and the commercial inspection services.
The most complete cooperation of all these agencies is necessary in
order to solve these important problems most quickly and thus reduce
as far as possible this great economic loss.
EXPLANATION OF PLATES IX-XI
PLateE IX. Two flasks of Missionary strawberries kept two days at ordinary
room temperature. a. Containing fruit in natural condition free from wounds.
b. Containing same quantity of fruit inoculated with Rhizopus.
PLATE X. Three wounded strawberries above; three sound berries below.
All were sown with spores of Rhizopus. Photographed after two days at ordinary
room temperature. The three wounded berries entirely destroyed, the three others
sound.
PLATE XI. Pile of spoiled cranberries discarded in sorting and screening. Loss
chiefly due to fungous disease and to the effect of ripening processes and their pro-
ducts or smothering.
BROOKLYN BOTANIC GARDEN Memoirs. VoLUME I, PLATE IX
SHEAR: STRAWBERRY LEAK
BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME I, PLATE X.
SHEAR: STRAWBERRY LEAK
VoLUME I, PLATE XI.
BROOKLYN BOTANIC GARDEN MEmoIRS.
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TUBERS WITHIN TUBERS OF SOLANUM TUBEROSUM
F. C. STEWART
New York State Agricultural Experiment Station, Geneva
At the New York Agricultural Experiment Station, in 1915,
several bushels of seed potatoes not needed for the spring planting
were left over summer in a cellar. The potatoes were of the variety
Sir Walter Raleigh. They were stored in slatted crates which were
piled one above another three crates deep in a single row along the
cellar wall. The cellar was cool, moderately damp and dimly lighted.
Its floor and walls were of cement.
Fic. 1. A new tuber protruding from a slit in the side of an old seed tuber of
Solanum tuberosum. Nat. size.
No attention was given the potatoes until the latter part of Sep-
tember. It was then observed that instead of producing sprouts in
the usual manner they had formed large numbers of new tubers.
Some of the new tubers were in sessile clusters of several small tubers
423
424 BROOKLYN BOTANIC GARDEN MEMOIRS
each while others were borne singly on sprouts one to three centi-
meters long and were of considerable size.
The formation of new tubers directly from old ones in this manner
is so common as to attract little attention. The unusual features of
the present case were: (1) The large size of the new tubers. Many
of them had a weight of 25-30 grams, several of 50-60 grams, and
one weighed 67 grams. (2) The formation of new tubers within old
Fic. 2. A new tuber protruding from a slit at the bud end of an old seed tuber.
Nat. size. (Compare Fig. 3.)
ones. Fifteen of the old tubers had large new tubers protruding from
slits in their sides (Figs. 1-2). In one instance a new tuber weighing
about 28,grams was wholly included within the parent tuber. This,
like most of the protruding new tubers, was considerably flattened
by the pressure to which it had been subjected during its growth.
This phenomenon of large new tubers within old ones was a sight
calculated to excite wonder in the beholder. To the mycologist it
suggested the bursting of the volva in the egg stage of the phalloids.
STEWART: TUBERS OF SOLANUM TUBEROSUM 425
To one familiar with Gager’s interesting paper! on ingrowing sprouts of
potato tubers it appeared probable that tubers forming on ingrowing
sprouts had enlarged until the pressure produced became sufficient to
rupture the tissues of the parent tuber. Upon dissection of the
tubers this was found to be true.
Fic. 3. The specimen shown in Fig. 2 with one side cut away to show the
origin of the ingrowing sprout which bears the new tuber. A slight change in pose
has brought to view a second new tuber. Nat. size.
The old tuber shown in Fig. 2 was carefully dissected to determine
the origin of the sprout bearing the new tuber which was emerging
from a slit at the bud end. It was found to have started from the
“eye’’ on the right side of the old tuber where the two external, sessile
new tubers are seen. Fig. 3 shows the same tuber, in a slightly
different position, with the tissue cut away so as to expose the new
tuber and the sprout bearing it. In accomplishing this one of the
1Gager, C. Stuart. Ingrowing sprouts of Solanum tuberosum. Bot. Gaz. 54:
515-524. I9I12.
426 BROOKLYN BOTANIC GARDEN MEMOIRS
small external tubers was removed. The changed pose in Fig. 3
brings to view a second new tuber not shown in Fig. 2.
The larger of the new tubers was borne on a very short branch
three centimeters from the point of origin of the sprout; and the
smaller one on a similar branch about a centimeter beyond. The
course of the ingrowing sprout was perpendicular to the surface of
the parent tuber at the point of origin. Apparently, the direction
of growth had been inward from the beginning. Close observation of
this and some other specimens revealed nothing to indicate that the
sprouts had started externally and turned inward. Whether it was
the tip of the sprout or the expanding tubers which first broke through
the cortex cannot be determined in this case; but in other specimens
(among them the one shown in Fig. 1) it was clear that the new tuber
had been responsible for the rupture of the cortex.
The ingrowing sprouts exhibited the lenticel-like openings observed
by Gager. These signify nothing except that the sprouts were
formed in a humid atmosphere. The fibrous roots observed by
Gager were lacking and the sprouts were but slightly branched.
The strange behavior of these tubers cannot be ascribed to low
vitality. This is shown by the fact that a large number of tubers
from the same lot were planted and a good stand of vigorous plants
obtained.
In the main, these observations agree with those made by Gager
and add nothing to them except to show that tubers of considerable
size may form on ingrowing sprouts and produce a striking freak of
nature. The internal tubers observed by Gager were small ones.
THE DUPLICATION OF A LEAF-LOBE FACTOR IN
THE SHEPHERD’S-PURSE!
GEORGE HARRISON SHULL
Princeton University
In two previous papers (Shull, 1911, 1914) I have demonstrated
the existence of two independent Mendelian factors in the shepherd’s-
purse (Bursa bursa-pastoris), each of which produces the triangular
form of capsule. In the latter paper I discussed at some length some
of the criteria and the significance of such “‘duplicate’’ factors. I
gave also a practically complete list of the relevant literature which
had appeared before 1914, and called attention to certain miscon-
ceptions which had found expression in a number of the papers cited.
It is not necessary, therefore, in presenting a new case of dupli-
cation of factors in this species, to repeat at any length the discussion
in this earlier paper. It is important however to direct attention to
the discussions there presented, since several papers along similar
lines, or on closely related matters, which have appeared more recently,
do not include a reference to my paper, even when from the terminology
used it is evident that the authors have had it before them. Several
writers are now making the desired distinction between “duplicate”’
and ‘“‘plural”’ factors, and it is to be hoped that in the future, in the
interest of precision and accuracy, all those who discuss size-inheritance
and related phenomena, will abandon the expression “multiple”
factors because of its erroneous implications.
It has been shown (Shull, 1909, 1910, I911) that the form of leaf
in shepherd’s-purse is controlled by certain Mendelian genes which
have been designated Aa and Bb, the presence of A resulting in an
elongation of the primary lobes of the leaf, while the B gene divides
the leaf to the midrib, and brings to light certain characteristic second-
ary lobing. The several possible combinations of these genes give
the four rosette types: AB = heteris, aB = rhomboidea, Ab = tenuis,
and ab = simplex. ‘These four forms are illustrated in Figs. 1-4.
I have now studied the progenies of a considerable number of
wild Bursas from places as diverse as Chile, Hawaii, Japan, China,
1 Contribution from the Station for Experimental Evolution, of the Carnegie
Institution of Washington, and from the Genetical Laboratory of Princeton Uni-
versity.
427
428 BROOKLYN BOTANIC GARDEN MEMOIRS
(x71) (172) (183) (184)
Fic. 1.—heteris. Fic. 2.--rhomboidea.
(301) (302) (303) (304)
Fic. 3.—tenuis. Fic. 4.—simplex.
Fics. 1-4. Climax leaves from eight rosettes representing the four phenotypes in
pedigree No. 15406. Heteris and rhomboidea possess one or more B factors; tenuis
and simplex lack them.
SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 429
Australia, Tasmania, India, Ceylon, South Africa, the Sahara, and
from widely distributed points in Europe and North America, and find
that the forms everywhere fall into one or more of these four rosette
types. This does not mean, however, that with respect to leaf-form
there are only four biotypes of this species in existence, for nearly
every lot of material from a new locality presents minor details of
lobing which lead to their easy recognition as new and distinct biotypes.
In all of the earlier crosses between types respectively dominant
and recessive for either of the above-mentioned character-pairs, there
appeared in the F. close approximations to the monohybrid ratio,
3:1, or undoubted modifications of that ratio,—the modifications
being due, in most cases at least, to the facts (a) that the A factor has
been in some combinations not completely dominant, and ()) that
both A and B require for their manifestation a certain minimum oppor-
tunity in the way of favorable environment, including cultural treat-
ment.
These results having been well established while the situation in
regard to the capsules called for further extensive investigation, a
number of my cultures which concurrently involved the rosette char-
acters and the capsule characters, have been grown under conditions
not ideal for the development of the leaf lobes, though adequate for
the determination of capsule form. For this reason my records with
respect to the leaf types in certain families are of such incompleteness
as to make the recorded ratios of no particular value. In nearly all
cases, however, a small portion of each pedigree has been given suf-
ficiently good treatment that the composition of the several families
with respect to the rosettes could be inferred with small probability
of error.
The discovery to be detailed below, that in certain races there are
two independent Mendelian factors which affect the leaf-form in
identical ways, each dividing the leaf to the midrib and bringing out
the secondary lobing which is seen unmodified in rhomboidea and modi-
fied by the action of the A factor in the case of heteris, has revived my
interest in the inheritance of the rosette characters and investigations
are now in progress which I hope will give in time a full insight into
the composition of the rosettes with respect to the major factors affect-
ing the leaf lobes.
The duplication of the gene which produces the triangular capsule
has been found almost universally distributed geographically, as will
be shown in detail in a later report, but the duplication of the leaf-
lobe factor, B, appears to be relatively much less frequent.
Before presenting the evidence of dimery in respect to the B
lobes of the leaves, it will be advantageous to have before us the
29
430
BROOKLYN BOTANIC GARDEN MEMOIRS
cases of monomery with respect to this character, as indicated by
the occurrence of monohybrid ratios in the F: from crosses between
plants respectively having and lacking the characteristic ‘B’’-lobing.
TABLE I
Composition of F, Families which Show Monohybrid Ratios, from Crosses Involving
the Presence of the B Factor in Wild Biotypes of Shepherd's Purse
Origin
New Carlisle, Ohio
Chicago, Illinois
Cardiff, Wales
Groningen, Holland
Pedigree Numbers
Pi Fi F,
== 040 | 054
040 056 06130
13148 15361
X |14359 |15362
1376 15363
= 040 | 0514
0515 | 0693 |07203
1337
x |14357 [75358
13214 15367
X |14361 |15368
13179 15369
| 1315 15324
X |14348 |15325
1376 15326
1317
15327
>< fu
mo 4349 15328
1317
1338 5330
1319 15331
X {14352 |15332
1338 15333
1333
X |14353 |15334
1376
1335 15340
X |14355 |15341
1376 15342
120 |13219 |14523
. | rhom-
heirs b oidea
II4 53
= || Aig
194 58
81 17
163 70
99 am
94 are
a7
— 20
= 49
= 28
164 54
40 10
40 7
24 5
128 49
103 43
1875. 35
24 II
28 18
108 23
=e | 847
— | 104
a 66
= II
al ATE
tenuis
Ab
sim-
plex
ab
16
72
Ratio
B:b
2.65:
3.01:
3-45 :
2.88 :
2.58 :
3:01 3
2154n:
BES Sc
1.96 :
SHULL:
TABLE I (continued)
DUPLICATION OF A LEAF-LOBE FACTOR
431
Origin
Berlin, Germany
Landau, Germany
Vicenza, Italy
Peking, China
Pedigree Numbers
Py
13218
x
1376
13222
x
1376
13223
x
13208
13226
x
1376
056
x
059
059
Fy F2
15373
14365 |15374
15375
14368 |15382
ONT 3383
15385
14370
BIO rs a87
15394
14373 |15395
(15396
0688 06212
06196
0689 Garay
12347 |13182
12348 |13189
13190
112349 |I3I19I
13192
13200
1235 13201
140 |15586
Pee)
1542C
14385 15421
[Baden
heterts
rhom-
boidea |
>
aB
56
33
25
tenuts
Ab
9.33 :
2.54:
6.25 :
All of the cases of monomeric B lobes which I have thus far demon-
strated in wild plants are shown in Table 1.
2 These families may possibly have a duplication of the B factor, but the evidence
for.such duplication is much less adequate than is the evidence for the occurrence
of a 3: I ratio in one or more F2 families from these same crosses.
It should be
observed however, that both 15:1 and 3: 1 ratios might occur in different F»
families from the same cross, since the original wild plant used inthe cross might
have been homozygous for one B factor and heterozygous for the other.
432 BROOKLYN BOTANIC GARDEN MEMOIRS
Several of these ratios deviate widely from the 3 : 1 ratio, but in
each such case the inclusion in the monomeric group has been based
on some special consideration. In some cases one or more pedigrees
from the same parentage gave a convincing approximation to the 3 : 1
ratio when grown under favorable conditions; in other cases small
samples of the families have been grown under good conditions, and
the inclusion of the particular pedigrees in one or another of the tables
has been based upon the constitution of these small well-grown samples
regardless of the indecisive ratios displayed by the family as a whole,
when grown under conditions which tended to suppress the dominant
leaf characters here under discussion. In still other cases a number
of families belonging to later generations have been grown and have
given full confirmation of the classification of the wild biotype from
which the pedigree in question originated. Because of the pre-
liminary character of the present report, it is not considered neces-
sary to present in greater detail, the evidences in support of the
conclusion that the families included in this table have a single B
factor. It need only be stated that families whose records are inde-
cisive for the particular point at issue, have been included for the
sake of completeness, and to avoid the immorality of arbitrarily
selecting for presentation those cases which are deemed to support
convincingly the author’s hypothesis.
Seeds of a specimen of shepherd’s-purse, received March 31,
1911, from Tucson, Arizona, through the kindness of Dr. D. T.
MacDougal have yielded a pedigree line which has given me much
difficulty in the classification of the rosettes, even under the most
favorable environment I could provide, owing to the fact that in this
particular strain there is so strong a tendency to precocious develop-
ment of the stems that the leaf characters even in the climax leaves,
are frequently of the relatively undifferentiated juvenile type. It
was just in this difficult material that, during several years, two facts
impressed me with the probability that there were present in this
strain two independent factors corresponding with the B factor of
the above notation. These facts were (a) the occurrence of a rela-
tively small number of tenuis (Ab) plants in two Fy, families (11413,
11414) derived from a cross between the Tucson biotype and a speci-
men of B. bp. tenuis from the eastern United States. No exact
count of the tenwis rosettes in these F2 families was made, but it
was noted that these fenwis rosettes were sufficiently distinct from the
rest of the family, that their number could probably have been deter-
mined with small degree of error. Only desultory attention was being
given at that time, however, to rosette characters, owing to seemingly
insurmountable difficulties of classification in this biotype, and to my
SHULL: ‘DUPLICATION OF A LEAF-LOBE FACTOR 433
interest in certain peculiarities of inflorescence and capsule characters,
which will be discussed elsewhere. (b) The second fact which sug-
gested the probable occurrence of duplication of the B factor in this
strain was the frequent preponderance of rhomboidea-like plants in
the pure-bred families. A large portion of the plants which, because
of precocious development, did not reach the full typical adult con-
dition, showed, nevertheless, highly developed rhomboidea characters,
even when no trace of the elongated A lobes was discernible. Other
plants in the same pure-bred families showed conspicuous elongation
of the primary lobes, thus making certain that the A factor was
present. If two B factors were present, namely B and B’, associated
with only one A factor, the greater ease with which the A character,
as compared with the B character, was suppressed by unfavorable
conditions, would be readily comprehensible.
With this clue to a possible interpretation of the rosette characters
in the Tucson strain, especial note was taken of the F2 families from
crosses between the same Tucson biotype and B. Heegeri simplex,
but in every case the number of individuals which fruited with rel-
atively undifferentiated ‘“‘juvenile’’ condition of the rosettes was so
great that the F. ratios gave no fully convincing proof of the correct-
ness of the hypothesis. The results of such crosses are given in
Table 2.
TABLE 2
The Composition of the Fz, Progenies from a Cross between Bursa bursa-pastoris heteris
from Tucson, Arizona, and B. Heegeri simplex, grown at the Station for
Experimental Evolution
WE es : heteris |rhomboidea tenuis simplex Ratio
Pp F F AB aB Aéb ab B:6
1 1 2
11505 13202 87 44 | 76a | 4 6.55 :1
m< 12353 13203 189 oor 9 18 12 $50 5300
11425 13204 76 Foy 5 | 13 9 OnvAucat
Fs 13205 | 18 Tl 7 (0) WALA, Se
oS sane 13206 45 42 4 8 7.25: 1
11424 354 13207 35 18 6 4 5.30: 1
13208 142 122 33 14 5202 il
diotals..- <... Boh oh eed ods! [le okemex ease ta | 592 364 97 51 6.46 : 1
HERTDOCLEO Sle cat eter ell tage tse 771 257 iyi 18s 15.00: I
The results in the several families were quite consistent, but the
deviation from the expected 15 : 1 ratio were in all cases very con-
siderable. If it is taken into account, however, that the demon-
strated difficulty in genetical studies with this Tucson strain arises
434 BROOKLYN BOTANIC GARDEN MEMOIRS
from the suppression of dominant characters it will be obvious that
these ratios are modifications from a higher ratio (e. g., 15 : 1) rather
than from the 3 : I ratio to which the empirical results in fact more
closely approximate.
A new attempt to test the constitution of the rosette in the shep-
herd’s-purse of Tucson, was made with seeds received from Dr.
Forrest Shreve on November 29, 1914. The same care was given to
the culture of the families involved in this new experiment as was
exercised in those recorded in Table 2. The better greenhouse facili-
ties available at Princeton as compared with those at Cold Spring
Harbor, where the previous cultures had been grown, made it possible
to secure a closer approximation to the expected ratios, as seen in
Table 3, the deviation being the same as before in direction but less
in amount.
TABLE 3
The Composition of the F, Progenies from a Cross between Bursa bursa-pastoris heteris
from Tucson, Arizona, and B. Heegeri simplex grown at Princeton University
Cee es) heteris |\rhomboidea tenuis simplex Ratio
l AB aB Ab ab | B:6
P, | F, F>
ee | 14387 | 1543 143 76 Il 8 11.53: 1
1338 15424 14 | of | 2 I | 7.00 1s
ie io. ee ney Wr meee Weer 157 owl). 4ct2 9 10.91 : I
Bapected. 0. ve een ee nee 185 Or I2 4 I5.00:1
Fortunately for genetical studies on the rosette characters of
shepherd’s-purse, such extensive suppression of characters occurs in
nearly all of the other biotypes which have been under observation,
only as a result of distinctly unfavorable environmental conditions.
The study of a large number of these biotypes from other regions,
in crosses with B. Heegeri simplex, has brought to light several other
cases of probable duplication of the B factor, as shown in Table 4.
The cultures in Table 4 also show for the most part distinctly
defective ratios, due certainly in the main to the fact that they were
being grown in an extensive study of the capsule determiners, and that
in consequence suitable conditions were not provided in many cases
for full development of the rosette characters. Here and there a
ratio closely agrees with the expected F2 ratio, 15 : 1, the best fits
being seen in certain families grown from seeds from Bremen, Germany.
By chance these families from Bremen grew under more favorable
conditions than many of the other cultures listed in this table and
this fact doubtless explains in part why they show a closer approxima-
SHULL: DUPLICATION OF A LEAF-LOBE FACTOR
TABLE 4
435
The Composition of F, Progenies from Crosses of Bursa Heegeri simplex and Members
of Wild Biotypes of B. bursa-pastoris which Probably Possessed Dimeric B lobes
Origin
Pedigree Numbers
Fy
Groningen,
Holland
Berlin
Bremen
Wales
13237
x
1376
1337
x
1338
14354
14367
14368
14372
14374
14377
14378
14379
14383
14357
F,
15337
15338
15339
15379
15380
15381
15384
15391
15392
15393
15397
15398
15401
15402
15403
15404
15405
15406
15407
15408
15409
15410
15411
15412
15413
15415
15416
15417
15355
rhom-
Rarer Iter sy ae oe ee ae
aB
I12 27 20 2 Gra2ncnle
58 20 12 4 4.65 21>
189 109 14 13 11.04 : I
= 286 = 35 Bhitg, Bi
— 56 == 5 12051
42 8 4 I 10.00 : I
216 86 II 7 16.78 : I
60 13 16 20 2.03:: 1°
53 16 15 7 BulAmcnie
208 38 8 10 1320740
170 26 15 2 PUSS cc
72 25 2 I Beessr ek
116 30 18 7 5ro4e: Te
162 5 2 2 41.75: 1
243 74 4 2 52.83 : I
218 69 14 9 T2*AGie UL
189 73, 13 5 14.56 :1
210 77 15 5 14.35 :1
= I51 == 17 8.88 : 1
= 103 = 12 8.58 : 1
a 106 = II 9.63 : 1
== 282 = 18 T5:670-a1
== 167 = 12 136020201
= 262 == 61 AlseXoy 8 1
a 198 = 23 8.61 : 1
a 202 = 27 FURST
= 298 = 26 Te OFsar
Fe 257 Mi eo eng i5 3 ee
== 85 — 6 DAW 0
’ These families passed the winter in the open field, and suffered considerable
injury.
ratio.
ratios much nearer to 15: I.
This doubtless accounts tor their very great deviation from the 15:1
Small samples from each of these families retained in the greenhouse vielded
Only for this reason are these families included here
436 BROOKLYN BOTANIC GARDEN MEMOIRS
tion to the expected ratios. In families No. 15405 and 15406, which
were given the most careful and detailed study, there was little evi-
dence of the suppression of dominant characters, except that in a
few specimens it was a little difficult to be quite sure whether the
A lobe was present or not and it is not improbable therefore that a
few heteris plants have been erroneously included in the rhomboidea
group, but this does not affect the ratios relative to the presence or
absence of the B factor. On the whole, the three families derived
from F, plants of pedigree 14378, showed the least marked tendency
to the suppression of the dominant lobing, and these families show a
close approximation to the expected ratio 45 AB :15aB:3Ab:1 ab.
The close agreement with this ratio in these families, indicates not
only the duplication of the B factor but also the independence of the
two B factors from the A factor.
While 15:1 ratios in the F2 give evidence of duplication, it is ©
highly important to carry the analysis forward at least into the Fs
generation in order to secure more convincing proof that the B factor
was really duplicated in the dominant parent of the original cross.
Until now the only families beyond the Fz, which have been grown from
material in which the B factor is duplicated, have been derivatives
from the earlier Tucson cultures and, as before, these families were
grown primarily for the study of the capsules, and only incidental
attention was given to the rosettes. The ratios in these families are
also defective, therefore, but they give, nevertheless, strong support
to the hypothesis that the B factor is duplicated in the Tucson plants.
These F; families are brought together in Table 5.
This table has been arranged into the three groups which are
expected in the F3; of a cross involving duplication of determiners.
In the first section are the families which bred true to the B lobing;
in the second section are those which split in the ratio 15 : I, and in
the third section are those which split into 3:1. The results may be
summarized as follows: 19 F3; families contained neither tenuis nor
simplex individuals, seeming to indicate that the 19 parents of these
had at least one of the B factors homozygous; 3 families showed
ratios which may be assumed to represent the 15 : 1 class, showing
that the 3 parents of these had both B and B’ present in the hetero-
as probably having a duplication ot the B lobe. As stated in footnote to Table 1,
15: 1 and 3:1 ratios might both occur in the F2 families grown from plants in a
single F, family, if the wild form used in the cross happened to be heterozygous for
one or the other of the B factors. Thus
BBB'b' X bbb’'b’ = BbB’'d’, yielding 15 : 1, and Bbb’d’ yielding 3 : 1.
This situation appears to have been realized in two cases, involving F, families
14357 from Wales and 14368 from Berlin. These two crosses are included in both
Tables 1 and a.
SHULL: ‘DUPLICATION OF ‘A LEAF-LOBE FACTOR 437
TABLE 5
' Composition of Fs Families from F, Parents Having B Lobes in Crosses between B.
Heegeri simpiex and a Biotype of B. bursa-pastoris heteris from Tucson, Arizona
ae ee heteris rhomboidea | tenuis simplex Ratio
AB ab Ab ab B:b
F2 F3
14476 57 A gras = 1:0
14477 49 155 | i = 1:0
13202 14479 14 9 a = 1:0
14482 22 155 = = 1:0
14483 30 98 = = 1 0)
14487 30 3 | — = TaO
14488 24 68 — = P= 0
| 14489 6 21 = = 180)
13203 | 14490 — 80 — 14 80:1
14495 = 30 = — 130
14496 = 245 == = 120
14499 2 17 — = Ler
14503 2 74. = = ae)
14506 = 115 == == D0
_ 14508 = 60 = = 1,20
14509 28 Ly = == Leo
14511 2 44 = SS DO
13208 14513 56 18 = = Leo
14514 = 13 = — LO
13202 14480 p 93 ; 216 | 16 at a ae 16:22 9°51
13203 14492 II 12 I = 23-00)211
__ 13204 14512 76 42— 5 3 Lislegsy Sit
14478 58 9 19 3 3-05 = I
13202 14484 = 125 — 82 152020
14485 36 I 18 = | 2.06:1
| 14500 59 104 12 2 | AORLE
14504 = 66 — 27 2.44 :1
Seen 14505 3 60 — 24 220200
14510 gI 61 36 | = An 22) ae
zygous state, 7. e., BbB’b’; while 7 families gave ratios which may be
appropriately referred to the 3 : 1 group, indicating that the parents
of these had but one of the B factors and this one heterozygous, e. g.,
Bbb’b’ or bbB’b’. According to theoretical expectation for the number
of families given in Table 5 these three groups should occur in the ratio
4 The occurrence ot one simplex in this family is of unknown significance. It
may represent an unusually defective 15:1 ratio, but the deviation is so much
greater than in any other family that other possibilities seem more likely to be true.
From all my experience with these pedigrees, it seems probable that this “‘s¢mplex”’
individual is merely a rhomboidea modified by the suppression of the B lobing. There
remain also the possibility of ‘“‘loss mutation”’ and of a technical error.
438 BROOKLYN BOTANIC GARDEN MEMOIRS
14:8 :8 instead of 19: 3:7. A nearer agreement might well have
been expected even with this small number of families. The dis-
crepancy is probably due in part to the small size of some of the fami-
lies. Thus among the families which were of necessity included in
the first section of the table there are six families, the largest of which
had no more than 33 individuals and if these families belonged properly
to the second group of the table, only I or 2 individuals of the recessive
type were to have been expected. That some of these small families
which contained no recessives, had them nevertheless potentially
present may be taken as a certainty. Only 5 such families need be
shifted from the first section of the table to the second section, to
bring about as perfect agreement with the expected ratio as is the-
oretically possible among 29 F; families.
A more positive demonstration of the duplication of the B factor
may be expected from the F; families derived from the Bremen bio-
types in which the suppression of the dominant rosette characters is
less extensive. Further experiments involving these biotypes are in
progress.
DISCUSSION
The discovery of a second case of duplication of determiners in
Bursa bursa-pastoris owes its chief interest to the facts pointed out
in one of my former papers (Shull, 1914), (@) that cases of actual
. duplication of genes appears to be rare, and (b) that there is some
likelihood that the duplication of factors may involve ‘a series of
special genotypic phenomena.”
It will be recalled that only in the red pericarp of wheat, yellow
endosperm of maize, the occurrence of a ligule in oats, and of triangular
capsules in shepherd’s-purse, was the demonstration of dimery con-
sidered adequate. Several new instances have been brought to light
more recently, and at least one of these must be admitted as fully
demonstrated (Ikeno, 1917). This relates to the quantity and dis-
tribution of chlorophyll in Plantago major, in which two seemingly
identical factors, G and H, determine independently the normal full
green pigmentation of the plant. Either of these two factors may be
entirely wanting, without modifying in any way the self-green color,
but when both are absent, the plants are conspicuously variegated
with white or pale yellowish blotches and stripes, the total quantity
of chlorophyll being considerably reduced.
Hallqvist (1916) has found the pinnatifid leaves of Brassica Napus
apparently produced independently by two factors, the recessive
undivided type reappearing in the F, in only one plant in 16. As the
lobed plants showed considerable variation the results in F3 will be
necessary to confirm the author’s conclusion in this case.
SHULL: DUPLICATION ‘OF ‘A LEAR-LOBE “FACTOR 439
Of cases in which two or more factors do not produce visibly
identical but only more or less similar results as in the black glume
color in oats (Nilsson-Ehle, 1908, 1909), there are many more in-
stances. These do not represent instances of duplication at all, of
course, though they may be expected to grade into cases which would
be indistinguishable from duplication. Several of the eye-color and
body-color factors of Drosophila appear to be of this nature, and some
real duplication may also be present in this group. Some of these
Drosophila characters should have been included in my former paper,
but they had not been to my knowledge cited as examples of ‘‘multiple’’
factors. They have since been so treated, and with obvious propriety,
by Morgan, Sturtevant, Muller and Bridges (1915). The characters
specifically mentioned by these authors are (a) pink eye-color which
is determined independently by factors associated respectively, one
with the sex (or X) chromosome, and the other with the ‘‘third’”’
chromosome; and ()) dark body-color, which is independently pro-
duced by two genes which have been designated “black”’ and “ebony,”
one in the “second”’ and the other in the “third’’ chromosome.
Black and ebony are not identical but merely so similar that their
separation is not practicable when associated in the same family.
Howard and Howard (1912, 1915) have shown that velvet chaff
of wheat is independently produced by two factors, LZ and S, but here
also the factors are clearly not duplicates of each other, for S produces
short hairs and L long silky hairs, while plants containing both factors
have a mixture of both types of hairs on the glumes. The same authors
have found the long awns of ‘“‘bearded’”’ wheat to result from the com-
bined action of two factors B and T, each of which produces short
awns in the absence of the other, but T produces shorter awns than
B and the T awns are most conspicuous in the distal part of the spike
while the B awns are more evenly distributed on the spike. In this
case the action of both B and T is cumulative, the fully awned
form appearing only when both B and JT are homozygous, 1. e., BBTT.
An exceedingly interesting case of duplication, should it stand
the test of further analysis, is reported by Gates (1915) in a cross
between Oenothera rubricalyx and Oe. grandiflora; for, starting with a
heterozygous type supposedly monomeric with respect to the char-
acteristic red pigmentation of the rubricalyx bud, he secured in the
F, two 15 : 1 ratios and two 3 : I ratios,in addition to one 4 : I and
four 5:1 ratios. In the F; he records 4 families with a 2:1 ratio,
one 3:1, two 4:1, four 15:1, and six pure rubricalyx (4. e., I : 0),
besides three families in which the pigmentation of all individuals
was intermediate. Gates interprets the several 15:1 ratios as
evidence that the R factor has become duplicated, but owing to the
440 BROOKLYN BOTANIC GARDEN MEMOIRS
notorious inharmonies between the inheritance ratios in the Oenotheras
and the expectation based on the usual Mendelian methods of segre-
gation and recombination, one may well suspend judgment regarding
this case as an instance of duplication, until it has been shown by
further analysis of one of these 15:1 ratios, that the rubricalyx
individuals will yield three kinds of families, characterized respec-
tively by the ratios I :0, 15 : 1, and 3: 1, and that these three kinds
of families are produced in approximately the ratio 7: 4:4. Unless
this should be the result of the further study, the 15 : I ratio noted in
several of the F. and F; families must have been brought about by
some combination of circumstances, other than the typical Mendelian
distribution of two duplicate factors for the rubricalyx pigmentation.
Gates discusses at some length two of the several methods by
which one may reasonably suppose duplication of factors to come
about. He seems to imply (Gates, 1915, p. 204) that my discussion
of this subject does not adequately cover the several possibilities. He
then proceeds to present two of the same possibilities as if they were
original propositions of his own. These several possibilities are (a)
the occurrence of independent mutations affecting in the same or
closely similar manner non-homologous chromosomes; (0) the mating
of non-homologous chromosomes; and (c) the transposition of parts
of chromosomes by what I have called a “‘sort of longitudinal crossing-
over’”’ (Shull, 1914, p. 139). Only the first two of these propositions
are considered by Gates and he agrees with me that both of the proc-
esses (a) and (b) have probably actually resulted in the duplication
of factors. He thinks that repeated mutations were responsible for
the duplication of red pericarp color in Nilsson-Ehle’s wheats, and
that mismating of chromosomes will explain the duplication which he
believes to have taken place in his Oenothera rubricalyx crosses.
Upon unpublished evidence Bridges (1917, p. 454) refers to two
cases of duplication in Drosophila which seem to result from essentially
the longitudinal rearrangement of genotypic materials that I had in
mind when suggesting the possibility of ‘longitudinal crossing-over,”’
though the details of the process as understood by him are somewhat
different. He states that a section from the mid-region of one X
chromosome appears to have been removed from its accustomed place
or locus in that chromosome, and to have become attached to the
end of the other X chromosome, its mate. The full account of this
case will be awaited with interest.
Accepting the validity of these several methods of duplication,
one may well ask in each specific case whether circumstances make
possible a judgment as to which method was probably responsible
for the duplication in question. I have assumed that the complexity
SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 441
of the structure of the triangular capsule of Bursa bursa-pastoris as
compared with the Heegeri type of capsule, is strong evidence against
the duplication of the factor for this complex character through inde-
pendently repeated mutations affecting different chromosome pairs
(Shull, 1914, p. 141). The character under consideration in the
present paper, namely the B lobing of the rosette leaves, appears to
justify the same observation. The production of leaves divided at
frequent intervals by sinuses reaching to the midrib, and bearing
characteristic secondary lobes and sinuses, involves the control of the
number and direction of cell divisions through very long and com-
plexly branched cell lineages, and it is scarcely conceivable that such
specific control of these long cell lineages should be exactly duplicated
by independent mutations affecting different chromosomes.
It appears to me much more logical to assume that such a rearrange-
ment of the genotype has taken place that the two B determiners
which are allelomorphic to each other in the homozygous monomeric
strains, assumed new positions, whereby they became associated with
chromosomes belonging to different pairs, and thus capable of inclu-
sion in the same germ cell.
As these two factors, B and B’, are apparently entirely independent
of each other, it may be taken for granted that they are associated
with different chromosome pairs. They could become thus asso-
ciated by either of the two methods, (b) or (c), but in the absence of
known linkage relations, there is nothing to indicate which of these
two methods has been the more probably responsible for the dupli-
cation of the B factor,—whether a rearrangement of whole chromo-
somes or the rearrangement of parts of chromosomes through a so-
called “longitudinal crossing-over.”
All these suggestions as to the origin of duplicate determiners
assume the duplication to be a derivative condition; but it may also
be in some cases the primitive condition from which monomeric and
recessive strains may have arisen as a result of repeated ‘‘loss’’ muta-
tions, as stated in my previous paper (1914, p. 137). Studies of the
geographical distribution of the duplicated factors may throw some
light upon the relative age of the monomeric and polymeric types,
for if wild biotypes almost universally possess the duplicated factors,
it may be assumed that this condition is either primitive or at least
relatively old, while a much restricted and more or less definitely
circumscribed range may be accepted as a criterion of relatively recent
origin from the monomeric condition.
In regard to the B leaf-lobe factor it will be noted by reference to
the tables, that plants showing the duplication (Table 4) have been
found at Tucson, Arizona, at Cardiff, Wales, at Groningen, Holland,
442 BROOKLYN BOTANIC GARDEN MEMOIRS
at Bremen and Berlin, Germany, and perhaps at Peking, China, while
monomeric B lobes (Table 1) have been demonstrated in strains from
all these places except Tucson, Arizona. Besides these places in
which the two types have been found associated together, the mono-
meric condition has been found at Chicago, Illinois, at New Carlisle,
Ohio, at Landau, Germany, and probably at Vicenza, Italy, in which
places duplicated factors for this character have not yet been dis-
covered. Excepting only Landau, Germany, these localities in which
duplication of the B factor has not yet been found, have been repre-
sented in my cultures by only one wild B-lobed plant from each
locality. It may be merely a matter of chance that the first plant
from each of these localities had but one of the B factors. It should
also be noted that from the only region in which monomeric B lobes
have not been found, namely at Tucson, Arizona, only two wild
plants have yet been tested, a number quite too small to give any
confidence in the suggested inference that no biotypes with monomeric
B lobes occur at that place. It is obviously necessary to make the
study of geographical distribution of these B factors much more
extensive before safe conclusions may be drawn as to the primitive
or derivative condition of the B lobe with respect to duplication.
This is a work in which many students might lend assistance by
crossing together the several wild biotypes from their own localities.
SUMMARY
The leaf lobes of shepherd’s-purse are controlled by Mendelian
factors A, producing elongated sharp lobes, and B which divides the
leaf to the midrib and brings to light certain characteristic secondary
lobing. The action of these factors is easily suppressed or obscured
by unfavorable environmental conditions, and the inheritance ratios
are usually more or less defective on this account. In previous papers
both of these characters have been reported to be monomeric, 7. @.,
each was found to be controlled by a single factor.
It is shown in the present paper that two factors, B and B’, exist
in certain strains and that these two factors produce the same char-
acteristic lobing of the leaves, but are inherited independently of each
other and of the factor A.
The biotypes having the B factor duplicated appear to be less
widely distributed than those which are monomeric with respect to
the B lobes. More extensive data are needed on this point, but if
the present indications are confirmed, the relatively less frequency of
the dimeric condition is taken to mean that the duplication of this
factor has taken place at a relatively recent date.
SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 443
The morphological complexity of the character produced by the
B and B’ factors is believed to indicate that duplication has come
about through a physical rearrangement in the genotype rather than
by a repeated mutation affecting in like manner chromosomes belong-
ing to distinct pairs.
My acknowledgments and thanks are due to the following corre-
spondents, to whom [| am indebted for seeds of the wild biotypes which
were used in the present study: D. T. MacDougal, F. Shreve,
J. M. Shull, C. A. Shull, A. H. Trow, Tine Tammes, G. Bitter, FE.
Baur, L. Gross, G. Molon, and T. Z. Chang.
LITERATURE CITED
Bridges, C. B. Deficiency. Genetics 2: 445-465. 1917.
Gates, R.R. On Successive Duplicate Mutations. Biol. Bull. 29: 204-220. 1915.
Hallqvist, Carl Ein neuer Fall von Dimerie bei Brassica Napus. Bot. Not. 1:
39-42; LOL:
Howard, A., and Howard, Gabrielle L. C. On the Inheritance of Some Characters
in Wheat I. Mem. Dept. Agr. India 5: No. I. 1912.
— On the Inheritance of Some Characters in Wheat II. Mem. Dept. Agr.
India 7: 273-285. I915.
Ikeno, S. Variegation in Plantago. Genetics 2: 390-416. 1917.
Morgan, T. H., Sturtevant, A. H., Muller, H. J., and Bridges, C.B. The Mechanism
of Mendelian Heredity. Pp. xiii + 262. New York: Henry Holt & Co.
IQI5.
Nilsson-Ehle, H. Einige Ergebnisse von Kreuzungen bei Hafer und Weizen.
Botaniska Notiser, pp. 257-294. 1908.
— Kreuzungsuntersuchungen an Hafer und Weizen. Lunds _ Universitets
Arsskrift, N. F. Afd. 2, Bd. 5, Nr. 2, pp. 122. 1909.
Shull, G. H. Results of Crossing Bursa bursa-pastoris and Bursa Heegeri. Proc.
7th Internat. Cong. Zoédl. 1907, Boston 1912. ‘‘ Advance reprint ”’ in 1910,
6 pp. 1907.
— Bursa bursa-pastoris and Bursa Heegeri: Biotypes and Hybrids. Carnegie
Institution of Washington. Publ. No. 112, pp. 57. 1909.
— Defective Inheritance-ratios in Bursa hybrids. Verh. d. naturf. Verein
Briinn 49: 157-168. IgII.
—— Duplicate genes for capsule-form in Bursa bursa-pastoris. Zeitschr. f. ind.
Abstamm.- u. Vererb. 12: 97-149. I914.
ISOLATION AND SPECIFIC CHANGE
EDMUND W. SINNOTT
Connecticut Agricultural College
Those regions of the earth which are so isolated biologically that
the dispersal of plants or animals between them and other areas is
difficult or impossible are characterized, as is well known, by large
numbers of species and genera which are peculiar to them or are
‘““endemic.’’ In general, the more definitely isolated the region the
higher is its proportion of local forms. Why isolation should be
associated so universally with the presence of these endemic types
is a problem which has excited speculation. It is evident that once a
local race is ‘established, isolation will operate effectively to maintain
it, both by preventing its dispersal abroad and by excluding invaders
which might supplant it. The difficult problem has been to account
for the actual origin of the endemic types themselves in the first place.
In an attempt to throw light on this problem a study has been made
of the floras of a number of islands which are isolated to a greater or
less degree from adjacent land masses and have developed a large
body of local species and genera of vascular plants.1
Certain elements of the endemic flora in all these islands are doubt-
less not of local origin but are ‘‘relicts,’’ remnants of types once much
more widely spread, which owe their preservation to freedom from
the keener competition of the mainland. These types do not con-
cern our problem. As to just how abundant they are we do not know,
but those forms which stand well apart and have no near relatives in
the islands or elsewhere are probably to be looked upon as relicts.
From the close similarity of most of the endemic species and genera
with others on near-by islands or on the adjacent mainland, however,
and from their frequent occurrence as groups of related forms, it is
evident that the bulk of the endemic element in these floras is actually
of local origin.
Several hypotheses have been put forward to account for the
origin of these endemic forms. Some investigators have pointed to
natural selection as the primary factor, believing that new types are
produced by this agency to fit exactly the peculiar conditions in each
region, rather than a wide environmental range. Others, less con-
1 These islands are New Zealand, Ceylon, Hawaii, the Galapagos, Juan Fernan-
dez, St. Helena, Sokotra and Mauritius.
444
SINNOTT: ISOLATION AND SPECIFIC CHANGE 445
vinced of the efficacy of selection, believe that each region has its
own characteristic environmental complex, different from that of all
others, which modifies directly the germ plasm of the animal and
plant types living under it and stamps upon them their local distinc-
tions. Both of these views regard the environment as the most
important factor in specific change and look upon isolation as the
agency which, through providing a comparatively simple and constant
environment, allows a much closer adjustment to it by the plant and
animal population than is possible on wider areas, and hence leads to
the production of large numbers of local species. Still another view
considers that most, if not all, of these endemic and peculiar forms
would have developed anyway under the progressive evolution of their
type, and owe their local character not to a dependence, direct or
indirect, upon a specific environment, but merely to the fact that
they have been unable to become dispersed abroad.
An analysis of the insular floras under investigation presents certain
facts which have a bearing on the problem. It makes evident, in the
first place, that endemism is by no means uniformly characteristic of
all the elements in the flora but that it occurs very much more fre-
quently in certain of the great groups of vascular plants than in others.
The vascular cryptogams, for example, which comprise an important
part of the vegetation of these islands, include but few species or
genera which are confined to any one island or island group. The
glumaceous monocotyledons—Gramineae, Cyperaceae and Juncaceae
—which are also abundant, are represented infrequently among the
endemic forms, though they are somewhat commoner there than are
the vascular cryptogams. It is in the petaloideous monocotyledons
and the dicotyledons that the great bulk of the endemics occurs
throughout all of these insular floras. Not only hosts of the species
but almost all of the local genera belong to these groups. Certain
families, like the Orchidaceae and the Compositae, often contain
almost nothing but endemic species. How great is this disparity in
the extent to which endemism occurs is evident from the following
table, which is an average of the eight island groups investigated.
Species
Genera
Endemic Non-endemic Endemic Non-endemic
Vascular Cryptogams............. 23.2% 76.8% 1.9% 98.1%
Glumaceous Monocotyledons.......) 31.4% 68.6% 2.2% 97.8%
Petaloideous Monocotyledons...... 59.0% | 41.0% 9:7% 91.3%
TUCORVIEGONS yA vitiec syste s Pants 61.7% 38.3% 11.4% 88.6%
What bearing have these facts on our problem of the origin of
local types? They offer little support, in the first place, to the theory
30
446 BROOKLYN BOTANIC GARDEN MEMOIRS
that natural selection has presided over the appearance of these new
forms, for groups which have developed few or no endemic species are
apparently as successful elements of the vegetation as are those in
which such species have been abundantly produced. In fact, Willis?
has gathered evidence from the flora of Ceylon which seems to show
that the non-endemic species are more successful, as a whole, than the
endemic ones, a fact which militates strongly against the theory of
selection. Of course we are confronted here, also, with one of the
major difficulties urged against natural selection, namely that it can
never create but can only eliminate.
Nor do our figures support the theory that local forms owe their
origin to the direct action of the environment, for such a theory can-
not well explain the abundance of endemic species in some groups
and their rarity in others. It may be argued that the vascular crypto-
gams and glumaceous monocotyledons are more primitive and slow-
changing types than the petaliferous groups, and are thus able longer
to resist the pressure of the environment and to maintain their original
characters. We have little evidence, however, that this is actually the
case. Ferns under cultivation seem to be very plastic, and our knowl-
edge of the genetics of the Gramineae, at least, does not indicate that
they are a particularly rigid group.
Both of these views look to the environment as the factor, either
direct or indirect, which is chiefly responsible for the origin of new
forms, and both are open to the objection (among others) that although
the whole flora is subject to the same environment, these new forms
develop only in certain groups. Our third alternative largely dis-
regards the environment. It looks upon the actual production of
new types as due to factors within the organism rather than in its
surroundings, and considers that the locally developed species and
genera in the floras under discussion would have appeared in these
regions whether isolation existed or not. Isolation is thus regarded
merely as the agency which keeps these new forms local and endemic
by preventing their dispersal beyond the place of their birth. Of
course such a theory allows for the play of selection in weeding out all
new forms which were distinctly unsuited to the environment under
which they appeared.
But is not this view also open to the objection which we have
offered to the others, that it cannot account for the rarity of endemism
in certain groups and its extreme commonness in others? A study of
the methods of reproduction in plants belonging to these two cate-
gories suggests an answer to this question. Vascular cryptogams in
the great majority of cases have bisexual gametophytes and are
* Willis, J. C., The evolution of species in Ceylon, with reference to the dying
out of species. Annals of Botany 30: 1-23. I916.
SINNOTT: ISOLATION AND SPECIFIC CHANGE 447
doubtless almost invariably self-fertilized. In the glumaceous mono-
cotyledons, although crossing is certainly not uncommon, it will
probably be agreed that self-fertilization is also the general rule.
In these two-groups we have noted that local species and genera are
very rare. In the petaloideous monocotyledons and in the great
majority of dicotyledons, on the other hand, the flowers are attractive
to insects and cross-fertilization preponderates. As far as our knowl-
edge goes, there are few petaliferous species which are not at least
occasionally cross-pollinated. These facts are significant when we
remember that it is among such forms that local types are produced in
great abundance. In short, our analyses of these insular floras sup-
ports strongly the theory recently emphasized by Lotsy and others
that hybridization is the most important factor in the production of
new forms; self-fertilized types changing slowly, cross-fertilized ones,
rapidly. The unequal development of endemism in certain floral
elements, therefore, which neither the theory of selection or that of
the direct effect of the environment can well explain, is thus under-
standable as the result of differences in methods of reproduction, and
is quite consistent with the theory that the production of new forms
is due primarily to internal causes.
The evidence brought forward by our study of isolated insular
floras therefore seems to indicate that isolation is not a factor in
evolution but simply in distribution; that it cannot create an endemic
species but can merely keep a species endemic.
SUMMARY
1. Isolated regions are characterized by possessing large numbers
of local, or endemic, species and genera.
2. In the insular floras investigated, endemism is not equally
distributed among all plant groups, the local species and genera being
rare among vascular cryptogams and glumaceous monocotyledons but
very common among petaloideous monocotyledons and dicotyledons.
3. This fact seems to indicate that the environment, whether
acting directly or by means of natural selection, has not been the
determining factor in the development of endemic forms.
4. Those groups which are poor in endemics are predominantly
self-fertilized, those which are rich in endemics, predominantly cross-
fertilized; suggesting that hybridization has been the most potent
factor in the development of these new forms.
5. Isolation is therefore regarded not as the factor which, by pro-
viding a simple, limited and peculiar environment, is responsible for
the origin of locally developed species and genera; but merely as the
factor which, by prohibiting dispersal, maintains the endemic char-
acter of local types which owe their origin to other causes.
THE RELATIONS OF CROWN-GALL TO OTHER
OVERGROWTHS IN PLANTS
ERWIN F. SMITH
Bureau of Plant Industry, U. S. Department of Agriculture
In the time assigned the most I can hope to do is to give the
barest outline of the suggested relationships. Some of these are well
determined; others are only suspected and are mentioned here as
hopeful lines of research rather than as definite conclusions. Indeed,
I am quite willing to admit that our work on crown gall has opened up
more problems than it has settled, but, one way or another, all of this
present uncertainty will make for progress and an eventual better
understanding of the whole mechanism of overgrowth. My own
belief is that all overgrowths are correlated phenomena, are the
response of the organism to essentially similar (but not necessarily
identical) stimuli, the visible difference in response when brought
about by parasites being due to number and location of the parasites,
age and kind of tissues invaded, and volume, direction, and velocity
of the stimulus exerted. In other words, in every case, I think the
stimulus is primarily a physical stimulus due to changed osmotic
pressures rather than a direct chemical stimulus. Overgrowths,
therefore, do not always involve the presence of a parasite although
as observed in nature parasites are probably responsible for most of
them.
I. Factors governing type of overgrowth in crown gall.
A. The host reaction depends on the type of tissue infected.
(1) Vascular vs. parenchymatic. For example, depend-
ing on the tissue in which it originates the vessels
in a tumor may be numerous or few, the paren-
chyma abundant or scanty.
(2) Nexus of cells stimulated, 1. e., unipotent, multi-
potent, or totipotent cells. Thus, according to the
tissues infected by the crown gall schizomycete, we
have it causing either organoid galls or histioid
galls.
(3) Rate of growth. The rate of growth depends on the
readily available supply of food and water, on the
age of the tissues when infected and on the species
448
SMITH: CROWN GALL AND OTHER OVERGROWTHS 449
attacked—some species are not subject to this
disease; old tissues respond slowly.
(4) Individual differences. There are, I believe, indi-
vidual differences in susceptibility as well as species
differences.
B. The host reaction depends also on the activities of the parasite
(Bact. tumefaciens) which are variable.
(1) Loss of virulence on culture media. The cause of
virulence is not known. The effect of long
continued growth on culture media is to
reduce the virulence of the organism and
finally to destroy it altogether.
(a) “Old” vs. “resistant” cultures of Paris daisy
organism. Our first isolation from Paris
daisy was extremely virulent in the begin-
ning but lost all power to produce galls in
about three years. Another isolation which
we called “Resistant Daisy” is now slowly
losing virulence at the end of three and one
half years.
(b) New vs. old cultures of poplar isolation. An
isolation from a poplar tumor was extremely
virulent for some time (several years) but
has now lost all power to produce tumors.
Along with this loss has come a progressive
thickening and toughening of the pellicle
on bouillon. This was true also of the daisy
isolation which lost its virulence.
Apparently this loss of virulence is not
correlated with loss of power to produce
formic acid for, according to the chemist,
the non-virulent poplar organism still pro-
duces that substance. Loss of power to
infect must be related, however, to loss of
some chemical or physical property once
possessed, and surely we ought to be able to
discover the exact nature of this loss. Some
strains of Bact. tumefaciens lose virulence
much sooner than others. One of our
strains (from hop) is still virulent after nine
years on culture media.!
(2) Not every isolation is a distinct strain. 1 speak of
‘strains’? only when I know that cultural
1 Ten years, as this now goes through the press.
450
BROOKLYN BOTANIC GARDEN MEMOIRS
and other differences exist; otherwise, I
speak only of “‘isolations.’”” There are cer-
tainly two of these crown gall strains, and
probably many.
(3) Feeble and virulent strains exist 1n nature, 1. e., there
as variation in virulence of colonies from the
same source—hop, carnation, rose, sugar beet,
etc. The author believes that the crown
gall bacteria not only lose virulence on cul-
ture media but also in the tissue of the gall.
Examples are: (a) of three colonies selected
as typical from plates poured from a hop
tumor in 1910 only one proved infectious;
(b) of six colonies plated from a witch broom
on carnation, all of which looked alike and
typical for crown gall, only one was found
to be able to cause tumors when inoculated;
(c) of seven colonies selected as typical from
a plate poured from a rose gall only three
proved infectious, and of these two were
actively infectious, while the other was only
feebly so; (d) of five colonies selected as
typical from a plate poured from a tumor on
Pelargonium none proved infectious, al-
though in advance we felt quite sure of all
of these colonies; (e) of thirty colonies
selected from plates poured from natural
tumors on sugar beets only five were infec-
tious and all feebly so (Bul. 213, pp. 192-194
and Plate XXXVI).
The first and natural supposition when a
culture has lost virulence is that some in-
truder has displaced the right organism; and
when only certain colonies on a plate are in-
fectious, that the others are intruders how-
ever much they may resemble the right
organism. I cannot say that we have abso-
lutely excluded this hypothesis, to which I
held strongly in the beginning, but we are
gradually coming to believe that it does not
explain all the facts.
Il. Some other types of plant galls.
(1) Nonparasitic intumescences. These can be produced in
several ways: by overwatering and exposure to
SMITH: CROWN GALL AND OTHER OVERGROWTHS 45]
excessively moist air; by exposure to very dilute
vapors of ammonia or of formaldehyd; by applica-
tion of a great variety of soluble substances not
the products of parasites, e. g., copper salts; by
painting the surface with vaseline and other insolu-
ble substances which interfere with the gas exchange;
by freezing lightly (Harvey), etc.
(2) Non-cancerous bacterial tumors—olive knot (due to Bact.
savastanot), beet tumor (due to Bact. beticola). In
these the bacteria are abundant and easily seen
occupying the vascular bundles and the spaces
between cells. Bacterial cavities are produced and
the overgrowths are only superficially like crown
galls. The secondary tumors are not developed
from tumor strands. When the bacteria are intra-
cellular the cells are destroyed.
3) Nematode galls. In galls due to Heterodera radicicola giant
cells, 7. e., several to many nucleate cells, are com-
mon. Parasitic nematodes which do not produce
galls. Here the anal excretions are voided outside
of the plant (Cobb). Occurrence of several-nucleate
cells in crown gall.
(4) Various fungous galls. Parenchymatic vs. woody; perish-
able vs. persistent; witch brooms (see newer work
on crown gall). Parenchyma strands (Dodge).
(5) Insect galls which show features resembling crown galls.
(a) Galls bearing leaves; galls bearing roots; galls
bearing modified trichomes. We have now suc-
ceeded in producing on internodes by bacterial
inoculation crown galls bearing roots, leafy shoots,
flower buds, and modified trichomes.
(b) Galls with cortex carrying bright colors—purple, red,
yellow. Crown galls produced on pale green balsam
stems show a red color in their cortex provided the
plants bear colored flowers, but not if they bear
white flowers. The production of red pigment in
the cortex has been seen also in axillary (teratoid)
crown galls developed on red-flowered Pelargoniums.
Etiolation. Excess of chlorophyll.
(c) Galls opening by opercula—strange but not more
so than twin-leaf fruits opening in a similar manner,
or than a double vascular cylinder in Ricinus with
death of intermediate pith and separation into two
cylinders. (Jour. Ag. Res. Jan. 29, ’17, pl. 37).
BROOKLYN BOTANIC GARDEN MEMOIRS
(d) Galls with very definite and distinct strata of gall
tissue—parenchymatic, vascular and protective,
e. g., cynipid galls. Kiister’s prosoplasmatic galls.
Many insect galls differ from crown galls in that
(1) the parasites are few or reduced to a single
organism, and (2) are extracellular, whereas in crown
gall the parasites are more numerous and are intra-
cellular. Many differences in structure, even of the
more complex galls, can be explained, I think,
by these two differences, especially if we assume
(3) that the kind of reaction depends on the volume,
direction, and velocity of the stimulus, its constant
or intermittent flow, and on location, distance,
and mobility or immobility of the source of the
stimulus. As in various insect galls so in crown
galls, there is a tendency toward the production
of more primitive tissues and of various anomalously
formed organs.
III. Crown galls are formed by extrusion of chemical substances. 1 have
recently produced galls with diluted crown-gall products and
this, it seems to me, suggests a new method of attacking gall
problems in general, especially those in which the gall para-
sites can be cultivated pure in sufficient quantity for chemical
analysis, e. g., various fungi. Striking results have been ob-
tained but many tests are yet to be made with the crown-gall
substances in various dilutions, mixed and separate on a variety
of tissues of responsive ages. Various types of cell growths
have been produced by the action of ammonia, acetic acid,
formic acid, aldehyd, etc. (all products of Bacterium tumefaciens,
the crown-gall organism) in less than killing doses, that is,
various degrees of hypertrophy and hyperplasia of cells and
mixtures of the two have been observed. Sometimes there is
great stretching of cells as in certain fungous and insect galls.
Giant cells in the animal pathologist’s sense of that word,
namely, cells containing several to many nuclei, such as occur
in the common nematode galls, are to be searched for in all
sorts of plant galls and to be produced, if possible, experi-
mentally, 7. e., with gall-forming substances. In due time we
shall be able, I believe, to get these multinucleate cells at will.
Probably they are weakened cells. Two very important things
to be determined are whether the size of the cell depends on
the volume or rate of movement of the stimulus or on the
kind of stimulus, and whether mixed stimuli applied in varying
proportions change the manner of cell reaction.
SMITH: CROWN GALL AND OTHER OVERGROWTHS 453
IV. Other effects of parasitic excretions. I believe also from what I
have seen and have obtained to some extent by experiment
that thyloses, fasciations, distortions of tissues, and various
duplications, simplifications and inverse tissue differentiations
are caused by the excretions of feeble parasites although in
nature probably, all are not so caused.
Finally, I would suggest that the application of chemical
substances in various dilutions to growing plants or plant
organs, such substances in particular as are known or suspected
to be produced by living organisms, or are present in soils as a
result of decompositions, may prove to be a hopeful way of
attacking certain unsolved and difficult problems in plant
pathology, e. g., the aetiology of the mosaic diseases, the
fo)
cause of various growth limitations, etc.
LITERATURE
Those who wish details on crown gall are referred to the following
papers:
For the older work: Science, N. S., April 26, 1907; Phyto-
pathology, Vol. I, No. 1, Feb., 1911; Science, N.S., February 2, 1912;
1* Congrés International de Pathologie Comparée, Tome II, Paris,
1912; 17th International Congress of Medicine, London, 1913, Sec-
tion III, General Pathology; and Bulletins 213 and 255, Bureau of
Plant Industry, to be had from the Superintendent of Documents,
Government Printing Office, price 40 and 50 cents, respectively.
For the newer work so far as published: The Journal of Agri-
cultural Research, April 24, 1916; The Journal of Cancer Research,
April, 1916; Science, N.S., June 23, 1916; The Journal of Agricultural
Research, January 29, 1917; Bulletin of the Johns Hopkins Hospital,
Sept., 1917; and Proceedings of the American Philosophical Society,
Nol. 56, 1917.
Separates of most of these papers may be had from the writer.
CONTACT STIMULATION
GEORGE E. STONE
Amherst, Massachusetts
The experiments presented here have been carried on since 1904,
at which time the writer observed some rather remarkable stimulated
growth responses induced in sunflowers when surrounded by wire
netting. At the time these observations were made we were carrying
on investigations relative to the effect of varying atmospheric electrical
potentials on plant growth, and for this purpose we made use of sun-
flowers established in large earthen pots or wooden boxes located at
different elevations in the open air. In some of these experiments
the plants were surrounded with wire netting and in contact with
the same but not with the soil, while in others (normals) no wire
netting was used. In some instances the wire netting was not only
in contact with the plants, but with the soil in which the plants were
growing; the soil being grounded by the use of copper plates in the
bottom of the boxes and by insulated wire, which led to the earth.
In other instances the plants were in contact with wire netting and
the soil, but were not grounded. The problem under consideration
at that time, however, more particularly concerned itself with the
influences of atmospheric electricity on plant growth, for which
purpose organisms of various kinds, including bacteria, were exposed
to elevations varying from thirty to sixty feet. Some of the earlier
experimenters have maintained that when plants were grown in the
free atmosphere surrounded with wires, they failed to develop, and
would eventually die in consequence of being deprived of the bene-
ficial effects supposed to be derived from atmospheric electricity.
In passing we may state that we have never observed any remarkable
mortality among plants in consequence of their being surrounded with
wires even when the experiments were performed at more or less high
elevation above the ground, and under ideal conditions for determining
the effects of atmospheric electricity on vegetation. Moreover the
growing of plants in conservatories where the electrical conditions of
the atmosphere are quite different from those out-of-doors demon-
strates the fallacy of this idea. On the other hand, we found that
plants were greatly stimulated by wire enclosures, especially when
they came in contact with the plants, and also the same stimulation
was noted when plants were grown thickly together and the leaves
454
STONE: CONTACT STIMULATION 455
touched those of other plants. An electrically charged atmosphere,
however, exerts a marked stimulation on plants and it is possible to
modify the function of organisms located at more or less high eleva-
tions by the use of metal coverings. The old idea that milk sours
more rapidly during thunder storms, and that plant growth is greater
following electrical storms has in reality a fundamental basis. The
discovery of contact stimulation led us to modify our methods of
studying the effect of electrical potential on plants since we found
that when the plants were not in contact with one another, or with
the surrounding wire mesh quite different results were obtained.
The observations and results obtained by contact of plants with one
another and with wires, etc., was so significant that we undertook the
investigation of this phase of the problem at that time, and have
devoted considerable attention to it since. Some of our earlier experi-
ments were conducted out-of-doors and parallel experiments were
carried on in a conservatory. Repeated tests of the air in our con-
servatory with a water-drip apparatus and electrometer have in-
variably shown that under ordinary weather conditions there exists
no atmospheric electricity in conservatories, the glass apparently
acting as a screen. The nature of the stimulation due to contact is
probably in no way associated with atmospheric electrical phenomena,
or at any rate, the growth responses do not appear to be identical
with those resulting from ordinary electrical stimulation. The
response to contact is induced by the use of various materials, such as
wire, twine, wood and metal stakes, excelsior, sphagnum moss, soil
particles or even by the plants being in contact with one another.
The same reactions are produced whether the different contact
materials used are suspended and only touch the leaves of the plant,
or whether they touch both the leaves and soil in which the plants
are growing. While there is no evidence to show that these reactions
are associated with any changes in the electrical tension of the at-
mosphere, surrounding the plants, they may, however, be connected
with electrical phenomenon. The reactions resulting from contact
stimulations are not unlikely quite primitive and universal to plants
and probably common to the lower forms of life in general. Probably
all organs will prove to be sensitive to contact but from our observa-
tion the leaves appear to be especially so. The nature of the reactions
appear to be fundamentally similar to those of touch, from which it
would seem the more highly differentiated reactions of tendrils and
wound responses, etc., originated.
456 BROOKLYN BOTANIC GARDEN MEMOIRS
Fic. 1. Showing growth of sunflowers surrounded by wire netting. Note
difference in size of plants which have penetrated through the wire-mesh enclosure,
a feature of common occurrence.
METHODS
In the study of the effects of contact of various materials with
plants we employed several methods and the experiments were carried
on under different conditions. With the exception of some of the
earlier experiments which were made out-of-doors, most of them were
STONE: CONTACT STIMULATION 457
conducted in a conservatory where the heat, light and soil conditions
were uniform and normal. Some of the experiments were carried on
in direct sunlight; others in darkness, while others again were con-
ducted under bell glasses where it was possible to maintain uniform
atmospheric moisture and transpiration conditions. Contacts with
wire were made with a two-inch-mesh galvanized iron wire netting,
and in some cases a one-inch-mesh wire was employed. These were
made up into cylinders 4 to 6 feet high and varied in diameter from
8 to 26 inches, according to the size and number of plants employed.
These wire cylinders were placed around the plants. In some in-
stances the plants were grown between trellises made of wire netting
placed from 6~10 inches apart, in which case the tips or free end of
the leaves of the plants came in contact with the wire on two sides.
Wooden stakes (dowels) and iron posts driven into the soil were also
employed as contact material, the dowels being used in some instances
to support loose twine which surrounded the plants and in other
cases they were used alone in various combinations. Fishnet made
of twine with a mesh of about two inches was employed in a similar
manner to that of the wire cylinders, and in some cases the plants were
more or less loosely tied up with twine. In the study of the effects
of contact on the stems and roots of seedlings, excelsior was employed
either loosely in long strands, or cut up more or less in fine shreds as
the nature of the experiment required. In the root, contact experi-
ments were made with soil particles which ranged from 16 mm.—0.I mm.
in size, the various grades of material being obtained by sifting through
sieves and bolting cloth. The particles were repeatedly washed and
sterilized and submerged in jars of water, the latter being previously
boiled to exclude air, inasmuch as the presence of air would greatly
modify the growth of the seedlings (1).
EFFECTS OF CONTACT OF PLANTS WITH ONE ANOTHER
When plants are grown close together, as is the case of some crops,
they often come in contact with one another and even in nature con-
tact stimulation plays an important rédle, particularly where certain
organisms grow so close to one another that they touch. The growth
of some coniferous trees is often such that they are much congested,
and in nurseries where many thousands of coniferous seedlings and
other nursery plants are grown close together a contact stimulation
may sometimes occur. We have, however, no authentic data based
upon experiments regarding the behavior of coniferous and other
seedlings grown under nursery conditions. Neither have we attempted
to differentiate growth which may be due to contact in thick stands
of forest growth from those originating from other causes, but some
458 BROOKLYN BOTANIC GARDEN MEMOIRS
gardeners and foresters have intimated that they have observed indi-
cations of an accelerated growth in height as a result of coniferous
seedlings coming in contact with one another. The stimulation effect
of contact, however, can be observed in the growth of crops and the
oie
ees
a
Fic. 2. Showing growth of tomatoes im situ in contact and not in contact with
one another.
method of close growing of certain economic plants has its advantages.
The configuration of plants, however, is greatly modified by close
growing, as may be observed in the handling of single-stemmed
chrysanthemums by florists, the growth of corn and various other
TABLE I
Showing Growth of Tomatoes (Lycopersicum esculentum Mull.) in Contact and Not in
Contact with One Another
Average Height and Diameter of Plants (Centimeters)
Percentage Gain by Contact
| |
| Diameter are z cel ;:
| | Height | Diameter
Normale crete ne are 19.41 | .46 |
Gontacthon.ne sate eee era | 31.10 47 60% 2%
plants. The stimulative effect of contact of one plant with another
is shown in Fig. 2. The tomatoes in this case were grown in a well-
lighted conservatory in soil similar in all respects and the water
supplied was such that each plant obtained similar amounts. (Cf.
Table 1.)
! 36 plants used.
STONE: CONTACT STIMULATION 459
RESPONSE OF PLANTS TO CONTACT STIMULATION WITH WIRES AND
TWINE
The investigations relating to the effects of contact with wires, etc.,
are given in Tables 2-9. All of these experiments were made in a
well-lighted conservatory, the plants being grown in a good uniform
grade of loam either in solid beds or in benches. In this series 1-6
plants were enclosed by wires, or dowels, and twine, and in some
cases only fish netting was employed. The normal plants were in some
instances grown free from contact with one another and in others not.
In the case of only a single plant being surrounded by wire netting
contact would occur only with the wire, whereas when two or more
Fic. 3. Showing growth of castor beans in contact and not in contact with
wire netting. Plants removed with as little disturbance as possible from original
position for photographic purpose.
plants were grown tolerably close together they would eventually be
in contact with each other as well as with the wire netting, etc. Hence
contact stimulation would result not only from the use of wires, etc.,
but from the contact of plants with one another, or in other words the
so-called normal plants were not in all instances free from contact,
inasmuch as when they were grown in groups they would eventually
touch one another and growth would be influenced. We therefore
have two series of experiments, namely: (a) those in which the
460 BROOKLYN BOTANIC GARDEN MEMOIRS
normal plants were perfectly free from contact with one another and
(b) those in which the so-called normal plants were more or less in
contact with one another. Both of these normal series were compared
with those in direct contact with wires, etc. The plants in this series
of experiments all show a gain in height by contact with different
material. The effects of contact with wire netting (2-in. mesh) as
compared with no contact whatsoever in the normal plants is given
Fic. 4. Showing growth of tomato plants in contact and not in contact with
wire,
in Tables 2-4. In the former experiment (Table 2) 12 plants were
utilized, all of which were grown separately, 6 being surrounded by
cylinders of wire netting and 6 grown free from any contact whatso-
ever. The plants in contact with wire netting showed a gain in
height of 31 percent compared with those of the normals and the same
percentage gain is given in Table 3, in which case the.contact plants
STONE: CONTACT STIMULATION 461
TABLE 2
Showing Growth of Sunflowers (Helianthus annuus L.) in Contact with Wire Netting.
Average of One Experiment with Twelve Plants?
Percentage Gain
Average Height in Height by
(Centimeters) Contact
Normal 4 a tise pat eee: 131.9
Contactiwites. sys aee e 173.0 es la
TABLE 3
Showing Growth of Sunflower (Helianthus annuus L.) in Rows between Wire Netting.
Average of Two Experiments with 54 Plants?
Average Dimension and Weight in Centimeters and Grams
; Internodes A Percentage Gain in
Height | Length Weight Height by Contact
Mos eee ee uw ccs ve 2 114.1 | 6.9 307.9
@ontact wire. ... 2.0205. 151.2 8.9 302.9 31%
TABLE 4
Average Dimension and Weight in Centimeters and Grams
Height Diameter Ratemotes Weight Percent Gain in
Stem EAE | Length Height by Contact
INommalas sa... 73 1.8 9.2 | 7.8 457
Contact wire...| 99 1.5 8.1 1232) 284 35%
were grown between parallel rows of wire netting, the netting being of
sufficient distance apart to come in contact with the leaves of the
plant. Neither the normal plants nor those in contact with the wire
touched each other. The experiments with the castor beans shown
in Table 4 were identical with those given in Table 2. This showed a
gain of 35 percent in the contact plants over the normal. The re-
maining 5 tables (5-9) show the effects of surrounding plants with
wire netting, dowels and twine, and fish netting. Since in this series,
from 2-6 plants are grown close together, they also show to a certain
extent the effects of contact with one another. Both the normal and
contact plants were in pairs in Tables 5, 6 and 8a while 7 and 8b were
arranged in threes and those in Table 9 contained 6 plants. In
Table 7a, a wire netting was suspended overhead by twine and did
not come in contact with the soil. From the data given in these
tables it is hardly possible to draw exact deductions as to the relative
* 2Normal plants separated from one another.
%’ Normal plants separated from one another.
4 12 single plants used, 6 in contact and 6 not in contact.
31 3
462 BROOKLYN BOTANIC GARDEN MEMOIRS
value of the various methods employed in inducing responses to con-
tact stimulation. Deductions, however, based upon a large series of
experiments not included here justify us in stating that surrounding
plants loosely and irregularly with twine does not produce the same
degree of stimulation or response, as by the more thorough contact
derived from the use of other material such as wire netting with a
uniform mesh, or, in other words, plants react more pronouncedly to a
larger contact surface than to a relatively smaller one, although there
probably exists a definite size of mesh which would produce the best
result, and this would undoubtedly vary with different species. There
appears, however, to exist some difference in the degree of stimulation
arising from the same size mesh, as shown by the behavior of some
species when the contact is applied to the leaves. The leaves, for
example, of the sunflower and corn do not respond so freely as those of
tomatoes and the castor bean presumably because the leaf apices
are the most sensitive as in the case of tendrils. The latter species,
possessing different type leaves, would appear to act differently on this
account.
TABLE 5
Showing Growth of Castor Bean (Ricinus communis L.) in Contact with Wire Netting’
Internodes | Ae
| Percentage Gain in
| Height | Diameter Stem Si) NAS |) :
Nes iene | | Height by Contact
INormallma-p rss) 3253 1.06 10.3 Bur | 152 |
’ Contact wire...| 55.1 1.18 11.5 4.6 | 200 ‘(| 70%
The data derived from these experiments are not sufficient to allow
of deductions being drawn which would be of any value in determining
the relative value of the various-sized meshes in stimulating growth.
In some cases where galvanized iron netting with a one-inch mesh was
employed, the stimulation appeared to be equally as great as with the
two-inch mesh. Neither is it possible by these tests to determine
accurately the difference in the degree of stimulation which resulted
from the use of wires and that from the contact of the plants them-
selves. In all cases where single plants were employed they were
removed far enough away from one another so as not to touch. The
stimulated growth, therefore, was due entirely to the material which
surrounded them. On the other hand when plants were grown in
such a manner as to touch one another there existed two sources of
contact. In the experiment shown in Table 9, six sunflower plants
®12 plants used. Plants in pairs.
STONE: CONTACT STIMULATION 463
were grown close together (about 8 inches apart) and the two series
of normal plants which consisted of twelve or six in each set were in
contact with one another throughout their period of development.
Hence, the percentage gain in growth in height is due to the additional
contact of the plants to the various materials which surrounded them.
The largest increase in growth is shown in Table 5 but this data was
obtained only by the use of 12 plants and probably is exceptional.
When closely grown plants are surrounded with wire netting, etc., and
especially large-leafed plants such as the common sunflower the
leaves do not have an opportunity to assume their normal shape and
in this way there occurs a tendency to shade the stems. The dis-
placement of the leaves by the wires thus shading the plants or the
slight shadow cast by the wire enclosures apparently has little effect
on the growth in length of the internodes, inasmuch as the same
reactions can be obtained by growing the plants in darkness. The
data concerning the results given in the tables (2-9) follow.
TABLE 6
Showing Growth of Castor Bean (Ricinus communis L.) and Corn
in Contact with Twine®
TONEEE Dimension, etc., in SCE IEE»
Experiments No. of Plants Height Internode: Woden mpc Gain in
Used eanterber Meni Height by Contact
Faye Mormal. 66 se. ae.: 24 ees 7 Ary (eae a
Contact twine.... O2167 TI) a13:0 12%
Corn (Zea Mays L.)
feremermal 2. ..s::si6 7, 36 132.9 TOO be lus teed |
Contact twine.... 145.6 Wenn || away | 9%
TABLE 7
Showing Growth of Sunflower (Helianthus annuus L.). (a) Average of Twelve Plants
in Contact with Wire Netting, (b) Average of Twelve Plants in Contact with
Stakes (Dowels) and Twine’
Percentage Gain
Average Height in Height by
Experiments (Centimeters) Contact
(@) aINonmally. oye eee Tai,
Gontactawitess.) eer 164 19%
ND) pNiOninialiaty Acwieens « ettdak one 132
Contact stakes and twine.178 34%
6 Plants in pairs. 24 plants in (a), 36 in (0).
7 Plants in triplets.
464 BROOKLYN BOTANIC GARDEN MEMOIRS
TABLE 8
Showing Growth of Sunflower (Helianthus annuus L.) in Contact with Wire Netting®
Average Dimensions and Weight in Centimeters and Grams
Experiment | ; Internodes P Gai
eraeiene | eee ai = Weight in Helga
| Number Length by Contact
(@) Normality. ae | 108.80 1.35 14.2 6.8 | 241.5
Contact wire..| 138.91 1:33 14.8 10.4 267.9 27%
(>) Normealeeeence y telitaG: 0.80 19.7 4.2 90.6
Contact wire..| 109.2 0.84 24.5 4.5 92.6 33%
TABLE 9
Showing Growth of Sunflower (Helianthus annuus L.) Surrounded by Wire and Twine.
60 Plants Used in Each Test. Experiment Made in Greenhouse in Benches?
Average Dimension and Weight in Centimeters and Grams
jab | Percent Gain in
Height Diameter | Internode Weight Moisture | Height by
Length | Percent Contact
Normale eo 132.5 1.59 8.03 PUI 87.6
Contact wire......| 151.0 1.46 9.01 216.0 89.6 | 15%
Contact twine..... 155.0 1.43 8.71 210.0 90.4 16%
Contact fish net...) 159.0 1.40 10.50 160.0 9g1.8 | 20%
The average diameter of the stems of the normal was 1.2 cm. and
for those in contact 1.24 cm. The average number of internodes
for the normal was 13 and for the contact plants 13.6; while the
average length of internodes for the normal was 6.8 cm. and that for
the contact plants 8.9 cm. or a gain of 30 percent. The average
weight for the normal plants was 243 grams and for the contact 216
grams. The diameter, number and length of internodes was slightly
greater in the contact than in the normal. The average weight was
I2 percent greater in the normal than in the contact plants. The
moisture contents of the plants were greater in the contact plants
than in the normal ones. The most important difference between
the normal and contact plants is in the length of internodes.
RESPONSE OF PLANTS TO CONTACT WITH EXCELSIOR
All of the experiments with excelsior were made with seedlings and
were carried on in darkness. These were made in large flower pots
containing either soil or sawdust. The excelsior was packed loosely
over the soil or sawdust as the case might be at the time the seeds
8 12 plants used in each experiment. (qa) Plants in pairs; (0) in triplets.
® Plants in sixes.
STONE: CONTACT STIMULATION 465
were planted, to a height of about Io inches and often less, depending
upon the nature of the plant in use. The excelsior was cut in lengths
varying from 1 to I0 cm. or according to the nature of the plant
employed. Since these experiments were all made in darkness and
the plants were in all cases covered with receptacles, such factors as
light and transpiration were controlled, and under these conditions
the seedlings were dependent largely on the reserved material con-
tained in the seeds. There was, however, no contact of the plants with
each other. Moreover in this series, the same contact material was
used throughout, namely excelsior, and any specific reaction which
might arise from the use of different materials in contact with the
TABLE I0
Showing Growth of Horse Beans (Vicia Faba L.) in Contact with Excelsior”
Average Height and Weight in Centimeters and Grams
Experiments Roncoee Gan by
No. ae Height Weight
Height Weight
me Motiial. ...o2% .isteaces ae 149 14.03 | 0.96
Contact excelsior....... To:20s0 |) ees 29% 33%
Mo eiNonmal se 62 aS 374 15.10 0.74
Contact excelsior....... 1G2500) | L:O5 18% 41%
(Ce)) aN \oy sins) Gees 125) L732) || wte37
Contact excelsior....... 20.41 | 1.50 17% 9%
PAWOTACeMOnMall. =... 40.6... 0% 15.48 1.02
BARE Ca 5. olor sginendlh 3 3%, 2 19.03 1.61 23% 58%
TABLE II
Showing Growth of Hypocotyls of Lupines (Lupinus albus L.) in Contact with Excelsior.
Average of Two Experiments with 128 Plants!
Average Height of Hypocotyls (Centimeters)
Percentage Gain
Hypocotyls by Contact
Normale: ear coats rae 19.28
Contact exeelsior:<: . .i6.c0 21.65 12%
plants would not be present in these. There would of course occur
variations in the stimulus imparted to the different species owing to
difference in the relative degree of contact of the plants with the
excelsior—a feature which would be determined by the fineness,
compactness and amount of the excelsior employed, as well as by the
10 Experiments in Which light was excluded.
11 Experiments in which light was excluded.
466 BROOKLYN BOTANIC GARDEN MEMOIRS
surface area of the organs in touch with the same. Inasmuch as the
plants used were seedlings and a larger number were employed than
in the preceding series, the results are likely to be more uniform.
On the other hand it should be noted that with the use of seedlings
grown under good heat conditions (in which case they would develop
rapidly) the duration of stimulus would be much less prolonged, and
the ultimate effect of contact on the configuration of the plants
would be less pronounced. The most marked stimulating effect of
contact would occur in general in those experiments which were the
most prolonged, namely with the larger seedlings. With the excep-
tion of corn all of the measurements given are either of the hypocotyls
or stems. The results of these experiments are given in Tables 10-15,
all of which show a stimulated growth due to contact with excelsior.
The average weight in all instances where determined was greater
in the contact than in the normal. In one water determination with
lupine, there was 4 percent more moisture in the contact-stimulated
plants than in the normal. In the experiments with corn (Table 14)
measurements of leaves were taken as well as the cotyledons although
it was our original intention to include in our measurements the
cotyledons only, since when the leaves break through the cotyledons
the growth of these organs is greatly retarded. The data giving the
measurements of leaves in the corn, however, have a limited value as
the growth of the leaves displayed more or less erratic behavior. In
one experiment the leaves protruded above the excelsior and conse-
quently they were not in contact. In another instance the leaves
showed a very decidedly accelerated growth in the normal plants
which was caused by a brief and accidental exposure to light. By
taking proper precautions in further experiments this did not occur
again. Some of the more sensitive cotyledons of the Gramineae are
characterized by a marked growth correlation following mutilation
or decapitation of the cotyledons in etiolated seedlings, and similar
reactions occur to etiolated seedlings which are exposed to light.
The function apparently of the cotyledons is to protect the true
leaves in protruding upwards through the soil and as soon as they
are exposed to light their growth ceases rather abruptly. Correlated
with the retarded growth of the cotyledons is a greatly accelerated
growth of the leaves which may amount to over I00 percent increase
in two or three hours. The exposure of the cotyledons to light even
for a brief period is sufficient to check their growth and greatly ac-
celerate the development of the leaves and this is what happened on
one occasion.
The normal plants would be the most affected by any such exposure
as they were not covered with excelsior, while the contact plants
STONE: CONTACT STIMULATION 467
TABLE I2
Showing Growth of Peas (Pisum sativum L.) in Contact with Excelsior. Average of
Two Experiments with Eighty-two Plants’
Average Height of Plants (Centimeters)
Percentage Gain
Stems by Contact
InGaAs Be gee ee ae oie ee 13.07
Contact excelsior.......... 16.66 27%
TABLE 13
Showing Growth of Cucumbers (Hypocotyls) (Cucumis sativus L.) in Contact with Ex-
celstor’
Average Height of Hypocoty] (Centimeters)
Percentage Gain
Hypocotyls by Contact
Normale water ete dove. 15.04
Contact exeelsion. y2h05% 04: 16.30 8.3%
were to a more or less extent protected from light by the excelsior.
One of the experiments with lupines which was carried on at the same
time behaved in a similar manner to that of corn. In one case 31 out
of 36 of the normal plants developed leaves averaging 7 cm. in length
while none developed in any of the excelsior contact plants—a feature
due to the same cause, namely, to a brief and accidental exposure of
the plants to light.
TABLE 14
Showing Growth of Cotyledons and Leaves of Corn (Zea Mays L.) in Contact with Ex-
celstor. Average of Two Experiments with 72 Plants'4
Aves Tenth of Gaby ledons and Leaves (Centimeters)
| Percentage of Gain by Contact
Cotyledons Leaves Pa ;
| Cotyledons Leaves
IGRISIINANIL, cous, sich a) ckojeholte tees 8.41 ee | as a7 |
(Contact excelsior....-- 0.0... 3 10.07 18.44 19% 4%
TABLE 15
Showing Growth of Turnip (Hypocotyls) (Brassica Rapa L.) in Contact with Excelsior.
Average of Two Experiments with 194 Plants
Average Height of Hypocotyls (Centimeters)
Percentage Gain
Hypocotyls by Contact
INOKIMa Mee, sea 8.57
Contact excelsior........... 9.66 11%
2 Experiments in which light was excluded.
18 Experiments in which light was excluded. No. of plants used 26.
4 Experiments in which light was excluded.
18 Experiments in which light was excluded.
468 BROOKLYN BOTANIC GARDEN MEMOIRS
RESPONSE OF Roots TO CONTACT STIMULATION
Excelsior
In the experiments so far enumerated no account has been taken
as to the effects which various substances coming in contact with
stems may have upon the growth of roots and other organs. It is
quite natural to expect that if one organ is affected by a stimulus
other organs will be, inasmuch as the organism as a whole responds
to stimuli of quite insignificant character. Incidentally, we observed
in our early experiments with seedlings grown in sawdust that the
roots in some cases respond when the serial portions of the plant were
in contact with excelsior. In I912 experiments were carried on for
the purpose of comparing the growth of roots in soil under different
conditions. These experiments consisted in the growing of roots in
boxes with glass sides. In one series the roots were grown in holes
or channels along the edge of the glass, and in the other series no
holes were provided. In both series there would be contact, but
where the roots followed the channels, there was less contact of the
roots with the soil particles than where they had to force themselves
through the soil. These experiments being limited, however, did not |
furnish data of any particular value, although from the more or less
meager data obtained, they seemed to indicate that the presence of
grooves or channels in the soil produces less stimulating effect on the
growth of roots than when more thorough contact exists. Most of
our investigations relating to the effects of contact on roots were
made with excelsior, and in some cases sphagnum moss and cocoa
fibers were used. In many experiments we have also utilized various-
sized particles of gravel, sand, and silt suspended in water. The
excelsior which we employed did not produce any bad effect on the
growth of roots when submerged in water. On the other hand it
-appeared to clear up the water in some instances, inasmuch as it
was apparent that it absorbed certain accumulated products which
sometimes occurred, and which were more or less detrimental to the
growth of the roots. The water containing the excelsior cultures
was clearer and possessed a more agreeable odor than those cultures
where it was not used. As the seeds were suspended over water on a
fine-mesh cotton cloth netting, these would occasionally become too
moist and in this way the water became more or less turbid, owing to
the presence of various extract substances derived from the same.
The presence of excelsior in the water had a tendency to prevent any
abnormality in the growth of the roots which might follow from the
presence of foreign substances, and in this respect it acted like sand
and charcoal in removing certain impurities such as copper sulphate.
STONE: CONTACT STIMULATION 469
TABLE 16
Showing Growth of Roots of Lupines (Lupinus luteus L.) in Water Containing Excel-
sior and Sphagnum Moss'®
Average Length of Primary Roots (Centimeters)
Treatment Roots
Normale cron ci eae 752
Sphagnumimosse 45-40. 2: 8.8
Excelsionicoarse::y-144-yaae 10.4
For the purpose of obtaining more uniform conditions for root
growth, we subsequently adopted larger containers provided with
loose partitions, in which case all the plants were subject to like con-
ditions. The excelsior experiments were made in either glass or
earthen jars with tap water which had been previously boiled to
exclude the air. The excelsior, which was of the ordinary commercial
form, was in long strands of more or less irregular shape and in cross
section was about I mm. in diameter. We employed three different
grades, namely, the coarse, loose, curly form as it is obtained com-
mercially, and the same cut from I cm. to 4 cm. long. The uncut
grades were packed loosely in the jars containing the water, while
the finer grades were much more compact and greatly increased the
contact surface to which the roots were subjected. In all cases the
excelsior was boiled before using it to exclude air, inasmuch as the
presence of air would be capable of modifying growth. The seeds
employed were of a good quality and were separated by sieves and
carefully selected before planting, which insured a uniform size and
TABLE 17
Showing Growth of Stems and Roots of Peas (Pisum sativum L.) in Water Containing
Excelsior. Average of Four Experiments
Average Length of Stems and Primary
Roots (Centimeters)
Treatment Stems Roots
INonmial hak: fa tal Gas Sort tasers Tez 4.5
Excelsionc0arse.cce ae ee LOS 7.0
Excelsior Ane 02.0 tenes boom « II.1 8.8
Secondary roots predominated in the fine excelsior, many in the coarse excelsior,
practically absent in the normal.
corresponding germinating capacity. The experiments were con-
ducted in a dark place, although in this series not in absolute darkness.
The amount of light, however, which prevailed was insufficient for
photosynthesis or for phototropic curvatures. Neither the stems nor
roots were in contact with one another, hence, any stimulating growth
16 5 plants used in each test.
17 One hundred and sixty plants used.
470 BROOKLYN BOTANIC GARDEN MEMOIRS
which followed was due to the excelsior alone. All of the plants were
in the seedling stage and the duration of the experiments in no case
exceeded fifteen days. The investigations relating to contact stimu-
lation of roots were limited to the use of a few species which show
different types of reaction. The results of these experiments are
shown in Tables 16-18, although several other species were employed
which are not included here. In all cases, the ‘“‘excelsior coarse’’
implied the loose commercial form which came in contact with roots
occasionally, while ‘‘excelsior fine’’ was in these particular experiments,
cut up into lengths 4 cm. long and furnished considerable contact.
When such plants as lupines and peas which possess strong and fairly
good-sized primary roots were in contact with excelsior the reaction
was characterized mainly by an accelerated growth of the primary
root system, together with considerable secondary root development
whereas in the case of delicate roots, such as mustard (Table 18 ), the
growth of the primary roots is less and the secondary root development
is greatly accelerated. Practically little or no difference existed in
the growth of hypocotyls and stems in any case, although this feature
is not always constant. In both the lupines and peas (Tables 16 and
17) there occurred a considerable accelerated growth of the primary
roots. In the latter case (Table 17), which represents an average of
four experiments, there was a decided increase in the number and
length of secondary roots from the normal to the fine excelsior. In
one experiment where the secondary roots were counted, their average
number was as follows:
INorinial. sy cee cen bot rete oe aie ie Ree EES 17
Goarsevexcelsioncn apy uyee ee ete at cicusieuste sete are 32
inlesexcelSIOrs ays, sessieete Sas oe eae OE eile ene ee 40
This feature was more pronounced in the mustard, however, where
the primary roots in contact with the excelsior were much less de-
TABLE 18
Showing Growth of Hypocotyls and Roots of Mustard (Brassica alba Boiss.) in Water
Containing Excelstor'®
Average Length of Hypocotyls and
Primary Roots (Centimeters)
Treatment Hypocotyls Roots
Normals: f:)2t ees ta eee eR L250 II.1
ExcelsiOricoarsen. aerseieneeine ie 6.8
EsxeelsiorGines.. 20s. es s1ae 11.9 8.9
veloped (Table 18). In this case there were no secondary roots on
the normal plants; many in the coarse excelsior and very numerous
18 20 plants used in each test.
STONE: CONTACT STIMULATION 471
and well developed on those in contact with fine excelsior. The
number of secondary roots or those in contact with the fine excelsior
averaged 18 to a plant and the ratio of the total length of the entire
root system of the normal and fine excelsior was 1 to 6 in favor of
the latter. Measurements made of the total surface area of a single
typical root from one of the normals and one of the fine excelsior showed
that the total surface area of the latter was over three times that of
the normals. These experiments show, at least in young seedlings,
that roots respond to contact and that the response is confined very
largely to these organs, although more than one type of growth
correlation may occur. They indicate also that different species will
respond to contact in a different manner. In other words, secondary
root development is stimulated more in some species than in others
by contact, and this excessive development of the secondary root
system is correlated with a lesser development of the primary root
system.
Soil Particles
Since roots are sensitive to contact to various materials it would
naturally be supposed that the nature of the soil constituents or
particles would exert an influence upon growth and configuration of
plants, and particularly upon the root itself. As contact is effected
by the surface area involved, variation in the size and shape of the
soil particles would be expected to produce different results. Conse-
quently, a series of experiments were carried on, but not completed,
with the idea of determining what effect, if any, soil particles have
on the growth of roots, and how the various-sized particles effect
development. For this purpose we had at our disposal several care-
fully prepared grades of gravel, sand, silt and clay which had been
sifted through sieves and bolting cloth. The size of the particles
ranged from 16 mm. to 0.1 mm. and in some cases to.05 mm. The ex-
periments were conducted in glass jars filled with water previously boiled
to exclude air. Each jar was completely filled with some particular
grade of material which had previously been thoroughly washed with
water and sterilized. We thus had a medium in which the particles
of gravel, sand, etc., were surrounded by water, and as far as possible
free from air. A fine-mesh cotton netting was placed over the jars
on which rested the seeds, and as germination took place the radicles
penetrated downward between the submerged soil particles. All
experiments made with soil particles in water were carried on in
darkness in a moist chamber where transpiration was limited and
the temperature condition alike. The plants were in fact under
identical conditions throughout, except as regards the substratum.
472 BROOKLYN BOTANIC GARDEN MEMOIRS
The normal or check series were run in water alone. The results
given by these experiments, which were limited in numbers, are
similar to those obtained by the use of excelsior in contact with roots
in water, namely, the various grades of gravel, sand, and silt gave
rise to different reactions on the part of the organism which resulted
in a stimulated growth correlation, as exemplified in the development
of the primary organs and secondary root system. The results ob-
tained by the use of mustard, peas and soy beans, etc., in the experi-
ments so far carried on, indicate generally that there exists little
difference in the height of the hypocotyls or stems of the normals,
and those in contact with the different-sized particles of soil constit-
uents. The growth in length of the primary root is lessened and the
secondary root system is greatly increased as we approach the finer
grades of contact material. In some species with relatively large
roots the coarser particles, namely 16-8 mm., appear occasionally to
stimulate primary organs, but this reaction is variable, and associated
with growth correlations as manifested in the more or less increased
development of the secondary root system. The reaction of roots
to different-sized particles will undoubtedly be found quite variable.
Species with relatively large and strong primary roots such as the horse
bean, peas, etc., react differently from seedlings with a delicate root
system, such as the mustard and turnip. The roots of the latter
species, even when grown in contact with excelsior or soil particles,
produce a marked secondary root system characterized by little or
TABLE I9
Showing Growth of Hypocotyls and Roots of Mustard (Brassica alba Boiss.) in Contact
with Different-sized Particles of Sand and Gravel in Water9
Average Length of Hypocotyls and
, Primary Roots (Centimeters)
Size of Particles
(Millimeters) Hypocotyls Roots
Worntals. ao Wetter eae 8.7 1207,
ROrSS velar ee bee Lk eee 8.2 12.6
cop hs NAR eRe oes Pome, Clacc'e-p on ier 8.7
AT AE eat SOY ty EN ere Oro 11.5 7.6
Poe tated Hite Ae RR EA LE ae bs tye 11.9 4.4
ORG Gam ERY aS og Se ao olot 11.6 4.3
OPS O25 ane ceo ee 11.2 4.2
O:2'5=Onli ras Se aaah eee 03:7 3.4
no primary root development. The same tendency exists in species
characterized by larger roots to increase their secondary root system.
On the other hand, the primary root system is greatly accelerated by
contact with excelsior in species with large roots. (Cf. Tables 16-17
and 18.) The experiments with mustard given in Table 19 show
19 15 plants used in each test.
STONE: CONTACT STIMULATION 473
much difference in the growth of hypocotyls and roots in the different
grades. This feature is associated with the greatly accelerated
development of the secondary root system, and extending from the
coarser grades to the finer ones. Other than the production of
secondary roots in the plants in the 16-8 mm. grades there was little
difference between the growth of the latter and those of the normal.
There were no secondary roots in the normal grades or water cul-
ture plants in this case, although they were fairly well established in
the 16-8 mm. grade, from which grade the increase in numbers and
total length of the secondary roots were quite noticeable. The
average development of the primary and secondary organs as well as
the surface area of the same was greater in the contact plants than in
the normals. Soil particles and excelsior submerged in water have a
similar stimulating effect on mustard as will be seen by comparing
Tables 18 and 19.
THEORETICAL CONSIDERATIONS
While the general tendency of plants and plant organs is to avoid
contact with one another, the histological units or cells which com-
posed the individual are in contact with one another, and the same
holds true to a certain extent with different organs when in the em-
bryonic or bud stage. It is the exception rather than the rule to find
the various members or organs of different plants, or even those of
the same plant such as roots, branches, leaves, etc., in contact with
one another, or in other words it appears to be a universal law in
nature that the various organs of plants occupy space by themselves.
Uniformity and regularity in the arrangement of cells and organs is
more common to the lower than to higher organisms, since in the
higher organisms this feature is sacrificed to some extent by biological
necessity and adaptation. Primarily the arrangements of organs in
plants or angles of divergence are determined by laws which are
common to gravitational and electro-magnetic phenomena, and the
‘ arrangement of the various organs of plants appears to be determined
by the action of these forces upon their ultimate structural units,
molecules, micellae, atoms, electrons, or whatever they may be.
The angles which various organs assume in plants closely resemble
those which are illustrated in the formation of certain types of crystals,
and the behavior of iron filings under the influence of a magnet.
Plants are susceptible to all of the common environmental influences
which surround them, but the modus operandi of these various external
agencies on protoplasm is little known and especially concerning the
mechanism and nature of conductivity of impulses. The reaction to
contact results from a mechanical impulse, inasmuch as when the
474 BROOKLYN BOTANIC GARDEN MEMOIRS
external cells of the organism come in contact with solid particles a
reaction follows. To affirm, however, that the stimulus is mechanical
in nature does not explain anything, because we know nothing of
the nature of the so-called mechanical impulse. The bombardment
of organisms with electric waves (negative electrotropic response)
may be of the nature of a mechanical impulse and such may hold
true for other forms of radiant energy. Phototropic, thermotropic,
electrotropic and contact stimuli may not materially differ from one
another in the nature of their action on organisms, that is, in the sense
of acting ina mechanical manner on plants. The stimulation resulting
from such contact is apparently transmitted to the living zones which
induce definite adjustments or coordinations of the vital processes,
the nature of the response being determined by the nature of the
stimulus involved and the organs stimulated. The external cells
of a leaf, for example, coming in contact with an object would probably
react to the stimulus through the cuticle and exterior cell walls. The
reaction of plants to contact is probably one of the most primitive
forms of responses and quite universal in the vegetable kingdom.??
The nature of the response to contact resembles mostly that of touch
or at least a primitive and rudimentary form of this sense.
There are several types of contact stimulation that have long been
recognized such as occur in the response of tendrils, tentacles, stamens,
etc., when brought in contact with different substances. These
reactions are associated with different types of irritability. Most of
the known reactions to contact such as are illustrated by tendrils,
etc., are closely associated with biological adaptations, and as such
they have been subject to considerable modification. It is not at
all improbable, however, that these various types of reactions are
modifications and differentiations of a more simple and_ universal
form of contact response. The various forms of response movements
which are associated with irritability have been classified under the
so-called tropistic, nastic, and taxic reactions. The first type of
response is characterized by the movement of organs toward a definite
position in response to a stimulus, while nastic reactions are those
which are independent of the direction of the stimulus. Taxic reac-
tions are those shown by motile organisms. ‘There are several different
types or manifestations of irritable phenomenona such as_ photo-
20 Some experiments made with mucor in Petri dishes containing beef extract
and fine particles of quartz sand showed contact responses. Both series of Petri
dishes contained the same amount of beef extract, but in one the fine sand was added
for the purpose of obtaining contact with the mycelium. The sporangia were black
and mature in the Petri dishes without sand, while in the dishes containing the fine
sand, the aerial hyphae were much more elongated and the sporangia light in color
and less mature.
STONE: CONTACT STIMULATION 475
tropism, geotropism, etc., which are characterized by both positive
and negative reactions, but for our purpose it is necessary to consider
only a few which are somewhat similar in character, and which have
not unlikely originated from a simple fundamental type.
(a) Contact Stimulation
Probably a universal type and one of the most fundamental forms
of response in plants, which is characterized by a stimulated growth of
various organs due to contact of one organ with another, or with
various substances. The extent of the reaction is determined by the
degree of contact, and probably influenced by the nature of the
contact substance.
(b) Haptotropism
Illustrated by various tendrils reactions; tentacles of Drosera.
(c) Haptonastie
Illustrated by certain types of contact of some tendrils and tentacles
of Drosera.”
(d) Setsmonastie
Illustrated by movement of stamens of Centaurea, Berberis, and
Mimosa leaves.
(e) Thigmotaxts
Reaction of motile organisms (Chlamydomonas) resulting from
contact of cilia with different substances.
(f) Wound Stimuli and Reactions
Wound stimuli in general caused by mutilation, presence of organ-
isms, and direct contact of plant members with one another induce
various reactions, the nature of which depends upon the host plant,
and nature of the cause responsible for the same. The general
response to wound stimulation is manifested by the production of
abnormal growth due to active cell division, and in some cases enlarge-
ment of the cells. These reactions are characterized by local effects
and the reactions are often disproportionate to the stimulus.
In some types of reactions there would seem to be involved more
than one class of phenomena. For example, the haptotropistic reac-
tions of tendrils, or at least the subsequent effects involved in the
formation of the spiral, etc., by the stimulus, and which is manifested
*1 Stark (3) has observed that many plants and some roots react to contact
stimulation similar to tendrils when rubbed with a stick or cork.
476 BROOKLYN BOTANIC GARDEN MEMOIRS
by the production of mechanical tissue would seem to be similar to
that induced by other irritable phenomenona. In the formation of
the spirals in tendrils there is considerable stress to be overcome and
the constantly increasing weight of the various members of the plant,
such as is associated with the development of foliage, fruit, etc.,
together with the effects of rain, swaying movements due to wind, is
most admirably met by the production of mechanical tissues. These
additional strains are taken care of by the same type of response as that
induced by the stretching of plants with weights. Since the leaves of
plants appear to be quite sensitive to contact it is readily conceivable
how the more specialized haptotropistic responses to contact exempli-
fied by tendrils could readily be evolved from simple contact irritability.
There are, however, a large class including various types of reaction
phenomena which do not fall under tropistic, nastic and taxic responses.
These are the so-called wound reactions—traumatotaxis (reaction of
cell nucleus). Traumatotropisms, illustrated by decapitated root, and
positive galvanic currents on roots, are apparently direct responses to
wound stimuli, as probably are the reactions first observed by Darwin
resulting from the attachment of different substances to the root tip.
In wound reactions, however, contact is involved to a greater or less
extent, and more or less injury and abnormal conditions are associated
with thés class of phenomena. Most of the responses following wounds
are local in their effect, although the organism as a whole may be
affected even from relative insignificant mechanical injuries, as shown
by the modification in the developments and functions of the several
organs in mutilated plants.
Representative types of this class are seen in the various accelerated
growths produced by insects, fungi, bacteria, mechanical injuries, etc.,
and generally the reaction continues long after the primary or excitory
cause has disappeared. Local accelerated growths, however, do
not always follow as a result of the intrusion of pathogenic organism
inasmuch as the nature of the response is determined to a large extent
by the character of the tissue affected. For example, eel-worm
infestation of roots may give rise to galls, whereas on stems such
a response may not necessarily follow and even on roots these reactions
differ. In many so-called wound reactions the degree of response is
disproportionate to the stimulus responsible for the same. This is
illustrated by feeble lightning discharges on trees, in which case the
stimulus (lightning causing burning) lasts but a few thousandths of a
second. The reaction, however, to such insignificant injury—often
hardly perceptible and characterized by the destruction of a few
cambium cells, may manifest itself for years in an accelerated growth
of the annular rings adjacent to the injury. The flow of tissue in
STONE: CONTACT STIMULATION 477
sucker growth around stumps, the enclosure of twigs, grass, etc., by
mushrooms, and the imbedding of tree guards illustrate stimulated
growth responses not generally associated with wounding. When
more or less long stubs remain following the destruction of limbs on
trees, they act as a stimulus, and large masses of tissue ultimately
accumulate around the base of the same. If, however, the stub is
removed close to the tree, healing follows, and when the callus unites
growth becomes normal. Again the feathery growths, consisting of
small twigs which are common on the trunks of elm trees, give rise to
the same type of reactions, namely, the formation of excrescence or
tuberous growths at their point of junction to the tree. The presence
of blocks of wood on trees to prevent the contact of guy wires with
the bark often stimulates the flow of tissue, and while there is appar-
ently no injury associated with this connection there exists a more
or less severe tension or pressure on the bark. The tendency of
tissue to grow over even loosely attached tree guards and wire attach-
ments, even when it would require but little force to dislocate them,
is universal. Signboards placed on trees, although often insecurely
attached, are sometimes entirely obliterated by a greatly stimulated
callous growth, and tree guards loosely attached to trees often become
imbedded in the tissues, when it would require but little exertion on
the part of the underlying tissue to dispose of the same. It may be
assumed, however, that the presence of nails driven into the wood
for the purpose of holding the sign in place would act as a stimulus,
but the same reaction follows if the nails are not present and the sign
merely held securely by means of wires. Moreover, the growth
response here conforms to the sign as a whole rather than to any par-
ticular point of attachment by the use of nails, etc. Similar response
occurs in the growing of roots around large masses of gravel in the
soil or when limbs or roots come in contact with one another, in which
case natural grafts may occur. The tissue, however, exterior to that
which is in contact is often destroyed in natural grafts, etc., which
would have the same effect as wounds; although even here the growth
stimulation is much greater than that resulting from mere wounding.
In this type of response there are two factors involved, namely con-
tact and growth acceleration. The reaction, however, is local and does
not differ materially whether injuries occur or not, as is shown by
growth stimulations arising from other causes, such as frost cracks,
etc. There are other similar types of reactions apparently differing
somewhat in the nature of the response to contact from those enumer-
ated, namely, those arising from restricted growth. Contact in such
cases occurs, but injury may be absent, although abnormal growth may
result. This type seems to be characterized in their manner of re-
sponding by a greatly increased osmotic tension of the cells.
32
478 BROOKLYN BOTANIC GARDEN MEMOIRS
When roots are growing between large boulders and are restricted
in their development, the flow of tissue is such that the root may
become enormously flattened. The reaction in such cases is similar
to that produced by a wound, although evidence of wounding may be
entirely absent. Such roots, however, often lift enormous weights
which would indicate that considerable osmotic tension exists in the
cells. When rapidly developing organs, such as a squash, for example,
is placed in a harness and subject to weight, it will assume a much
distorted shape which illustrates stimulated growth phenomena.
Again, when ferns and even delicate mushrooms push through con-
crete they show little or no evidence of wounding, although the reac-
tions in such cases are characterized by a large increase in the osmotic
tension of the cell, sometimes equaling 50 atmospheres (4). The
experiments of Pfeffer (5), in enclosing roots in plaster casts, thus
restricting growth and greatly increasing the osmotic tension of the
cells, are typical of this class of responses. To what extent, if any,
cell fusion of sexual elements, development of attachment organs in
fungi and algae, and outgrowths in spirogyra filaments when in con-
tact with certain crystals, are related to contact, is problematical since
in some instances chemotropic phenomena would have to be carefully
differentiated from any other which might prevail in interpreting such
phenomena. Also to what extent contact stimulation may influence
cell enlargement and cell division resulting from the intrusion of
foreign elements would be merely a matter of speculation at the
present time. There are, however, numerous instances of cell enlarge-
ment associated with crystals (raphides), pathogenic and non-patho-
genic organisms, which resemble contact stimulation, although the
recent important contributions on this subject by Dr. E. F. Smith (6)
would indicate, in some cases at least, that cell responses to pathogenic
organisms are associated with chemical or physical phenomena. Some
of the responses of plants associated with pathological phenomena are
not characterized by cell stimulation, but with color reactions due
apparently to excretions from organisms. Intense color reactions
are also associated with marked cell proliferation caused by chemical
substances absorbed by plants from the soil, as shown by the reactions
of Platanus Orientalis to the toxic properties contained in illuminating
gas. These reactions, however, as are similar ones in poplars and wil-
lows, which develop large masses of parenchyma under the bark and
cause rupture of the same, are associated with two factors, namely,
the direct effect of the toxic substances on the cambium inducing
rapid cell division, and decrease in the tissue tensions of the cortex
following the collapse of the same by poisons. Various chemical
substances (banding substances) applied to trees produce local growth
STONE: CONTACT STIMULATION 479
acceleration, but these apparently affect the outer tissues, and to a
certain extent the underlying vital layer. Dr. H. von Schrenk (7)
and Dr. E. F. Smith (6) have shown that ammonia compounds cause
intumescences in cauliflower. The absorption of the chemical sub-
stances by the roots in some cases produces a rupturing of the epidermal
tissue which is followed by cork formation, and in cases of malnutri-
tion excrescences are often formed on the fruit. Finally, contact
stimulation has an important bearing on experimental work, especially
with that done with plant food, fertilizers or soils, inasmuch as stakes
and wires are often employed as support. Any differences existing in
the contact of the plants, either by the use of supports or that occa-
sioned by the proximity of plants to one another, thus causing contact,
would be effective in modifying the results. The relative height and
development of two plants which would affect the contact surface
would also be important factors in experiments of this nature. Indeed
in this respect we have noticed on more than one occasion experiments
where differences existing in the degree of contact would account for
all the variations in the growth of the plants that were supposed to
be due to other causes rather than to the particular treatment which
they received.
LITERATURE CITED
1. Hall, A. D., Brenchley, W. E., and Underwood, S. M. The Soil Solution and the
Mineral Constitutions of the Soil. The Jour. of Agr. Sci. 6: 278-301. Sept.
IgI4.
2. True, R. H., and Oglevee, C. S. The Effects of the Presence of Insoluble Sub-
stances on the Toxic Action of Poisons. Bot. Gaz. 39: I-21. 1905.
3. Stark, Peter. Ber. d. d. Bot. Ges. 338: 380.
4. Stone, George E. The Power of Growth Exhibited by Ferns. Bull. Torrey
Bot. Club 36: 221-225. May 1909.
5. Pfeffer, W. Druck und Arbeitsleistung durch Wachsendepflanzen, p. 93.
6. Smith, E. F. Mechanism of Tumor Growth in Crowngall. Jour. of Agr. Res. 8:
Jan. 1917.
7. Schrenk, H. von. Intumescences Found as a Result of Chemical Stimulation.
Ann. Rept. Mo. Bot. Gard. 16: 125-148. 1905.
DUPLICATION AND COHESION IN THE MAIN AXIS
IN CICHORIUM INTYBUS
NB. SLOUmr
New York Botanical Garden
The terms duplication and cohesion may be used to designate a
very special type of fasciation which the writer has observed in the
variety of chicory cultivated under the name “red-leaved Treviso.”
The most distinctive characteristic of this type of fasciation is the
duplex nature of the main axis. From a single root a double
stem arises with the two parts, as a rule, strongly united. The
duplex nature is indicated by a pair of grooves which extend in the
direction of the long axis and round out the two stem-elements. This
is quite well shown in number 5 of the accompanying plate XII.
This type of fasciation differs from the banded and cone types
most usually seen in that here two stem-elements of equal size and
rank are clearly in evidence throughout a segment of unequal
diameters, giving a symmetry that is bilateral. Also the fasciation
is confined to the middle and lower portions of the stem and decreases
upwards, the main axis often becoming at its apex quite symmetri-
cally simplex.
I have been unable to find in the literature any reference definitely
mentioning fasciation of this particular type. It appears not to have
been noted and described even in this rather well-known variety of
chicory.
In the degree and the extent of the duplication there is much
variation. The most extreme condition of duplication is seen in the
complete separation of the two stem-elements with each perfectly
formed and without lesions, as is shown in no. 7 of Plate XII. In many
cases the two elements are indicated only by grooves which extend
from close to the base of the plant to a height of about three feet.
The length of the grooves and the corresponding segment of evident
duplication may, however, be reduced to mere traces, as is shown in
no. 3 at a or in no. 4 from a to J, as indicated. In a few plants of
this strain there is no evidence of duplication and the phyllotaxy is
of a single and regular spiral only.
Each stem-element has its own phyllotaxy in so far as this can be
expressed. In the most pronounced duplex condition there are clearly”
two separate spirals in evidence. The phyllotaxy of one is not a
480
STOUT: DUPLICATION AND COHESION ~ 481
continuation of the other and more branches and leaves are produced
upon the two than upon a single stem. Branches and leaves occa-
sionally develop, however, from the very center of the grooves and the
bases of leaves thus placed often extend across a side of both stem-
elements. Such leaves and branches are always simple as far as I
have observed. In the uppermost portions of the stem the phyllo-
taxy is often completely regular, but immediately above the seg-
ment of noticeable duplication the phyllotaxy is irregular and this
irregularity may extend to the apex or terminate in the banded type
of fasciation.
Torsion is frequently seen. The two elements may twist about
one another in spirals with the two parts equal in development and
the spirals in the same direction. In other cases the two stem-ele-
ments exhibit incompatible growth either as to direction of torsion
or rate of growth, or both, and mechanical lesions result. These tear
apart the two elements or often cause one to break, thus producing
much irregular and twisted development of the main axis. Various
conditions which thus arise are shown in Plate XII. Usually the lateral
branches above such lesions are poorly developed (no. 9), the more
vigorous growth being seen in the lower laterals.
In this strain of red-leaved Treviso, irregularities of development
are seen in the growth of the cotyledons. For most seedlings two
quite normal and separate cotyledons are produced as shown in nos.
1, 2, and 4of Text Fig.1. In numerous cases even of sister plants the
two cotyledons are fused; only the basal portions of the stems may be
fused as in nos. 5, 6 and 15, or the fusion may extend upward
toward the apex as shown in nos. 7, 8 and g. In some cases
what appears to be only one cotyledon is in evidence, as shown
in nos. 10, II, 12 and 16; either the two have become completely
fused or the growing point of one failed to develop. The very
complete series of stages of fusion leading to a single cotyledon
suggests that the initial growing points of the two cotyledons may
completely fuse or perhaps are never differentiated. There are also
numerous cases of crumpled and irregularly developed cotyledons
either in one or both of a pair or in a single one, as may be seen in
ee. 3, 13, 14, 15 and’ 17.
In only two cases have I observed any tendency to an increase of
cotyledons; in each of these seedlings one of the cotyledons was some-
what bisected, as shown in nos. 19 and 20. The duplication seen in
the main axis is not accompanied by duplication or increase in the
normal number of cotyledons.
In a very few instances there is no development of the plumule.
This may occur in seedlings having two normal unfused cotyledons as
482 BROOKLYN BOTANIC GARDEN MEMOIRS
shown in no. 4, as well as in seedlings having only one cotyledon, as
showninnos.16and17. Such plants usually diesoon. In some cases,
however, adventitious buds arise, but as far as observed these make
only a feeble growth. The drawings 16) and 17) show such feeble
adventitious growth of seedlings nearly three months old, at which
time many sister plants had rosettes measuring ten inches in diameter.
WO NIP BoP YW
=f ‘an4
Po Le PY
13 14
Vat Be
Fic. 1. Young seedlings of the variety red-leaved Treviso. Scale about one-
half natural size.
1 and 2. Two normal separated cotyledons with plumule.
3. Two cotyledons, both somewhat crumpled.
Two cotyledons; no plumule.
Cohesion at base of cotyledons only.
Cohesion at base of cotyledons; one crumpled.
7, 8,9. Decided cohesion.
10, II, 12. Only one cotyledon present.
13. Two cotyledons; separation involved some lesion.
14. Much crumpled and poorly developed cotyledons.
15. Some cohesion at base.
16a. Single cotyledon; no plumule. 6. Same seedling two months later with
adventitious bud.
17a. Single much crumpled cotyledon with seed coat attached at apex; no
plumule. 6. Same seedling two months later.
18. Much fusion of cotyledons.
19, 20. Decided lobing of one cotyledon.
15
Qy Cn
The growth interrelations of reduction or fusion of cotyledons with
duplication in the main axis are not clear. In the former there is
a fusion or reduction in the number of organs normally formed; in
the latter there is a tendency to the production of two main stems
STOUT: DUPLICATION AND COHESION 483
instead of one, accompanied by their cohesion or incomplete separation.
In both the factor of cohesion is present, but in the cotyledons it leads
to reduction in the number normal to the species, while in the stem it
tends to reduce the duplication to the single stem-element that is
normal to the species.
The occurrence of the sort of duplication described above has now
been observed in four generations. Thirty-five plants of the 1913
crop of the variety red-leaved Treviso were grown from commercial
seed. In 14 plants the main axes were decidedly duplex, the length
of the segments showing duplication ranging from a few inches to
about three feet. The stems of the other 21 plants showed no sign
of duplication. All plants of this generation which were tested were
found self-sterile from physiological incompatibility, so it was not
possible to obtain self-fertilized seed. Numerous crosses were at-
tempted between plants, but only one cross attempted was compatible.
In 1914, 12 plants were grown as progeny of the cross between two
plants which were quite alike in exhibiting duplication of rather inter-
mediate development. In all 12, duplication developed. The vari-
ability in degree of duplication was marked and ranged from very
slight indications to very decided cases of torsion and lesion. One
plant developed too late in the season to be tested for self-fertility,
but the other 11 were all self-sterile. However, some compatible
crosses were made from which 43 plants were grown in IQI5.
Of the 1915 crop, 39 of the 43 plants exhibited duplication to some
degree. In one plant the two elements were completely separate from
the very base upward (no. 7). In several plants torsions and lesions
were strongly developed. Four plants were apparently not in the
least fasciated and in each of these the phyllotaxy was regular and
single.
In the 1916 crop, 150 plants were grown. Two of these were froma
self-fertile plant (the only one of the 1915 crop that was found to be self-
compatible in any degree). The 148 other plants of this generation
were from six different crosses involving eight different parent plants, all
of which, however, exhibited duplication in some degree. This genera-
tion was descended from three generations of parentage that exhibited
duplication. There was duplication to some extent at least in 144 of
these plants. Sixappeared to have a single stem-element with regular
phyllotaxy; these six plants were distributed among the offspring of
three different crosses.
One plant of the 1916 crop exhibited a noticeable fasciation of the
ribbon type in the upper branches in addition to duplication and
cohesion in the main axis. The tips of the main branch and of various
laterals were broadly flattened and the branching was reduced so that
484 BROOKLYN BOTANIC GARDEN MEMOIRS
flowers and short spur branches were clustered at the extreme ends,
giving a peculiar rosette-like appearance. The two types of fasciation |
were quite distinct on the plant, both as to general appearance and
location.
From the performance of these pedigreed cultures, it seems clear
that the character of duplication and cohesion persists in successive
generations of this variety of chicory to such an extent as to appear
strongly heritable. It is not completely so, for a few normal single-
stemmed plants do occur quite irregularly in various generations and
lines of descent.
The heredity of the character of duplication has also been tested by
crossing plants of the I915 crop showing typical fasciation with a
plant of wild stock (plant A) which had a short, slender, main stem.
No tendency to duplication has been seen in plants of wild stock which
have been grown or in plants of various generations derived by crossing
the wild plant A with plants of the cultivated variety known as Barbe
de Capucin. This cross here in question involved, therefore, on one
side parents with duplication, and on the other a plant of a stock
free from duplication.
Thus far 81 plants of an F, generation have been grown; nineteen
of these had the wild plant A as a seed parent. Of these 81 plants
only three possessed the grooves which are seen in most typical cases
of duplication of the main axis. In only one of these were the grooves
pronounced (see no. 10); in the other two there were only slight indi-
cations of grooves (see no. 11). In 78 plants there was not the slightest
indication by grooves of any duplication. However, in tracing the
phyllotaxy from base upward, irregularities were seen in 48 plants.
Two leaves or branches were often opposite or the direction of the
spiral would appear to shift from left to right or vice versa. In 30
plants the spiral of the phyllotaxy proceeded very regularly from base
to tip in a way that indicated a normal single stem-element (see no.
12). Of the 19 plants having the wild plant for a seed parent, 7 were
quite normal and in 12 there was a mixed or irregular phyllotaxy.
Judged by performance in the F, generation, the character of
duplication is only incompletely and partially dominant. An inter-
mediate type is frequent in which the only suggestion of a duplex con-
dition of the main axis is seen in an irregular phyllotaxy.
I cannot at the present time contribute any information regarding
the sources, causes, or nature of the stimulus operating in duplication
nor any definite facts regarding the attending anatomical development.
When this paper was presented it was suggested by Dr. Erwin F. Smith
that possibly infection by some organism, bacterial or otherwise, was
necessary to the development of duplication as here observed. If this
AUNOOIHD NI NOILVIOSVYY *LNOLS
“WX 3LW1d 4 AWNIOA “SHIOWSIA) NAGYVS OINVLOG NA1IMOOYUG
STOUT: DUPLICATION AND COHESION 485
should be the case it is evident that while the susceptibility is very
decidedly limited to the variety it can be transmitted directly or
indirectly though incompletely to te numbers of the offspring of a
hybrid generation.
SUMMARY
An unusual type of fasciation occurs in the variety of chicory known
as red-leaved Treviso. It consists of a very decided duplication in
the main axis of the stem, giving two stem-elements with, however, a
decided cohesion of the two. In this variety of chicory there are also
various irregularities in the development of cotyledons and plumule.
All degrees of fusion between the two cotyledons are in evidence; in
some seedlings only one cotyledon is present. Occasionally no plu-
mule develops.
The character of duplication and cohesion of the main axis is
strongly but not completely heritable. There is wide variation in
the degree of duplication and a few plants with a normal unduplicated
main axis occur.
In an F, hybrid generation of crosses between plants with dupli-
cation and plants of wild stock which exhibit no tendency to such
fasciation the character of duplication is incompletely dominant both
as to degree of expression and number of plants affected. An inter-
mediate type is strongly in evidence in which the only indication of
duplication is seen in a mixed and irregular phyllotaxy.
EXPLANATION OF PLATE XII
Nos. I-9 inclusive are of stems of plants of the variety red-leaved Treviso;
no. 7 is of the 1915 crop, all others are of the 1916 crop.
No. 1. Stem showing no duplication. Phyllotaxy regular.
No. 2. No duplication but stem thicker than in no. 1. Phyllotaxy regular.
No. 3. Stem of small plant. Short segment of duplication at a.
No. 4. Duplication from a to b; phyllotaxy above a is irregular with tendency
for branches to be paired.
No. 5. Very decided duplication from a to b. Lesion separates the two stem-
elements in lower center. Stem-elements parallel below but twisted above.
No. 6. Decided duplication with parts much twisted.
No. 7. The two stem-elements separate from the base. No lesions.
No. 8. Longitudinal lesions strongly developed. One stem-element much
contorted.
No. 9. Much torsion with transverse lesion of one stem-element.
Nos. 10, 11 and 12 are F; hybrids of red-leaved Treviso X unfasciated plant of
a wild variety.
No. to. Duplication evident in lower two thirds of stem.
No. 11. Duplication indicated by a very slight but broad groove near base.
Phyllotaxy mixed and irregular especially near base.
No. 12. No duplication. Phyllotaxy regular.
A QUANTITATIVE STUDY OF RAUNKIAER’S GROWTH-
FORMS AS ILLUSTRATED BY THE 400 COM-
MONEST SPECIES OF LONG ISLAND, N. Y.
NORMAN TAYLOR
Brooklyn Botanic Garden
The value of sorting species of plants into different categories,
based on their growth-forms, has been pointed out so often that
there scarcely seems further need of going over the subject again.
The weakness of such a sorting and the percentages based on it, due
to the fact that species, not individual plants, are considered, is ob-
vious. Such percentages as have been published show not so much
an actual response to climatic factors, as they do the multiplicity of
forms that may have been developed. For most regions that is all
that can be done, as anything like a plant census of a given region is
usually impossible. Yet upon such a census, or some approximation
to it, there could be based percentages of different growth-forms that
reflect more accurately than any species percentage the actual climatic
response of vegetation to climate.
The importance of getting, if possible, some growth-form per-
centages that should be quantitative rather than those based on
species only resulted in a study of the flora of Long Island, N. Y.,
with this in view. The island is roughly 120 miles long and 12-16
miles wide and, excluding ferns and their allies, has about 1120 species
of native plants. It is diversified as to vegetation, as there are good-
sized areas of “scrub,” mostly oak and Ericaceae, considerable de-
ciduous forest, some extensive “pine-barrens,’’ salt marshes, a small
prairie, and the downs at Montauk and Shinnecock.
In a general study of the flora and vegetation of the island, distri-
bution maps for each of the riative species were made and have been
posted up for several years. Such maps indicate actual collections
represented by specimens in herbaria, field notes by the writer, all
published records of species and descriptions of different vegetative
areas by nearly all who have written about Long Island for the last
250 years. From Daniel Denton’s History of New York, through
the period when numerous Quaker journals were issued, down to the
modern observations of professional botanists, these records have
been accumulated. The opportunity, therefore, of getting something
486
TAYLOR: RAUNKIAER’S GROWTH-FORMS 487
like a comprehensive view of the flora and vegetation of the island is
excellent.
Within the last few months it has been possible to separate the
distribution maps of the species into two groups. The one which
interests us just now is the smaller, consisting of the 400 commonest
species. These make up the great bulk of the vegetation of the island,
the other group of about 719 species being scattered and nothing like
so common.
When these four hundred species are sorted into the different
growth-form categories of Raunkiaer and their percentages reckoned,
we find them grouped as follows :!
Numbers of Percentages of the
Growth Form Species Commonest Species
INI GAC abe tree oe a tye 6 1.50
VIS epee Orotte mate a ve as ree 12 3.00
MIG aoe ait cea nae Shr 34 8.50
Ie eee ete, SR aa nn 17 4.25
(CR eke ae oe re eae 29 7.25
1, Ie adie Sot Ne din Pai dincreit heat os oh 120 30.00
Ge er eee, ee, 5 aes: 84 21.00
Lill Lee eel oe BRE eer 5) 6.75
dE ce og chor eee ide eee ote 57 14.25
Stem succulents... 15/15 aoe 8 2.00
Parasites. pietticets ae oe 6 1.50
The amount of deviation from the normal spectrum of Raunkiaer
or from the growth-form percentages of the total flora of Long Island,
or from the percentages of the local flora area? should show an inter-
esting relation. The figures are as follows:
| MG | Ms | MC | N | CH H G He |r
a — —} — |
Normal spectrum.......... 6 | 17 | 20 | 9 | 27 3 I 13
WMOCAIMOLA se sesenmeee ele sc 14,704 12,588 86
Nigicmany (CBINARE) Bee apie co NOren eee dete aie Cte eee Mech sic 857 741 86
Meeseomnde (SLELANSSOM) fi cis cok, oe tee ss odes eseeiee 221 200 90
Biligcimenclionda(Simmons)its . sneer. coe ar ae 76 71 93
BARGE SA NV ALINIng et ale )ie on. 64 chee teil maa eaesrs 164 150 gI
SrereLTIM (nize A ceo hy sic Ms ic, Gaatove ise ana roa Maun Ree arava > 4,481 3,554 79
CORE CCUINLIT) en cnareer oy ctisves ote ese z crore RBS neta met eae 1,461 1,161 79
Svcalhy (CUK@ yet] 0%=50(2) a eae an ee 1,697 1,295 76
men iM OSE) Ped chpes dhe praiesim ite denen ate eats Obs 2,949 2,477 84
MlonanOnientalis (B01SSIEM))... acs. 06 ee os ee wee es 9,771 8,110 83
MePePMR TESTICLE). 6 oss cele wa ea ga due ele eines Sea 7/ 1,861 57
iciegzalll (AWG A Rp en Aree el aPRe Met pater Mien fh 15,981 H,092)| "26
Dio pAmazon Valley: only). 00650576822! bo 33 ane 2,209 265 12
Brarshevvest, Indies, (Grisebach)). . 50924. .4.5.4-2 2% 2,249 675 30
Tropical Africa (Oliver and Thiselton-Dyer)........ 8,577 35001 | Ae
Emimcuietnciat (HOOKEer)) eo nat «sec Pte n ee eee 10,454 4,344 42
Bombay. (Cooke), Lowland only... ..........4:25- lp at,249 487... 39
Wmaper Gangetic Plain (Duthie), 3. 1: 62.425 act. sie | 1,084 583 54
Mera OTM MIEN): 5. i isn des share aciae apne ae oe T7080 Glos | a7
eprom (ioorders) ss kh, SRN. SPE ane eB ee 3,188 867 27
Dnrenenast ndies:(Miduel)). ct) eases esee asc ade 6,398 | 1,599 2:
Maier remus (IIe) ois tie wip tard mpegs andres tea antes 252 553 17
meen (BENCHAM) o.oo vist ota v kjake its sole Wee we 728 293 40
vs anil ees (GUI eri BD AR hae eaten ifcmeec nesters iSteicacenlenae ates tie 333 106 32
When it is remembered that these figures do not include mono-
cotyledons they add additional weight to those given for the local
flora area, the total Long Island Flora, and for the 400 commonest
species on the island. Summarized these figures show the following
percentages of herbs:
490 BROOKLYN BOTANIC GARDEN MEMOIRS
PERCENTAGES OF HERBS
1. Normal spectrum, including all categories......... 48%
2. Average of 15 North Temperate floras as listed
above, excluding monocotyledons............... 82% or about 90% counting
monocotyledons.
3. Average of 13 Tropical floras as listed above,
excluding “monocotyledons: . 23s. on ne oe 31% or about 38% counting
monocotyledons.
4. Australia’ (excluding monocotyledons)............ 30% or about 35% counting
monocotyledons.
5. New Zealand’ (excluding monocotyledons)......... 55% or about 70% counting
monocotyledons.
G:pleocalsioracarea,allicateconiess sears eee eer ae 79%
7. Total Long Island flora, all categories............. 83%
8. 400 commonest Long Island species, all categories .78%
Items 2, 3, 4 and 5 do not count monocotyledons and a careful esti-
mate of these monocotyledons shows that they make up form 1/5-1/3
of the floras of the regions mentioned. This monocotyledonous ele-
ment is overwhelmingly herbaceous and it adds a great deal to the
percentages of herbs in these items. Upon these figures the per-
centages, counting the monocotyledons, which are of course included
in Raunkiaer’s normal spectrum, are estimated as shown in the table
above. The average of the percentages, including monocotyledons,
in items 2, 3, 4 and 5 is 58 percent which is as near an estimate to
the relation between herbaceous and woody species as we can get for
the areas mentioned. These, with the exception of South America,
make up the great bulk of the flora of the world and 58 percent of
herbs as against 42 percent of woody plants can, for the present, be
set down as a fair estimate. This is a clear 10 percent above the
combined herbaceous percentages of Raunkiaer’s normal spectrum,
and very much nearer the percentages of the northern regions gen-
erally, where herbs predominate.
In other words, the evidence from large areas, based on species,
and from a small area like Long Island, based on frequency of indi-
viduals, points unmistakably to the necessity of shifting some of the
Raunkiaer growth-form percentages. Herbs make up the great
majority of the vegetation in North Temperate regions, and, as we
have seen, even in the tropics and southern hemisphere their bulk is
by no means insignificant. Yet in the face of these figures, and of
those of as complete a plant census of Long Island as we can get, the
total herbaceous element according to Raunkiaer should be only 48
percent. As we have seen this is much too low for anything like a
true vision of the relation between herbs and woody plants in the
whole North Temperate region, and it is 10 percent lower than the
SANT OtGih tC. sa 5S3) boos
TAYLOR: RAUNKIAER’S GROWTH-FORMS 491
figure for nearly the whole world. This is the chief point made here.
For, if the percentages of woody and herbaceous species, as shown by
Sinnott and Bailey, and percentages based on an approximation to a
plant census as shown by the figures from Long Island are not in
substantial agreement with Raunkiaer’s system, then it follows that
that system does not, as yet, show what has been claimed for it. Just
what reshifting of the percentages in the normal spectrum is necessary
lies outside the scope of this paper. It seems evident, however, that
they need more study and over large areas, preferably based on plant
censuses such as has been attempted on Long Island. With the com-
pletion of such studies we might have in the revised percentages more
accurate data as to climatic response than is possible at present.
THE ANCIENT OAKS OF AMERICA
WILLIAM TRELEASE
University of Iilinots
While studying the oaks which are now so striking a component of
the vegetation of North America, I have found it necessary to form
some idea of the history of Quercus before our own day. Neither time
nor opportunity has offered for basing this on a reexamination of the
scattered materials that have served for the classic studies of Les-
quereux and Newberry, or for the later work of Hollick, Knowlton,
Berry and Cockerell; but from a careful examination of descriptions
and illustrations I have tried to bring into some sort of orderly as-
semblage the scattered facts that have been observed and described (1).
So far as I know, only two of our fossil species (Q. consimilis and
Q. paucidentata, both of the Eocene) are known in fruit (2); the others,
though exceptionally with twig remnants, are represented usually by
dissociated leaves, sometimes well preserved but frequently only in
fragments showing little detail. These materials have been referred
to Quercus because of the general appearance and especially the
venation of the leaves when this is ascertainable. It is not sur-
prising that misapprehension should have existed occasionally as to
the age of clay and similar deposits in which some have been found,
or that some of them should have been transferred to genera of other
families, even, as a result of further study; indeed a considerable
number of these fossils appear to have been called oaks rather because
they could be called nothing else than for any very positive other
reason (3).
When the American fossils were first studied, the genus Quercus
was made to include a number of forms that are segregated now in
the genus Dryophyllum (4), held to be prototypic of the family Fagaceae
rather than of its dominant genus, Quercus. The natural early tend-
ency, as would be expected of conservative and experienced botanists
working with isolated and fragmentary leaves, was to stretch the
limits of species so as to recognize identities of Old and New World
species, rather than to see dependable differences in such a repre-
sentation, especially in a genus recognized as unusually variable in the
foliage of even individual trees of existing species. None of the
recently described species has been identified with a European form,
and most of the earlier identities have been discarded, sometimes by
492
TRELEASE: THE ANCIENT OAKS OF AMERICA 493
those who had believed in them at first (5). At present, only the
following identities with European species stand, and these, appar-
ently, because their representations have not been reexamined:
CRETACEOUS—Q. hieracifolia (Kas.), Q. straminea (Col.); EocENE—
Q. Chamissonis (Alaska), Q. doljensis (Wyom.), Q. drymeja (Oreg.),
Q. eucalyptifolia (Col., N. Mex., Miss.), Q. Godeti (Mont.); Muro-
CENE—(Q. elaena (Col.), Q. Steenstrupi (Calif.).. One European species,
Q. Gaudini, very indefinitely reported as American, seems to have no
ascertainable significance.
Unfortunately until recent years nomenclature has been treated
independently in the several branches of natural history, even in
different groups of the same kingdom. Under this uncorrelated
procedure, fossil and existing species have been independently named,
with the result that a given name may refer sometimes to the former
and sometimes to the latter although no idea of identity or even rela-
tionship within the genus has been intended in their designation.
No procedure appears sensible except the restriction of a given bi-
nomial to a single species, and the acceptance of such a name as
valid from its earliest publication, whether for a fossil or extant
species. Application of this procedure causes a considerable number
of changes among the names of American fossil oaks, as well as among
species that are now living (6).
On this continent, as in the Old World, the earliest appearance of
Quercus is in the Cretaceous, for which 48 nominal species are known
from scattered deposits in the Atlantic States of New York, New
Jersey, Maryland, North Carolina and South Carolina; in Kansas and
Nebraska in the Plains region; in Wyoming, Montana, Colorado and
New Mexico in the Rocky Mountains; in Utah in the Great Basin;
and, quite isolated, in Vancouver in the northwest. None of these
species is known to have survived Cretaceous time, and none. bears
striking resemblance to any existing oak, though holly-like leaves were
found then as now.
For the Eocene, 56 nominal species are reported from scattered
deposits in Canada and (perhaps questionably) Mississippi in the east;
from North Dakota, Wyoming, Montana, Colorado and New Mexico
in the interior; and from Oregon, Washington and Alaska in the
northwest. No species is known to have survived into the Miocene,
and none appears to be related to existing species, though holly-like
leaves are represented among these fossils.
The nominal species for the Miocene number 42 and they have
been found in scattered deposits from Maryland, the District of
Columbia and Virginia in the east; and from Colorado, Montana,
Idaho, Oregon, California and Nevada in the west. One of these
33
494 BROOKLYN BOTANIC GARDEN MEMOIRS
Miocene oaks, of California, has been held to be varietally related
to the existing golden oak of California, and is known as Quercus
chrysolepis montana; but little can be said for or against this reputed
relationship. Except for this, none of the Miocene oaks is thought
to have survived.
Little is known of the Pliocene in North America, and it may be
that the sparing deposits in Maryland and Alabama that are supposed
to be of this horizon may be open to some question whether they are
not of more recent age. The 4 nominal species of Quercus that have
been found in them are distinctly more like modern oaks than any-
thing that preceded, but identities with existing species are not
clearly evident to me (7). In South America, several fossil oaks from
the Pliocene have been described, not evidently related to existing oaks,
from localities far from any existing species (8). At present only
four oaks occur in South America; these, which grow in the interior
mountains of Colombia, form a natural group which appears more
closely related to some of the Costa Rican oaks than to any others that
are now known (9).
If the term Pleistocene be used to designate glacial or later deposits
in which fossils are found, it is to be assumed that these fossils will
be very similar to if not identical with existing species. Scattered
deposits of this kind have been examined from various points in the
Atlantic region—Canada, New Jersey, Pennsylvania, Maryland,
Virginia and Kentucky; and from California in the Pacific region.
From these deposits 20 oaks have been named. Two of them, Q.
predigitata Berry and Q. pseudo-alba Hollick, are separately designated
as the ancestral forms respectively of Q. digitata or falcata and Q. alba,
both of which are held to be represented by other Pleistocene material.
A third, Q. abnormalis Berry, may have been a teratological bifid form
of Q. Phellos, which is known in its normal form from Pleistocene
deposits. Concerning a fourth species, Q. Glennit Hollick, I must
admit a serious doubt as to the horizon to which it is ascribed. The
remaining 16 species, into which I have merged Q. abnormalis, Q.
predigitata and Q. pseudo-alba, are easily identified with species now
living in the regions in which they have been found fossilized and, as
would be supposed from this, all of these Pleistocene oaks are from
the Atlantic region, except Q. chrysolepis, which was collected in
California.
Even a cursory inspection of the many illustrations of fossil oaks
that have been published shows that collectively or for any given
period they present a multiplicity of leaf forms more or less com-
parable with what is known for existing species; indeed Professor
Cockerell, who has given much attention to the point, finds in Quercus
TRELEASE: THE ANCIENT OAKS OF AMERICA 495
a good illustration of an ample but inherently limited range of vari-
ation within which in the passage of long periods of time the same
general cycle of forms has appeared repeatedly.
It has seemed to me worth while to arrange the principal leaf types
of the Cretaceous and Tertiary fossils comparatively without regard
to horizon; and for convenience of reference rather than as implying
relationships, those of general comparability are brought together
by the following key:
Leaves entire.
Oblanceolate-obovate, large. (Pl. XIII.) MAGNIFOLIAE.
Lanceolate or oblong, moderate. (PI. XIV.) SIMULATAE,
Leaves toothed.
Teeth numerous, small and sharp.
Leaves elongated, moderate. (PI. XV.) FRAXINIFOLIAE.
Leaves short and broad, small. (Pl. XXII.) SPURIO-ILICES.
Teeth sparse or coarse.
Leaves broad, moderate. (PI. XVI.) DISTINCTAE.
Leaves elongated.
Rather large, not pointed. (PI. XIX.) CASTANEOPSES.
Moderate, acuminate. (PI. XIX, XX.) PAUCIDENTATAE.
Small, not pointed. (PI. XXII.) SPURIC-ILICES.
Leaves crenate or repand.
Rather elongated and small. (Pl. XXII.) MYRICAEFOLIAE.
Broad, moderate.
Some teeth acute. (PI. XVIII.) SUSPECTAE.
Teeth all rounded. (PI. XVII.) DALLIEAE.
Leaves incised or lobed.
Lobes or shoulders 2 or 3, toward the apex. (PI. XXII.) BICORNES.
Lobes or divisions several, not apical.
Leaves small, few-lobed. (Pl. X XI.) LAMBERTENSES.
Leaves moderate or large. (PI. X XI.) LOBATAE.
These foliage-groups scarcely appear to me comparable with
existing oaks except for the Magnifoliae, which suggest certain large-
leaved white oaks of Mexico and Central America; the Simulatae,
which may be held to resemble more or less closely some of the entire-
leaved black and white oaks of the United States and tropical America;
the Spurio-ilices, which parallel the holly-leaved black oaks of Cali-
fornia, and white oaks like Q. Douglasii, many of the scrub oaks of
the table-land, and the dwarf live oaks of the Gulf States; the Pauci- °
dentatae and Castaneopses, somewhat suggestive of existing white
oaks with chestnut-like foliage; some of the Lobatae, comparable
with the white oaks of the United States which have lobed foliage,
as well as with some of the existing chestnut oaks; and the Lamber-
tenses, which resemble if they differ from existing black oaks. Of the
lobed forms, one only, Q. ursina, apart from these, at all recalls our
familiar incised black-oak foliage to my eye.
496 BROOKLYN BOTANIC GARDEN MEMOIRS
The groups which find no existing parallel in the genus may be
questioned as really representative of Quercus. The Fraxinifoliae
are not known since early Tertiary time, but they constituted about
II percent of American Cretaceous species referred to this genus, and
about 17 percent in the Eocene. The Distinctae formed about 15
percent through the Cretaceous, Eocene and Miocene, with changing
species. The Suspectae constituted about 20 percent of the whole
in the Cretaceous, but only about 10 percent in the Eocene, and they
fell to some 4 percent in the Miocene. Neither of the groups that I
have called Myricaefoliae and Bicornes is known to have had more
than a small and transient representation—Q. praeundulata in the
Cretaceous and Q. Ramaleyi in the Miocene for the former, and
Q. bicornis and Q. negundoides in the Eocene for the latter; and the
Dallieae are represented only by Q. Dall of the Eocene.
The ancient foliage types more or less comparable with those of
today show the following relative abundance at different times, so
far as records go: The chestnut type, now most largely represented,
with nearly half the existing American species, formed one tenth of
the whole in the Cretaceous, over a third in the Eocene, and about
one seventh in the Miocene: over a fourth of the known Pleistocene
oaks have this kind of foliage. About 16 percent of the known
living American oaks have lobed leaves, and nearly half of those
known from the Pleistocene are of this general kind; though about a
third of those known for the Miocene are of this type, none of them
has bristle-tipped lobes so far as I know; and in the Eocene only
about 3 percent are found to have had lobed leaves. The pungent
or holly-like type, now constituting about 4 percent of the whole
and represented by one form in the Pleistocene, contained about 15
percent of the Miocene and 9 percent of the Cretaceous ferms, though
it is not yet recognized in Eocene deposits. As might, perhaps, be
expected, entire-leaved oaks, now represented by over a third of the
known species, have been abundant throughout the history of the
genus, and nearly a third of the Cretaceous, a fifth of the Eocene, and a
fourth of the Miocene and of the Pleistocene, species possessed this
type of foliage, which today is often found associated with holly-like
or comparably toothed leaves, often in the same species or even on
the same individual.
It does not seem profitable to attempt to draw climatic inferences
from what I can see in these fossil oak leaves. Some of the entire
leaves appear to have been rather coriaceous, as in certain semi-
xerophytic species now living on the Mexican table-land, and these
and the holly-leaved forms may have been somewhat xerophytic.
Most of the leaves look as if they might have belonged to mesophytes.
TRELEASE: THE ANCIENT OAKS OF AMERICA 497
One group only, the Paucidentatae, even remotely suggests a rain-tip
in its acumination, but the Eocene bicornis and negundoides are some-
what pointed, and some of the more deeply divided forms have acute
lobes, though these do not appear to have been more than mucronate.
One of the characters largely relied on by palaeobotanists is the
venation of leaves. The sigi¥ificance of this has “been insisted on by
Oersted (10) and especially by von Ettingshausen, in the discussion
of Quercus. Some years before his death, this distinguished Austrian
botanist published an extensive tabular comparison of the venation
of existing American oaks and (chiefly European) fossils ascribed to
the same genus (11). For one interested in the Old World fossils,
the table should be most instructive, since it often brings into asso-
ciation a number of fossil species through comparison with a single
living one. On the other hand, the assembling of several existing
species through comparison with a single fossil is suggestive of re-
semblances which might escape notice otherwise and which may indi-
cate some sort of relationship between them.
As a general thing, these venation associations corroborate con-
clusions of affinity based on other considerations, as for instance Q.
macrocarpa*! and stellata* in comparison with Q. Buchii; Q. digitatat,
Kelloggiit, * Leanat and rubrat in comparison with Q. cruciata; and
Q. Douglasii* and stellata* in comparison with Q. cymaena (12).
It is interesting to see that Q. virginiana* is brought into com-
parison with Phellost under Q. elaena, and with imbricariay under
Q. chlorophylla, for even good botanists have found difficulty in
distinguishing between the leaves of some of our Southern live oaks
and the black oaks with willow- or myrtle-like foliage.
Less fortunate associations appear to be those in which unrelated
species are thus brought together; e. g., Q. marilandicat and Warsce-
wiceii* through Q. Zoroastri; Q. magnoliaefolia* and nigra} through
Q. sinuatiloba; Q. Garryana* and ilicifoliat through Q. liriodendroides;
QO. undulata* and Wislizenit through Q. firma; and, especially, Q.
chrysolepis,t grisea,* lanceolata} and laurinat through Q. lauriformis.
Though not necessarily the most abundant at any period, or the
most typical in the Fagaceae, the most synthetic of the many leaf-
forms shown by past and present oaks appears to me to be the sub-
pungent or holly-like type. Those who are familiar with the existing
Californian oaks know with what ease toothing passes into the entire
margin in Q. chrysolepis and its allies, and into the lobed outline in
Q. Douglasii; and it is very frequent in juvenile forms.
‘Engelmann’s confidence in foliage characters was shaken by the
*
1In this comparison, species of Leucobalanus are indicated by *, those of
Erythrobalanus by t, and those of the intermediate Protobalanus by f.
498 BROOKLYN BOTANIC GARDEN MEMOIRS
existing Q. undulata, as he understood that species, in which were
comprised entire, frequently toothed, and very deeply lobed forms,
some of which he was disposed to segregate varietally as others have
done specifically, but all intergrading, as he saw it, as forms of one
single extremely polymorphic species (13).
Similar polymorphism is shown by the many forms of associated
European Tertiary oak leaves studied by von Ettingshausen and
included in his conception of Q. palaeo-ilex. Guided by venation .
studies, he was able to see in this species a foreshadowing of all of the
modern types of oak foliage (14). In the absence of other knowledge
than we now possess, I am disposed to think that in the holly-like form
we may see a starting point for the successive reevolution of the various
forms of leaf that Quercus has presented in the several geological ages,
and now presents.
REFERENCES
1. An indispensable aid in locating first references to American
post-carboniferous fossil plants, in this as in other genera, is Professor
Knowlton’s catalogue of Cretaceous and Tertiary plants, published as
Bulletin no. 152 of the United States Geological Survey—though it
is now antiquated.
2. Fruit of QO. consimilis and Q. paucidentata is figured by Hollick
in Monograph 35 of the U. S. Geological Survey, Pl. 43.
3. Nominal species of Quercus subsequently transferred to other
families than Fagaceae are Q. anceps Lesq. (Diospyros ambigua),
QO. Benzoin Lesq. (Persea Leconteana), Q. californica Lesq. (Mespilo-
daphne pseudoglauca), Q. chlorophylloides Knowlt. (Pisonia chloro-
phylloides), Q. elkoana Lesq. (Carpinus grandis), Q. Lyellit Lesq.
(Nectandra lancifolia), Q. microdentata Hollick (Dillenites microdenta-
tus), Q. Mudgit Lesq. (Protophyllum Mudegi), Q. myrtifolia Lesq.
(Sophora Lesquereuxt), Q. platania Lesq. (Platanus cordata), Q.
retracta Lesq. (Myrica bentonensis), Q. Saffordi Lesq. (Banksia Saf-
fordi), Q. semialata Lesq. (Antsophyllum semialatum).
4. Nominal North American species of Quercus subsequently
transferred to Dryophyllum are Q. crassinervis Lesq. (D. tennesseense),
Q. gracilis Newb. (D. subfalcatum), Q. Mooru Lesq. (D. Moorit).
5. The following European fossil species of Quercus are now believed
to be represented by segregable American forms: Q. acrodon (Q.
Lesquereuxiana Knowlt.), Q. angustiloba (Q. prae-angustiloba Knowlt.),
Q. chlorophylla (Q. chlorophylloides Knowlt.), Q. furcinervis (Q. furci-
nervis americana Knowlt.), Q. Johnstrupu (Q. raritanensis Berry),
Q. Laharpi (Q. fraxinifolia Lesq.), Q. mediterranea (Q. peritula Cock.),
Q. pyrifolia (Q. florissantensis Cock.), Q. voyana (Q. distincta Lesq.).
TRELEASE: THE ANCIENT OAKS OF AMERICA 499
6. The following American fossil oaks require renaming because
their names are preoccupied in the genus: Q. affinis Knowlt., not
Scheidw. (Q. clarnensis n. nom.), Q. Breweri Lesq., not Wats. (Q.
Berryin. nom.), Q. cuneata Newb., not Wang. (Q. Newberryi n. nom.),
Q. elliptica Newb., not Née (Q. washingtonensis n. nom.), Q. latifolia
Lesq., not Steud. (Q. dryophyllopsis n. nom.), Q. laurifolia Newb.,
not Michx. (Q. Penhallowi n. nom.), Q. montana Knowlt., not Willd.
(Q. Cockerellii n. nom.), Q. occidentalis Knowlt., not Gay (Q. van-
couveriana n. nom.), Q. pandurata Heer, not H. & B. (Q. alaskana n.
nom.), Q. salicifolia Newb., not Née (Q. Eamesi n. nom.), Q. sinuata
Newb., not Walt. (Q. prae-undulata n. nom.), Q. Turneri Knowlt.,
not Willd. (Q. prae-dumosa n. nom.).
7. North American oaks ascribed to the Pliocene are Q. cates-
baeifolia Berry, Q. lambertensis Berry, Q. nigra L. fide Berry, OQ.
prae-virginiana Berry. See Berry, Professional Paper no. 98L, U.
S. Geol. Surv., p..200.
8. Brazilian Pliocene oaks are Q. brasiliensis Krass., O. Hussakii
Krass., Q. prae-mespilifolia Krass., Q. pseudodaphnes Krass. See
Krasser, Sitzungsber. K. Akad. Wien. 112!: 854.
g. A summary analysis of existing American oaks is given by the
writer in Proc. Nat. Acad. Sci. 2: 626-9.
10. Oersted, Bidrag til Kundskab om Egefamilien i Nutid og
Fortid (Vidensk. Selsk. Skr. Naturvid.-math. Afd. 5 ser. 6: 335),
Copenhagen, 1871: Chénes de 1’ Amérique Trop., p. 6.
11. Von Ettingshausen, Denkschr. K. Akad. Wien. Mathemat.-
Naturwiss. Classe. 63: 126. This discussion stands in relation with a
series of three largely illustrated ‘‘Beitrage zur Erforschung der
atavistischen Formen an lebenden Pflanzen und ihrer Beziehungen zu
den arten ihrer Gattung”’ (op. cit. 54: 245-254; 55: 1-38; 56: 47-68),
published jointly with Krasan.
12. The following names of European fossil oaks employed in von
Ettingshausen’s comparison tables are preoccupied in the genus: (Q.
affimis Sap., not Scheidw. (Q. paramoea n. nom.), Q. crassipes Heer,
not H. & B. (Q. pachypoda n. nom.), Q. undulata Web., not Torr.
(Q. cymaena n. nom.).
13. Engelmann, Trans. Acad. St. Louis 2: 372; Bot. Works. 389.
14. Von Ettingshausen, /. c. 125.
500 BROOKLYN BOTANIC GARDEN MEMOIRS
EXPLANATION OF PLATES XIII-XXII
PLATE XIII. MAGNIFOLIAE.—CRETACEOUS: 1. Q. Wardiana, Lesquereux,
Monog. U.S. Geol. Surv. 17. pl. 7—EoOcENE: 2. Q. magnifolia. Knowlton, Monog.
U. S. Geol. Surv. 32. pl. 88.
PLATE XIV. SIMULATAE.—CRETACEOUS: I. Q. montanensis. Knowlton, Bull.
U.S. Geol. Surv. 163. pl. I. 2. Q. coriacea. Hollick, Monog. U. S. Geol. Surv. 35.
pli 19. 3. Q. Eamesi, n. nom. Hollick, Monog. U. S. Geol. Surv. 35. pl. 1 (as Q.
salicifolia Newberry, a preoccupied name). 4. Q. glascoena. Lesquereux, Monog.
U. S. Geol. Surv. 17. pl. 6. 5. Q. Morrisoniana. Lesquereux, Rep. U. S. Geol.
Surv. 8. pl. 17.—EocENE: 6. Q. neriifolia. Lesquereux, Rep. U. S. Geol. Surv. 8.
pl. 31.—MI0cENE: 7. Q. convexa. Lesquereux, Rep. U. S. Geol. Surv. 8. pl. 45 B.
8. Q. dayana. Knowlton, Bull. U. S. Geol. Surv. 204. pl. 6. 9. Q. simulata. Knowl-
ton, Rep. U. S. Geol. Surv. 18%. pl. ror. 10. Q. elaena. Lesquereux, Rep. U. S.
Geol. Surv. 8. pl. 28.—Horison?: 11. Q. cinereoides. Lesquereux, Rep. U. S. Geol.
Surv. 75 pls 20,
PLATE XV. FRAXINIFOLIAE.—CRETACEOUS: I. Q. flexuosa. Hollick, Monog.
U.S. Geol. Surv. 35. pl. 19. 2. Q. banksiaefolia. Hollick, Mon. U.S. Geol. Surv. 35.
pl. 18. 3. Q. Holmesit. Lesquereux, Rep. U. S. Geol. Surv. 1874. pl. 8 (as Q.
salicifolia).—EoOcENE: 4. Q. fraxinifolia. Lesquereux, Rep. U. S. Geol. Surv. 7.
pl. 20. 5. Q. Haidingert. Lesquereux, Rep. U. S. Geol. Surv. 7. pl. 20. 6. Q.
consimilis. Hollick, Monog. U. S. Geol. Surv. 35. pl. 43. 7. Q. drymeja. Les-
quereux, Rep. U. S. Geol. Surv. 7. pl. 10; 8. pl. 54. 8. Q. Crossii. Lesquereux,
Proc. U. S. Nat. Mus. ro. pl. 2.
PLATE XVI. DisTINCTAE.—CRETACEOUS: I. Q. alnoides. Lesquereux, Monog.
U. S. Geol. Surv. 17. pl. 7. 2. QO. Lesquereuxiana. Lesquereux, Rep. U. S. Geol.
Surv. 7. pl. 19 (as Q. acrodon).—EocENE: 3. Q. viburnifolia. Lesquereux, Rep. U. S.
Geol. Surv. 7. pl. 20.—MIocENE: 4. O. distincta. Lesquereux, Mem. Mus. Comp.
Zool. 67. pl. 2—UNPLACED TERTIARY: 5. Q. aquamara. Ward, Bull. U. S. Geol.
Surv. 37. pl. 2. 6. Q. carbonensis. Ward, Bull. U. S. Geol. Surv. 37. pl. 9.
PLATE XVII. DALLIEAE.—EOCcENE: Q. Dailii. Lesquereux, Proc. U. S. Nat.
Mus. 5. pl. 8. .
PLATE XVIII. SuspecTAE.—CRETACEOUS: I. Q. suspecta. Lesquereux, Monog.
U.S. Geol. Surv. 17. pl. 47. 2. Q. dryophyllopsis n. nom. Lesquereux, Rep. U. S.
Geol. Surv. 1874. pl. 6 (as Q. latifolia, a preoccupied name). 3. Q. Cockerellii n. nom.
Knowlton, Bull. U. S. Geol. Surv. 257. pl. 17 (as Q. montana, a preoccupied name).—
EocENE: 4. Q. pseudo-alnus. Lesquereux, Rep. U. S. Geol. Surv. 8. pl. 53. 5. Q.
Culveri. Knowlton, Monog. U. S. Geol. Surv. 322. pl. 87.
PLATE XIX. CASTANEOPSES.—EOCENE: I. Q. castaneopsis. Lesquereux, Rep.
U.S. Geol. Surv. 8. pl. 28. 2. Q. groenlandica. Hollick, Monog. U. S. Geol. Surv.
35. pl. 54.—PAUCIDENTATAE.—EOCENE: 3. Q. clarnensis. Lesquereux, Rep. U. S.
Geol. Surv. 8. pl. 53. (as Q. furcinervis).—4. Q. nevadensis. Lesquereux, Mem. Mus.
Comp. Zool. 6°. pl. 2.
PLATE XX. PAUCIDENTATAE.—EOCENE: I. Q. yanceyi. Knowlton, Monog. U.
S. Geol. Surv. 32”. pl. 89. 2. Q. paucidentata. Hollick, Monog. U. S. Geol. Surv.
35. pl. 43. 3. Q. pseudo-castanea. Unger, Palaeontographica. 2. pl. 35.—MIOCENE:
4. Q. Osbornii. Lesquereux, Rep. U. S. Geol. Surv. 8. pl. 38.
PLATE XXI. LosBatarE.—MIoceneE: 1. Q. Merriami. Knowlton, Bull. U. S.
Geol. Surv. 204. pl. 6. 2. Q. pseudo-lyrata. Lesquereux, Mem. Mus. Comp. Zool.
6. pl. 2. 3.Q. Milleri. Berry, Journ. Geol. 17: 24. f. 3. 4. Q. duriuscula. Knowl-
ton, Bull. U. S. Geol. Surv. 204. pl. 8—LAMBERTENSES.—MIOCENE: 6. Q. chapmani-
BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME |, PLATE XIll.
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TRELEASE: THE ANCIENT OAKS OF AMERICA 501
folia. Berry, Prof. Paper U. S. Geol. Surv. 98L. pl. 11. 7. Q. Lehmanni. Hollick,
Md. Geol. Surv. Miocene. pl. 483.—PLIOCENE: 8. Q. lambertensis. Berry, Prof.
Paper U. S. Geol. Surv. 98L. pl. 11. :
PLATE XXII. BIcORNES.—EOCENE: I. Q. negundoides. Lesquereux, Rep. U.S.
Geol. Surv. 7. pl. 21. 2. Q. bicornis. Ward, Bull. U. S. Geol. Surv. 37. pl. 9.—
MyRICAEFOLIAE.—CRETACEOUS: 3. Q. praeundulata. Hollick, Monog. U. S. Geol.
Surv. 35. pl. 31 (as Q. sinuata Newb., a preoccupied name). 4. Q. antigua. Hollick,
Monog. U. S. Geol. Surv. 35. pl. 13.—MIOCENE: 5. Q. Ramaleyi. Cockerell, Bull.
Torr. Bot. Cl. 33: 309.—SPURIC-ILICES.—CRETACEOUS: 6. Q. Haydeni. Lesquereux,
Rep. U. S. Geol. Surv. 7. pl. 19. 7. Q. spuric-ilex. Knowlton, Monog. U. S. Geol.
Surv. 17. pl. 48.—MIocENE: 8. Q. Applegatei. Knowlton, Rep. U. S. Geol. Surv.
20°. pl. 1. 9. Q. distincta. Lesquereux, Mem. Mus Comp. Zool. 62. pl. 2 (as Q.
voyanda).
THE ABSORPTION OF CALCIUM SALTS BY SQUASH
SEEDLINGS*
RODNEY H. TRUE AND R. B. HARVEY
Bureau of Plant Industry, U. S. Department of Agriculture
In the course of a former investigation! carried on with the white
lupine (Lupinus albus L.) as a test plant, it was found that the rate of
absorption of electrolytes by seedlings from solutions of the usual
mineral nutrients was influenced to a great degree by the chemical
character of the substances offered. In simple solutions it appeared
that the usual salts of potassium were not absorbed producing rather
an outgoing current of ions from the seedlings into the solutions;
in magnesium solutions a small absorption took place in the more
dilute solutions, while in calcium solutions absorption was much greater
and took place in the most dilute solutions. It appeared from the
behavior of the lupine in the solutions of calcium sulphate and calcium
nitrate that in each case the quantity absorbed is limited, even though
the supply may contain a large excess, and the effect of the anion
appears to be subordinate to that of the Ca ion in determining the
quantity absorbed.
It was found by preliminary experiments by the present authors
that this condition of things does not apply to all kinds of plants.
The common garden squash, sweet corn and soy bean were found
to behave quite differently with respect to the anion employed with
the Ca ion. In this paper the records of a series of experiments are
presented showing the absorption from a series of solutions of three
of the commoner inorganic salts of calcium by the seedlings of Cucur-
bita Pepo L. of the horticultural variety known as Early Prolific
Marrow.
Before considering the evidence on which the conclusions here
advanced are based a word concerning the method is in order.
Selected seedlings obtained from seeds germinated in chopped sphag-:
num were grown in carefully prepared solutions contained in prac-
tically insoluble glass beakers. The concentration of ions of each
solution was noted daily by taking conductivity readings by means
of an accurate wheatstone bridge. The temperature throughout was
* Published by permission of the Secretary of Agriculture.
1 True, R. H., & Bartlett, H. H. Am. Journ. Bot. 2: 278. 1915.
502
TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 503
maintained at 18° C. by automatic control so-accurate that the range
of variation was seldom above four tenths of a degree Centigrade
during the course of an experiment running a fortnight. The con-
tainers remained in darkness except during the short time required
for the determination of the conductivity which took place in rather
faint diffused light. Since it is obviously unsafe to draw conclusions
from a comparison of ohms, results were always calculated to concen-
trations expressed as gram-normals of the salt in question dissolved
in a million liters of water (grm. norm. X 10-°). The water was
obtained by twice distilling Potomac River water from glass with
electric heat in a laboratory from which gas was excluded. Each
experiment was usually continued until signs of deterioration began to
appear in the seedlings.
CALCIUM NITRATE
Several experiments were carried out with squash seedlings in
calcium nitrate solutions. Since they were in close agreement but
one is presented here, that running from May first to May fifteenth,
last, inclusive. The distilled water used in making up the solutions
had an initial conductivity equal to that of a solution containing 11.7
grm. norm. X 10-® Ca(NOs)o. Nine cultures each containing 5 seed-
lings and 500 cc. of solution were set up in a series ranging in con-
centration from 18.2 to 867.0 grm. norm. X 10-*. Daily observations
were made until signs of exhaustion began to appear. Since in the
cultures containing less than 50 grm. norm. the behavior of the seed-
lings varied so little in the different members of the series only a part
of the record is shown here in order not to confuse the table with
several nearly coinciding curves. In the curve representing the
record of the culture in distilled water a dashed line is employed
(Fig. 1).
It will be observed that in both distilled water and in cultures
containing calcium nitrate up to a concentration of 100 grm. norm.
x 10-° the solutions gain in concentration for two or three days, a
course which in the distilled water is followed by a very slight ab-
sorption until near the close of the experimental period. At no time,
however, were the plants able to regain any considerable proportion
of the electrolytes lost to the medium during the first few days.
With the dilute solutions of the salt (under 100 grm. norm.) this
period of leach gradually passes over into one of active absorption
as a result of which these solutions are reduced to a lower ion content
than the distilled water.
As the initial salt content of the solution is increased to approxi-
mately 500 grm. norm. the slight leach seen in the weaker solutions
504 BROOKLYN BOTANIC GARDEN MEMOIRS
fails to appear, immediate though slow absorption being the rule.
This intake gradually speeds up through the succeeding days, the
absorption being roughly related to the original quantity of salt in
the solution. This active absorption continues to the end of the
experiment at which time the appearance of the seedlings and the
SS ee
Jersger sie) | "| aia
a
\ SQUASH CaoNOLIEC| |_|
CA
NORMAL CONCENTRATION X_10°
S
SECON SL
SEENGRGRRSNGEe
oT tf eae 7 a 9 OT ee
Fic. 1. For explanation see text.
diminishing absorption indicate approaching exhaustion. At this
final stage of the experiment the plants have reduced the ion content
of solutions of the salt to an approximately like minimum, which lies
between 35 and 50 grm. norm. X 10-® Ca(NOs3)o. This concentration
lies at about 25 grm. norm. X 10~° below the concentration reached
TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 505
by the distilled water; and seems to represent a fairly well-defined
irreducible minimum. This point is marked by a lower ion content
than the similar minimum seen in the distilled water. It seems clear
that the substances giving to distilled water its conductivity are not
absorbed to as dilute a minimum concentration as is calcium nitrate.
It will be noted that only in the two more concentrated members
of the series are there any absorbable electrolytes remaining at the
end of the experiment, and therefore only in these cultures is the total
capacity of these plants to absorb this salt measured. In these cases
the total absorbed salt equals 565 and 713 grm. norm. X 10-* re-
spectively.
An inspection of the graphs shows that in nearly all solutions of
the salt there occurs a time at which the plants reduce the conductivity
of the solution to a minimal concentration, a point that may be
assumed for present purposes to represent that of maximum absorption.
In some cases experiments have been closed before this point has been
reached in a number of members of the series (usually the more con-
centrated ones), owing to the well-marked exhaustion of the plants
in a number of the cultures. In such a case the absorption maximum
may not have been reached, although probably in most cases it has
been approached.
It is interesting to compare the original concentration of the differ-
ent solutions with the corresponding concentration at the time of
maximum absorption. By this means one is able to ascertain how
much of the salt is absorbed or how much net loss the plants have
suffered calculated at the time of greatest absorptive efficiency.
Carrying out the calculation referred to for the cultures included in
this experiment the results seen in the following table (1) are obtained.
A glance shows that about 37.0 grm. norm. Ca(NOs)» are required
by five squash seedlings growing in 500 cc. of solution to enable them
to protect such ions as are mobilized from their reserves against the
leaching action of the distilled water. This stated in terms of the
quantity of Ca(NOs)2 per plant would be about 0.00028 grm. absorbed
during a period of about two weeks.
As the salt content is increased, absorption increases in approxi-
mately the same magnitude leaving an unabsorbable residue of
approximately 34 grm. norm. X 10-§ Ca(NOs)2. This minimum is
here calculated as Ca(NOs3)o, but it should be clearly understood that
the substances actually composing this residue may be, and probably
in considerable part are, of quite other composition. Indeed, the
substances indicated as Ca(NOs)2 at all stages of the experiment,
especially the later ones, doubtless consist in part of other materials,
largely coming by exosmosis from the interior of the test plants. The
35
506 BROOKLYN BOTANIC GARDEN MEMOIRS
TABLE I
Net Absorption by Squash Seedlings from Solutions of Ca(NOs)2
Original Concent. of Minimum Concent. of Maximum Net Absorp-
Solution Solution as tion Calc. as
Ca(NOs)2 X1076 Ca(NOs)2 X 10-6 Ca(NOs)2 X 107-8
HALO) Caen WOM 5 os ascons 50:0 grim. noOTmry.. .. 4... - — 38.0 grm. norm.
EO:2 fae erg yy ae Aigoy as ine Se — 24.008" z
Py etd Basen thos. Bylo) righ AS ai Pte - 95 “ *
BO! ie te ONE estore 32.1 A ME ete cree + 43 “ <
ayshioy separ Siu 37/0 SS phe oes Qe) e
Aiea PUR Ay cus Stmte 20:6— Bids coe cic Ov Dae 2
45h5 un iD BA teres Skee 26:30 PR, ieee eS 0:2". i
63:0) aN Roe o oe eylfey & ae Ss ot epeh e
96:2) an Ses cel By Oma rt bee iti 62:3. 9% +.
TO2,0us ee eR es 20160 Ase Mes 153207 S
351.8 * 3 re eer 39.1 : Serres es 21257 H ‘
[iGo anrg eM ES as tae Se aa AT Ole gee na any etch oe 470.7
692555 5 CioULE Ss Seen elas Eas 1250 ASTRA he renee 567.6 “ rs
86720) ae Saad tee 15220 memes peers he: AT aes oe
concentration indicated in any case merely states the net result of
ion interchange in a given culture stated in terms of such a solution of
the given salt as would have a like electrical conductivity.
As the quantity of salt is still further augmented, the unabsorbed
residue increases, indicating a clear surplus of this constituent. It is
interesting to note, however, that the unabsorbed residue does not
increase at a rate parallel with the quantity offered; the absorption
increasing also though at a lagging rate. The maximum quantity of
Ca(NO3)2 absorbed by five plants out of a concentration of 867 grm.
norm. X 10-* is about 714.1 grm. norm. This expressed in weight of
salt absorbed per seedling would be about 0.0055 grm. This may be
taken to represent approximately the maximum quantity of Ca(NOs)e
absorbed by a squash seedling living in darkness in a temperature of
18° C. during the time required by the seedling to exhaust its available
reserves.
CALCIUM SULPHATE
A somewhat similar experiment was made in which calcium sulphate
was used, in the hope that we might get some light on the comparative
effect due to the sulphate and the nitrate anions. A series of 14 cul-
tures was set up using squash seedlings as before covering a range of
concentration between distilled water and 830 gram normals of the
salt in a million liters. The experiment ran from July 12 to 23, 1916.
A graphic record® of the course of the concentration changes in the
several cultures is shown in the accompanying diagram (Fig. 2):
2 In order not to crowd the figure only one curve representing an original
CaSO, concentration less than 50 grm. norm. is given.
TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 507
A glance at the curves shows that in some respects the course of
absorption is markedly different from that seen in the case of Ca(NOs)2.
Alike in the distilled water control and in the solutions originally
containing 11.08 and 15.6 grm. norm. respectively, the solutions gain
ions and not even at the time of greatest absorption are the plants able
to reduce the ion concentration to that seen at the beginning of the
experiment. These quantities of the salt in question are insufficient
to prevent the solutions from acting practically like distilled water
ad
0; ~ —
PPR LTT TL [saasy asa vec. | |
as aa =Seatee : 850
2c eee
Ae eae
2 60\— Lf
S cn
Bo
S
§
4
2 sis 14 LSS ME SEI 4e:
Fic, 2. For explanation see text.
by withdrawing a preponderating quantity of ions from the seedlings.
When the concentration of CaSO, reaches about 30 grm. norm. in a
million liters, an equilibrium of some sort seems to be established
between the plant and the medium with the result that neither ab-
sorption nor leach is marked. A similar equilibrium point appears
at approximately the same concentration in the case of Ca(NQOs3)>
solutions. It is important to note that the concentration of the
calcium sulphate solutions at the time of maximum absorption in
508 BROOKLYN BOTANIC GARDEN MEMOIRS
these most dilute solutions is higher than is the case of those originally
having a greater salt content, seeming to indicate that at this extreme
dilution the plants are not only forced to yield ions to the solutions
but are unable by reabsorbing them to reduce the concentration to a
point lying much below the equivalent of 50 grm. norm. in a million
liters.
As the original concentration of CaSO, is increased to 31.3 and
52.1 grm. norm. respectively absorption increases to a point showing
.net gains by the plants. Leaching of ions by the plants, if it takes
place, is more than met here by their greater absorptive activity.
This activity when greatest reduces the residual ion content markedly
below that seen in the case of the originally more dilute solutions.
It seems that with the addition of even slightly larger quantities of
CaSOx,, the absorptive function becomes more active and is able more
nearly to exhaust the quantity of ions offered.
As the quantity of CaSO, is increased to concentrations rising
from 101.9 to 824.4 grm. norm. in a million liters the plants reduce
the ion content of all solutions but absorption even at its greatest
leaves a residue which increases as the quantity offered increases.
Absorption, while in general increasing as the quantity of ions present
increases, lags far behind the quantities offered. It thus comes about
that the curve representing the residual ion content approximately
parallels that representing the original ion content of the solutions.
From these data it would seem that when squash seedlings are
grown in darkness at 18° C. in solutions of CaSO, alone, they are
unable to absorb as many ions as they lose when the solution contains
less than about 30 grm. norm. of the salt in a million liters. When the
solutions contain somewhat more than this quantity of this mixture of
ions the plants can reduce the more dilute members of the series to an
ion content of about the concentration seen at this equilibrium point.
When the supply of ions is far in excess of absolute requirements, the
seedlings absorb greater quantities than in weaker solutions but the
quantity of residual ions left in the solution increases in nearly the
same proportion as the quantity offered.
In the following table (Table 2) are shown (1) the original con-
centration of each solution, indicating the quantity of ions offered
in each case; (2) the concentration of each solution at the time of
maximum absorption, or the residual ion content of each solution and
(3) the quantity of ions absorbed from each solution at the time of
maximum absorption.
TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 509
TABLE 2
Net Absorption by Squash Seedlings from Solutions of Calcium Sulphate
Original Concentration Minimum Concentration Maximum Absorption
as Grm. Norm. as Grm. Norm. as Grm. Norm.
CaSOs X10-6 CaSO X 10-6 CaSOs X 10-6
AMORA eet hha Sead ASO ey Siict achat aches — 38.7
PROS Mb ari Gis eae ays Ws Glnae opto Bole ice toe —sAQIZ2
1B Olends chegcnae WP CNA eet eas BOLOMe Ries ciate ee — 34.4
RTD aS Oo vents aiken. es es DULG A steve Qn dic bane Fi o.06, a15 3 ae ey!
Ise a Meee des tyra cena ee aie SRG kis SA RA eee aect se Poe 14.4
MOOR sc eee eens Cee ee OO) ier sie oer Bie seks 25.9
UA Ordre ae eles chore. aca (OPIN Oe 55s cei ACCORD Eat 57-4
TOTES eten aekc ne Sir oe eee INGA S alee ad erotic 51.3
Bil slr Pee tet y ous erons fag ees POI IP eto: ety Chere re Rie 57-6
AMAR sett SH ee Oke cs AG ais Seclice CE EE ce eae 78.8
Hide iran ceo hecri inte s & BASES cichoee Gia y eb ees 87.3
(OY Ia Le ote Ghote ine erat eee as 2 OM ba Mere tone ene ites 125.7
FR RL ON ae aieheoete arciee on Centar OTTO erecta r es, 112.0
SZ AP A eater heres test seas se LOARO ivattncre atte isle 119.8
2 CALCIUM CHLORIDE
In the foregoing experiments we have had to do with salts in which
both cation and anion are required for the normal growth.of higher
plants, and we find squash seedlings behaving toward them in sharply
contrasting ways. From solutions containing Oe and NO; ions root
absorption is very active. When ce ions are accompanied by SO,
ions, however, absorption is relatively small.
In view of these facts it is a matter of no small interest to test the
behavior of plants in solutions in which the Ca ion is accompanied by
the Cl ion.
A series of solutions was made up containing a graded series of
concentrations running up to 582.4 grm. norm. in a million liters.
The experiment ran from Aug. 9 to Aug. 26, 1916. A graphic record
of the concentration changes observed is shown in the accompanying
group of curves (Fig. 3). It is perhaps hardly nécessary to do more
than to point out certain of the more important features to be observed.
In general a very striking similarity to the corresponding set of curves
obtained from Ca(NQOs3)2 solutions may be noted. The solutions hav-
ing an original concentration of 32.5 and less show a loss of electrolytes
for the first five or six days after which time absorption begins and
continues until near the end of the experiment, but in none is the
loss during the early days fully regained, although in that originally
containing 32.5 grm. norm. the net loss is practically negligible.
At this concentration we find again evidence of a critical concentration
of some sort below which the plants can not absorb and above which
510 BROOKLYN BOTANIC GARDEN MEMOIRS
they are able to do so. This required minimum quantity is probably
of considerable physiological significance. At higher initial con-
centrations, from 53.5 to 384.0 grm. norm. loss of ions during the
early days of the experiment becomes less marked as the concentration
increases. Absorption usually begins more promptly and proceeds with
greater speed as the quantity of ions present increases until in all cases
but one the concentration of ions remaining in the solution is reduced
700 700
renee eeeee |
ee
[a | Ssewsy Gch. 1eCl aie
eee eee
650
600
550
~)
ede |e
>40
: ERAGRAERe
wi 400
ea ON EN No
Sa eS 350
S er at SS ey es
& 300
: 0
S
=
=
N
W
aN
Q
0’
N
ie)
to the unabsorbable minimum at an average concentration of approxi-
mately 27 grm. norm. X 10~®. Only when the original concentration
of 582.4 grm. norm. X 10~° is reached do we find the quantity of ions
offered greater than can be reduced by the seedlings to the unab-
sorbable minimum. These relations are seen in the numerical data
given in Table 3, in which the concentration relations are shown as
they exist at the time of maximum absorption. Here again the
similarity to the situation seen in solutions of Ca(NOs)s is striking.
If we are justified in concluding that the depressing action of the
SO, anion is responsible for the great reduction of root absorption
TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS Sit
TABLE 3
Net Absorption by Squash Seedlings in Solutions of Calcium Chloride
Criginal Concentration of Minimum Concentration of Maximum Net Absorp-
Solution Grm. Norm. Solution as Grm. Norm. tion as Grm. Norm.
CaCh X 10-6 CaCle X 10-6 CaCle X 10-6
AL SUOuie Okc e ale 28.5. 9b MOLm. aise actos — 34.3 grm. norm.
Os Seti t natasha 2 aT es fw ERE ENE 2 — 15.9 “
Toy Ok eset recone! 26:50. NTE ERS COR eps — 10.8 ‘ s
Berane” ieee ed at Bee) ft Pee ee = ini © ~
Be.5) Seah ta ote cos Ae sete tot ee ae + 108 “
Fist an Senet ie eee ae 20:50 a Meee ee ee: Gils)
m1G:5 °°.) RRS STI TS: TOMS Pe nyt, date 96:3 :
191.8 “ Pe eee ae eee 19.9 “ Map errtetS : 70 OM .
Sen = A ee erat aot ioe Sa ashe etry eee: 25 aes
Bao, ii Soha. oe eos eae Ti peony, sg teas aang 5
seen in the solutions of CaSO, when compared with that seen in solu-
tions of Ca(NOs)2 of a similar range of concentration, we are also
justified in concluding that the influence on process of absorption due
to the Cl arjion is as favorable as that exerted by the NO3 anion.
DISCUSSION OF RESULTS
It seems clear that for both the squash and white lupine in the
seedling stage the calcium ion favors the absorption of ions. The
lupine while finding this ion necessary does not absorb it in as large
quantities as does the squash. The maximum requirement per cul-
ture of five seedlings of the lupine under the conditions of these experi-
ments is not over 175 grm. norm. X 10-® Ca(NOs3)2 and about 125
Sem. tlorm. < 10-* CaSQ,.®
The squash on the other hand reduces an original concentration
of 518 grm. norm. Ca(NOs3)2 X 10-® to 47.8 grm. norm., thus finding
but a very small surplus present, and may absorb more than 700
grm. norm. when a concentration of 867 grm. norm. X I0~* is offered.
It is more quickly satisfied in the case of CaSQu,, this plant absorbing a
maximum of about 120 grm. norm. only from solutions increasing in
concentration up to about 825 grm. norm. X 10~*. It takes in there-
fore from an excess supply about as much CaSQ, as does the lupine.
This comparison gives additional evidence of the well-known
indifference of the white lupine toward calcium in several combinations.
From the evidence at hand it appears that in the presence of the
Ca ion, the effect exerted by the NO; and SO, anions on absorption by
the lupines is not markedly different. With the squash the anion
effect comes strikingly to the front. The Ca ion accompanied by the
§ Calculated from True, R. H., and Bartlett, H. H. Am. Journ. Bot. 2: 262 and
265. I915.
512 BROOKLYN BOTANIC GARDEN MEMOIRS
NO; ion is from four to five times as favorable for absorption under the
conditions of these experiments as the Ca ion accompanied by the SO,
anion. This seems to indicate a striking and specific difference in
the influence of these anions on the absorptive activities of the squash.
When the Ca ion is accompanied by the Cl ion, absorption is influenced
very much as in the case of Ca acting with NO; ions. The favorable
effect of NO; and Cl ions is contrasted with the action of the SO, ion.
The strong influence exerted by the specific characteristics of the
different species of plants is seen in the contrasting behavior of the
lupine and the squash in the presence of Ca accompanied by the NO;
ion.
It should be borne in mind that the probable physiological inter-
action of a given pair of ions is perhaps such as to make it unsafe to
speak strictly of the specific action of any single ion irrespective of
that of its companion ion or ions.
It is obvious from what has been here shown that any theory of
cell permeability which may be framed to account for the income
and outgo of the living plant with respect to electrolytes must reckon
with the striking differences that exist in the behavior of plants toward
even such fundamental factors as the required mineral nutrient ions.
INHERITANCE STUDIES ON CASTOR BEANS
ORLAND E. WHITE
Brooklyn Botanic Garden
Ricinus, though a monotypic genus involving only a single widely
recognized species (R. communis), possesses a multitudinous number
of forms, which from time to time have been temporarily ranked as
species. These forms breed true to‘ many of the numerous characters
which distinguish them, as shown by data obtained from growing
several generations of fifty or more types in the experimental breeding
plots of the Brooklyn Botanic Garden. Numerous crosses between
even the most extreme types have given perfectly fertile F,; and Fy,
generation hybrids.
Hybridization studies to determine the manner of inheritance of a
dozen or more of these characters have been followed through the
Fi, Fs, and, in some cases, the F3 generations. Several thousand
plants were involved in these studies.
MATERIALS AND METHODS
Seeds of the various types were secured through Farquhar & Co.
of Boston, P. Henderson & Co. of New York City, and from various
botanic gardens. Many of these types are known in seedsmen’s
catalogues as varieties or sub-species, and these, much to my surprise
(since the castor-oil plant is monoecious and wind-pollinated), bred
true immediately to many of their more prominent characteristics,
such as stem color, seed color and color pattern, and height. Further
observations on plants of different varieties grown close together
demonstrated that very little cross-fertilization took place (certainly
not more than five percent), even when conditions were most favor-
able. This rather unexpected tendency to self-fertilization in a
monoecious plant adapted apparently to wind-pollination is largely
due to the slightly earlier maturity of the male flowers and to the
comparative isolation of the flowers of each plant through the preven-
tion of air currents by the large leaf surfaces. As the stigmatic surfaces
of the female flowers become exposed and mature, the pollen from flow-
ers on the same plant has already fallen or falls upon them in small
clouds, thus insuring, to a large extent, self-fertilization.
Difficulty is experienced under Long Island climatic conditions in
making bagged inflorescences on outdoor cultures, set a normal amount
513
514 BROOKLYN BOTANIC GARDEN MEMOIRS
of seed. Because of this, the F; generation plants, in some cases, were
grown in isolated cultures, instead of being bagged. In this way
large quantities of seed were obtained for growing F»: populations.
In most cases, enough seed from bagged F, plants was obtained to
check up the F, results from the unbagged seed. A large number of
F; families were grown from seed of unbagged Fy, plants. The per-
centage of cross-fertilization among the Fy, individuals appeared to
be small.
EXPERIMENTAL WORK
Stem Color
Stem color in castor beans can be roughly classified into five cate-
gories (see Plates XXIII and XXIV)—bright green, green with reddish
blush on sunny side, carmine or rose red, mahogany red, and purple
(dark red). The development of each type of red coloration depends,
to some extent, on sunlight, particularly the red blush class. In
shade, plants of this class have green stems.
These red color types are similar chemically, so far as solubility
tests with their pigments are concerned, since all the red pigments are
soluble in water containing a trace of chloroform, but remain insoluble
in alcohol, xylol, or in pure chloroform. The red pigment is a sap
color, occurring in the epidermal palisade cells in the leaf and stem
(see Plate X XIII), also in parenchymatous areas of the stem, especially
in the pith in some varieties. The different shades are apparently
due (as observed microscopically) to different concentrations of the
pigment in the cells and to a difference in the amount of pigmented
cell area. From evidence thus far obtained, the writer is inclined to
regard the hereditary differences between the red types as due to the
presence and absence of several color dilution factors, each of which
modifies the expression of a red pigment producing factor common to
all, except, of course, the green-stemmed class. Still further modi-
fications in coloration appear to be due to the presence and absence
of a pattern restriction factor, in the absence of which the stems and
leaves are mahogany red, or purplish red if ‘“‘bloom”’ is present.
In crosses, red blush X other red blush varieties always gave
only red blush F, and Fy, offspring. Red blush varieties X green-
stemmed varieties and the reciprocal always gave all red blush in Fy
and approximately 3 red blush: 1 green in Fs.
Actiallyiobtalned s.r cit cent ieerneier 113 red blush: 43 green
Mheoretically-expectedas heer eeten 1 Alege 20,08
In F;, seed from unbagged F», green-stemmed plants generally gave
all green plants, while unguarded seed from F, red blush plants either
gave all red blush or both red blush and green-stem families.
WHITE: INHERITANCE STUDIES ON CASTOR BEANS 515
Red blush X mahogany and reciprocal gives rose or carmine-
stemmed plants in F,, and approximately 1 red blush: 2 rose: I
mahogany in F», the actual figures being 47 red blush: 144 rose:
47 mahogany, theoretically expected 59.5 red blush: 119 rose: 59.5
mahogany. No F3; generation of this cross has been grown, but from
the above ratio the plants with rose-colored stems are expected to
produce all three F. types again, while the other two types are ex-
pected to breed true.
As previously stated, there are forms with rose-colored stems that
breed true. These when crossed with red blush varieties give all
rose-stemmed F, plants, indistinguishable from their rose-stemmed
parent. In Fs, these produced 429 rose: 145 red blush, the the-
oretically expected proportions being 430.5 rose: 143.5 red blush.
Associated with the types of stem coloring are pigmented areas
in other parts of the plant. The mahogany-red-stemmed plants have
mahogany-red leaves and fruits. The rose- and red-blush-stemmed
types have green leaves with red or reddish-green midribs. The dark
purplish-red= (mahogany bloom) stemmed plants have dark purplish-
red leaves and fruits. Plate XXIII is a much enlarged micro-photo-
graph in natural colors showing the pattern and distribution of the
pigmented areas in the leaf of the mahogany type. The pattern
resembles one of the mottling patterns in castor-bean seed coats, and
possibly may be due to the same cause. It occurs only in the forms
with dark red or purplish-red leaves and stems, and may be regarded
as resulting from the absence of the pattern restriction factor previ-
ously mentioned.
Bloom
Bloom, similar in appearance to that on grapes, in castor beans
consists of a scale-like waxy substance, which, under the microscope,
resembles a piece of cracked frozen ground. It is easily rubbed off.
This covers the whole plant and is especially noticeable on the stems
and fruit capsules. In some varieties, it is produced more freely than
in others. Many forms are known which breed true to its absence.
When it occurs on plants with mahogany stems, a dark purple or
purplish-black effect is produced (see Fig. D, Plate XXIV).
Crosses of bloom X no-bloom give either complete or partial domi-
nance of bloom in Fy. In F2, approximately 3 with bloom: 1 no-
bloom were obtained (actual numbers being 1,108 bloom: 377 no-
bloom, the theoretically expected being 1,113 bloom: 371 no-bloom).
In’F3, seeds from unguarded Fy, plants without bloom produced only
plants without bloom. Seeds from unguarded Fy», plants with bloom
either bred true in F3; or gave 3 bloom: 1 no-bloom families. In one
x
516 BROOKLYN BOTANIC GARDEN MEMOIRS
cross of bloom X no-bloom, the F; plants had bloom, but were only
lightly covered as compared with their ‘“‘bloom”’ parent. In Fo, the
plants approximated a ratio of 3 bloom: 1 no-bloom as usual, though
many of those with bloom were lightly covered as in F}.
Dehiscent and Indehiscent Seed Pods
The seed pods or seed capsules of Ricinus, in most varieties, are
dehiscent, the seeds being thrown out of the mature ripe capsule
with great force. These are known as “‘poppers”’ in regions where
the plant is cultivated commercially. A few varieties have inde-
hiscent capsules, the seed being retained within the pod for several
months. This characteristic considerably reduces the cost of harvest-
ing the beans, as only two or three collections a season are necessary,
whereas ‘“‘popper”’ varieties must be harvested twice a week or more,
to prevent loss. The castor-bean plant matures its seed over a very
long season. The ‘non-popping”’ variety with which the writer
experimented has thin-walled, comparatively brittle capsules, while
all the popping varieties have capsules with thick, leathery walls.
The dehiscent characteristic is probably due to cells similar to those
found in the fern sporangium, which rapidly lose their moisture content
when the capsule matures and contract, thus breaking apart the cap-
sule and expelling the seed.
Crosses of “‘popper’’ with the ‘‘non-popper”’ variety gave all
“poppers’’ in F; and approximately 9 “popper”: 7 “‘non-popper”’ in
F. (the actual figures being 343 pop.: 259 n.-pop., the theoretically
expected being 338.4 pop.: 263 n.-pop.).
In F;, seed from unguarded F, “‘popper”’ segregates gave either all
“poppers,” or ““poppers”’ and “‘non-poppers.”’
Seed from unbagged “‘non-popper’’ F2 segregates produced in most
cases only “non-poppers.”’ The “popper’’ and “non-popper”’ vari-
eties involved in the crosses were very distinct, but the F, populations
were somewhat difficult to classify, as many of the ‘‘non-poppers”’
would, under very favorable conditions, slightly pop. These were
usually thin-walled, brittle capsules, showing, perhaps, that the nature
of the capsular tissue (thick, leathery or thin and brittle) modified the
‘“popping”’ or “non-popping’’ characteristics. On the assumption
that two pairs of characters are concerned in this cross, each of which
is primarily determined by the presence and absence of a single factor,
the results may be interpreted by regarding the ‘‘popper’”’ character
as due to the presence of both the factor for popping (A), and the
factor for thick, leathery capsules (B). In the presence of A and the
absence of B, the capsules would have thin, brittle tissues, but pop
slightly, although not sufficiently to class them as ‘“‘poppers.’’ In the
‘ LT
WHITE: INHERITANCE STUDIES ON CASTOR BEANS SLY,
presence of B and absence of A, the capsules would be thick and
leathery, but non-popping. When both A and B are absent, the
capsules would be thin, brittle, and non-popping. On this provisional
hypothesis, ‘‘non-poppers”’ of the aB class crossed with those of the
Ab class would give all AB or “‘poppers”’ in F; and a 9 : 7 ratio in Fs.
The two types used in the above-recorded crosses would be represented
by the formulae
I
AABB = “poppers,”
aabb = ‘“‘non-poppers.”’
Crosses of these would give a 9:7 F2 ratio, such as that actually
obtained.
Seed-coat Colors
Seed-coat colors in castor beans are white, brownish yellow,
various shades of red, gray, brown, and black. With one exception,
all forms, so far as the writer knows, have seed coats in which the
ground color.is modified by one of several mottling patterns, although
the mottling patterns are inherited, as in garden beans, independently
of the ground color. The nearest approach to a self color in the
writer’s collection is a black-seeded variety having in some cases very
few mottling marks and in others none at all. Efforts have been
made to discover a self-colored white-seeded variety, but so far with
no success.
In crosses, chocolate brown is dominant over black, red, white
and gray. No F» data are available from any crosses excepting those
of red X brownish gray and its reciprocal. The F; is brown on a
gray background. In Fo, segregates of various degrees of redness
appear as a minority. By counting all those F2 segregates with a
red cast, an approximation to a ratio of 3 brownish gray : 1 reddish
gray is obtained. The actual results are 172 non-red : 40 reddish
gray or red, the theoretically expected results being 159 non-reds : 53
red gray. No reds as brilliant as the grand parental type appeared,
showing that more than a single pair of factors is involved.
In Fs, seed from unbagged Fy» light red segregates gave all light
reds in the majority of cases. Seed of the same kind from medium
red F. segregates also bred true. Seed from unbagged brown F:2
segregates gave browns of various shades in some cases in F3, while
in others, browns, reds, and brown grays were produced. Reds as
brilliant as the red ancestor were secured from red F» segregates.
_ At least three types of seed-coat color mottling can be distinguished
definitely in castor-bean seeds. One is coarse-veined, one is fine-
veined, dotted and splotched, while the third is characterized by a
very few large splotches. When the seed-coat is black, the mottling
518 BROOKLYN BOTANIC GARDEN MEMOIRS
is obscured. As in the case of the stem colors; the different types of
mottling appear to involve the presence and absence of several pairs
of restriction factors. Only two of these patterns have been studied
in detail. These are the coarse- and fine-veined types. Crosses
between coarse and fine always give in F, all fine, indistinguishable
from the ‘‘fine’’ pattern parent. In Fs, approximately 3 fines : 1
coarse are obtained, the actual figures being 163 fine : 49 coarse
(theoretical expectation 159 fine : 53 coarse). Unbagged Fy. segre-
gates having coarse mottled seed generally breed true in F3, the few
cases where plants with fine mottled seeds have appeared being un-
doubtedly due to foreign pollen contamination. Unbagged Fe segre-
gates with fine mottled seeds either bred true in F; or gave both fine
and coarse-mottled progeny. Coarse-mottled X the large splotched
type gives a dominance of the former in F;._ No F2: progeny have been
grown.
INTERPRETATION
The inheritance of five of the sets of characters described in pre-
ceding pages—green and red blush stems, red blush and mahogany
stems, red blush and rose red stems, bloom and no-bloom, fine and
coarse seed pattern—so far as the F, and Fy», data are concerned, are
most simply interpreted as due to the presence and absence of a single
genetic factor in each case, making in all five genetic factors. The
inheritance of dehiscent and indehiscent capsules is assumed to involve
primarily two pairs of factors. No evidence of close linkage was found
between any of these seven pairs of factors, although the data were
taken with this end in view.
SEED SHAPE AND DIMENSION
Castor-bean seeds differ as to shape in being oval or orbicular
(about as long as broad). In crosses between varieties breeding true
to the two types, the F;, plants are all oval seed, while in Fe, orbicular
seeds are present in considerably over one fourth of the progeny sug-
gesting a 9 : 7 ratio.
Varieties of Ricinus vary remarkably in their seed dimensions
and weight. Some of the commercial varieties have seeds less than a
centimeter long, which run about 4,550 to a pound of 450 gm., while
the seeds of some of the large Zanzibar ornamental types are over 2.5
cm. long and run only 450 to a pound. Between these are numerous
forms breeding true to almost every gradation in size and weight.
A large number of crosses between these types have been made,
the F, plants showing various degrees of intermediacy. All the
WHITE: INHERITANCE STUDIES ON CASTOR BEANS 519
different F, plants of each were practically uniform as to seed size
(see Plates XXV, XXVI, and XXVII).
Most of the large-seeded forms and some of the very small-seeded
forms (with indeterminate growth period) require a long season to
produce mature seeds, so that even when Fy» populations were started
in the greenhouse in pots several months before planting out, only
about two thirds of the segregates matured seed. One year, attempts
to overcome this difficulty by growing the plants in 10 cm. pots for a
year were unsuccessful. The difference in seed maturity between the
outdoor and these pot-grown plants was very slight. Because of these
difficulties, several of the F2 populations shown in Plate X XVII repre-
sent only part of the segregates—the small- and intermediate-seeded
classes. In F.2 populations from small x large seed or the reciprocal,
small-seeded types similar to the small-seeded grandparent and even
smaller were obtained in every case, while in some of the crosses
involving nearly complete F: populations (Plates XXV, XXVI), the
large-seeded type was also obtained. In all crosses, as expected from
studies of size characters in maize, poultry, and other plants and
animals, numerous intermediates were present, so that a complete Fe
population represented a gradating series ranging from those similar
to or smaller than the small-seeded parent to those similar to the
parent with large seeds.
Seeds from unbagged Fy, small-seeded and large-seeded segregates
have given similar F; progeny, showing the extremes to breed true.
Various F»2 intermediates have also bred true in F3, while other inter-
mediates have given the whole F», series again. Still others have
shown very much less variation.
OTHER CHARACTERS
Numerous characters, other than those described in preceding
pages, have been studied from the standpoint of heredity, but not in
sufficient detail, to admit of interpretation. Crosses between low-
growing (dwarfs), early seed-maturing types with determinate growth,
and tall, late-maturing types with indeterminate growth gave inter-
mediates in Fy, which in F, gave all three types, though accurate
classification so far has been impracticable. Crinkled, much notched
leaved types crossed with ordinary leaved types gave either dominance
of the ordinary type or intermediates in Fy. Some types have a loose,
few-seeded fruiting spike, while others have a dense compact spike
with a larger number of pods. Crosses between them give either
intermediates or dominance of the loose spike. In Fe, both types
reappear, together with many intermediates.
520 BROOKLYN BOTANIC GARDEN MEMOIRS
HETEROZYGOSIS
F, hybrids between several of the forms produced, as in many
maize crosses, a much larger amount of seed than either parent, the
environment being practically the same for all. F, hybrids between
still other forms, however, failed to show this increased productivity.
This is also true of maize F, hybrids.
This increased productiveness in F,; should be, as in the case. of
maize and tomatoes, of great commercial value, since crossing castor-
bean varieties, where no particularly accurate results are desirable,
is very simple and could be done rapidly. Plants of the two types
to be crossed could be grown separately and one lot used entirely as a
pollenizer. A large quantity of pollen from the same spike matures
at the same time. Hence, these spikes could be cut off when nearly
mature and laid on paper sheets till the pollen was shed—a matter
of aday or two. The pollen could then be collected in a powder
gun or similar device and shot over the newly matured pistils each
morning. The male flowers on the plants used as_seed-bearers,
for the most part, can be easily rubbed off without injuring the
flower spike. The amount of selfed seed by this method would be
very small, most of the mature seed being crossed. As ordinarily
planted commercially, one bushel of beans running 1,500 beans to
the pound will plant anywhere from 6 to 40 acres, depending on the
distance apart and the number of beans planted per hill (1-3 beans).
Some of the commercial varieties run as high as 3,000 beans or more
per pound. Castor’ beans produce from 10 to 40 bushels per acre,
depending on the variety, soil, climate and length of frostless season.
With these facts in view, it seems unnecessary to urge the commercial
importance of using F; generation hybrid seed produced by the method
mentioned above. Experiments to determine which varieties crossed
together would give the greatest yields in a given locality should be
made in regions where the beans are grown commercially, since, as
previously stated, there is great variation in varieties as regards this
characteristic. This characteristic of increased productivity in Fy
progeny of certain varieties, combined with such characters as “‘non-
popping”’ and high oil content, should help toward putting castor-bean
growing on a better commercial basis in this country. No experiments
regarding increasing the oil content through “‘selection,’’ so far as
the writer knows, have been made. Varieties with seed yielding 30 to
45 percent oil are said to be already common commercially. Experi-
ments with, and chemical analysis of the innumerable varieties would
possibly give us strains with a much larger oil content.
BROOKLYN BOTANIC GARDEN MEMOIRS, VOLUME I, PLATE XXIII.
WHITE: INHERITANCE STUDIES ON CASTOR BEANS
BROOKLYN BOTANIC GARDEN MEmoirS. VOLUME I, PLATE XXIV.
WHITE: INHERITANCE STUDIES ON CASTOR BEANS
VOLUME I, PLATE XXV.
BROOKLYN BOTANIC GARDEN MEMOIRS.
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INHERITANCE STUDIES ON CASTOR BEANS
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WHITE: INHERITANCE STUDIES ON CASTOR BEANS
BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME I, PLATE XXVII.
WHITE: INHERITANCE STUDIES ON CASTOR BEANS
VOLUME I, PLATE XXVIII.
BROOKLYN BOTANIC GARDEN MEMOIRS.
Ss
STUDIES ON CASTOR BEAN
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~- WHITE: INHERITANCE STUDIES ON CASTOR BEANS 521
re EXPLANATION OF PLATES XXIII-XXVIII
PiaTe XXIII is a microphotograph of a highly magnified portion of leaf
epidermis of the mahogany type showing the red pigment pattern.
“~ PLatE XXIV illustrates the various stem colors. Fig. A is red blush and
green bloom; Fig. B is green; Fig. C is red blush and green without bloom; Fig. D
is mahogany red or dark red-purple with bloom; Fig. E is mahogany or dark red
without bloom; Fig. F is rose or carmine red.
PLATE XXYV illustrates inheritance of seed dimension.