Title: Contributions from the Botanical Laboratory, vol. 8
Place of Publication: Philadelphia

Copyright Date: 1931

Master Negative Storage Number: MNS# PSt SNPaAg242.4

CONTRIBUTIONS

FROM THE

Botanical Laboratory

OF THE

UNIVERSITY OF PENNSYLVANIA

VOLUME Yin

1930-193 1

PHILADELPHIA
1 9 S 1

CONTENTS

CLELAND, RALPH E. The cytology and life history of Nemalion
multifidum, Ag. (Annals of Botany, July, 1919.)

FOGG, J. M., JR. The flora of the Elizabeth Islands, Massa-

chusetts.

I. General discussion. (Rhodora 32:119-132, 147-161, 167-180,

208-221. 1930.)

II. Annotated list of the Vascular Flora. (Rhodora 32:226-258,
263-281. 1930.)

PLOWE, JANET Q. Membranes in the plant cell. (Protoplasma
12:196-240. 1930.)

I. Morphological membranes at protoplasmic surfaces.

II. Localization of differential permeability in the plant
protoplast.

See also under Seifriz, William.

SEIFRIZ, WILLIAM. The Spierer Lens and what it reveals in
cellulose and protoplasm. (Jour. Phys. Chem. 35:118-129. 1931.)

-The structure of protoplasm. (Science 73:648-649. 1931.)

Sc PLOWE, JANET Q. The effects of salts on the exten-

sibility of protoplasm. (Jour. Rheology 2:263-270. 1931.)

STEERE, WILLIAM CAMPBELL. A new and rapid method for
making permanent aceto-carmine smears. (Stain Technology
6:107-111. 1931.)

STYER, J. FRANKLIN. Nutrition of the cultivated mushroom.
(Aiiier. Jour. Bot. 17:983-994. 1930.)

TAYLOR, W. R. A synopsis of the marine algae of Brazil. (Revue
Algologique 5:279-313. 1930.)

WHERRY, EDGAR T. Asplcnium bradleyi erroneously reported on
Limestone again. (Amer. Fern Jour. Vol. 21, No. 3. 1931.)

The Eastern short-stemmed leatherflowers. (Jour. Wash.

Acad. Sci. Vol. 21, No. 9. 1931.)

-The Eastern short-styled Phloxes. (Bartonia, Vol. 2, No.

12. 1930.)

-A new spiral-orchid from the Southern states. (Jour. Wash.

Acad. Sci. Vol. 21, No. 4. 1931.)

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Tl^J.f^NATQRY NOTE. The following contribution is the abstract of a
paper printed in 1918. Through a series of accidents due to the dis-
turbed condition of the mail service at the time, the article that
should have been included in an earlier volume of the Contributions
was omitted. This abstract is now included here with a reference to
the original publication in order in a measure to fill the vacancy in
this record of the Department's work.

October, 1931.

Rodney H. True, Chairman,
Department of Botany

THE CYTOLOGY MID LIFE HISTORY OF !^]L\LION niLTIFIDU?.!, .'^.
(An abstract of a paper by the sane title in Annals of Botany, July, 1919)

Ralph E. Gleland

While the cytological situation in the tetrp.sporic red algae has be-
come fairly well understood in recent years, the seat of chromosome reduc-
tion has not been satisfactorily established in that group which does not
produce tetraspores, and in which no morphological alternation of genera-
tions occurs. The results obtained by V/olf e, '04, on Nemaliou, and by
Svedelius, '15, on Scinaia are contradictory, and the whole group has been
in need of a thorough study. In view of this situation the present invest-
igation has been made.

Cell structure. Nemalion possesses a true jyrenoid, whicn appearo as
a densely staining body at the center of a radiating chronp.tophore. In
cells in which photo synthetic activity is great, the pyrenoid is more prom-
inent than in cells in which this activity is not so great. Furtnernore.
the structure of the pyrenoid is quite different in the two types of cell.s.
In cells near the center of the thallus, in which the amount of photosyn-
thesis is reduced, the pyrenoid consists of a central body or hub, a series
of radiating strands passing out from this hub, to an outer ho llov/ sphere.
In cells near the periphery of the thallus the pvrenoid is solid, the re-
gion of radiating strands being replaced by a solid matrix. The pyrenoid
is probably a metabolic center, concerned with the reproduction of so-called
"Floridean starch".

The nucleus of the cell is much smaller than the chromatophore, or the
pyrenoid, and much less easily observed. Nevertheless, it has been seen in
the division stage, and the number of chromosomes was found to be 8, the
reduced number. The details of mitosis in the vegetative cells are essen-
tially similar to those observed in the cystocarpic filaments (see below).

Spermatogenesis. Antheridial cells are produced in large quantities
at the tips of assimilative filaments. Often a succession of anther ;.dia
may be formed at a single point. The contents of the antheridium slip out
as a naked spermatium, which is carried passively to the trichogyne. vjliile
attached to the trichogyne, the spermatium nucleus divides, and both ol the
resulting nuclei may pass into the trichogyne. A number of spermatia may
become attached to a single trichogyne. While several nuclei may enter the
trichogyne, only one succeeds in entering the carpogonium proper, which is
cut off from the trichogyne by a cleavage plane after entrance of the male
nucleus, ;

Oogenesis. The procarp arises as a lateral shoot from an ordinary veg-
etative branch. From the first it is easily recognised by its broader,
blunter outline. It consists normally of 4 cells, although from as few as
2 to as many as 6 have been observed. The trichogyne of the carpogonium
sometimes shows traces of a nucleus, although this is rare. It is Probable
that the trichogyne was originally an outgrowth of the carpogonium, and that
the trichogyne nucleus represents a second gamete nucleus which has lost its
reproductive function.

Fertilization, This is accomplished rather slowly, so that stages are
readily founl. The fusion of nuclei involves a fusion also of chromatic
nucleoli*

Reduction division. The reduction in the number of chromo somes occurs
with the first division of the zygote nucleus. The reticulum is seen to be
composed of threads which shorten and thicken, become parallel, and finally
break up into 8 bodies, which are the bivalent chromosomes. A character-
istic stage of diakinesis is developed in which the paired chromosomes stand
about in the nucleus, more or less evenly spaced. When the spindle is con-
stituted, it is found to be intranuclear, the nuclear membrane remaining
intact throughout metaphase. Eight chromosomes are separated to each pole.
There is, therefore, a segregation of homologous units ^ and the division is
a true reduction division. The cell is cut across horizontally after this
division, forming an upper sporo^enous and a lower hypogynous cell. The
upper cell gives rise to the gonimoblastic filaments of the cystocarp.

Development of the cystocarp. The divisions which follow the reduction
division have been studied in detail. A reticulum is formed out of sub-
stance contributed by the nucleolus, and this becomes concentrated in eight
regions to form the chromosomes. These are gathered to the equator of an
intranuclear spindle, are split and separated to the poles in the usual man-
ner. All of the divisions from the first to that occurring at the formation
of the first carpo spore have been followed, and these are all found in nor-
mal mitoses, involving the haploid number of chromosomes.

The carpospores are shed by the rupture of the distal wall of the term-
inal cells of the cystocarpic filaments, and the escape of the protoplasts.
Other cells are then buddea into the empty shells thus created^ so that a
series of carpospores may be produced from the tip of each gonimoblastic
filament*

Germination of carpospore. This has been followed closely, and the
mitosis has been observed, as a result of which the germination tube is cut
off from the spore. The haploid number of chromosomes is present in this
division. It is a matter of interest that the nucleus remaining behind in
the nearly empty spore cell, after the papilla has been cut off, occasionally
divides again, although the spore itself is not to survive, and both it and
the nuclei thus formed early degenerate. This supplementary nuclear division
suggests that the reduction division might occur at the germination of the
spore, but it is clear that the chromosome number in Nemalion is already re-
duced before this division occurs.

Discussion. The type of alternation of generation observed in the red
algae is different from that seen in the Archegoniates. In the latter, the
sporophyte is m new structure, interpolated between two gametophyte genera-
tions. It has no phylogenetic ancestry. Its origin took place simultan-
eously with the establishment of the diploid condition as a distinct phase
in the life history. In the red algae^ however, the phase which is chiefly
characterized by the presence of the diploid condition, in plants possessing
alternation of generations, is the tetrasporic plant, which is not a new
structure, but an old structure made over, through a delay in the time of
reduction. There is, however, a structure antithetic with the original gam-
etophytic structure, and that is the cystocarp. This, however, was in exist-
ence before the advent of the diploid phase, as is evidenced by its presence
in forma like Nanalion, in which the diploid phase has not as yet been estab-
lished. While the tetrasporic plant is homologous with the sexual plant,
therefore, in the red algae, the situation is complicated by the presence of
an antithetic structxire, which in those forms lacking alternation of genera-
tions is haploid, but in those having alternation of generations is diploid.
Neither "homologous" nor "antithetic^ can be applied accurately, therefore,
to the type of alternation characteristic of the red algae.

Reprinted from Rhodora, Vol. 32, July to December, 1930

CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY

XCI.

THE FLORA OF THE ELIZABETH ISLANDS,

MASSACHUSETTS

By John M. Fogg, Jr.

Dates of Issue

Pages 119-132 28 June, 1930

" 147-161 28 July, 1930

" 167-180 4 September, 1930

" 208-221 14 October, 1930

" 226-258 3 November, 1930

" 263-281 18 December, 1930

\

CONTRIBUTIONS FROM THE GRAY HERBARIUM OF
HARVARD UNIVERSITY— NO. XCI

THE FLORA OF THE ELIZABETH ISLANDS,

MASSACHUSETTS

John M. Fogg, Jr.
Part I. General Discussion

LOCATION

The Elizabeth Islands are formed by a partly submerged ridge of
morainal hills which extends WSW from Woods Hole, Massachusetts,
for a distance of about 16 miles. This ridge has been separated from
the mainland, as well as divided into islands, in comparatively recent
geological time. To the northwest this chain of islands is washed by
the waters of Buzzards Bay, while along their southeastern and
southern shores runs Vineyard Sound, a channel 4 miles wide which
flows between the Elizabeth Islands and Martha's Vineyard. To-
gether, the Elizabeths and Martha's Vineyard constitute Dukes
County, the former being included in (Josnold Township.

Seven main islands and twelve smaller ones make up the Elizabeth
Islands. Starting at their eastern end the seven principal divisions
are, in order: Nonamesset, Uncatena, Naushon, Pasque, Nasha-
wena, Cuttyhunk and Penikese. All of tliese lie in a more or less
direct line with the exception of Penikese, which is separated from the
main axis of the chain, being just one mile due north of Cuttyhunk.
Between Nonamesset and Naushon lie Monohansett and Buck
Islands, separated by narrow channels or *' gutters" which have been
bridged. To the north, and lying in Hadley Harbor, are Captain's
and Ram's Head Islands, the latter appearing on some maps as

120

Rhodora

[July

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 121

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Puritan Island. To the south of the gutters lie East Buck and West
Buck Islands, although here again confusion exists, as West Buck oc-
casionally appears on maps as " Monohansett. " The three Weepecket
Islands extend northward from the eastern end of Naushon and now
constitute a bird sanctuary. With the mention of Gull Island, a
small strip of sand lying east of Penikese, and Pine Island, imme-
diately to the northeast of Nonamesset, the subject of the minor
islands may be dismissed, for the remaining islets are too snudi to
have received formal names.

HISTORICAL INTEREST OF THE ELIZABETH ISLANDS

A unique historical interest attaches to the Elizabeth Islands
through the fact that upon the outermost of the chain was uuide the
first attempt to establish an English settlement in North America.
On the 4th of June (Old Style, May 25th), 1602, Captain Bartholomew
Gosnold, after having named Cape Cod and Dover Cliff (now Gay
Head), fixed upon the island of Cuttyhunk as the site of a future
settlement and, in honor of his sovereign queen, called it Elizabeth's
Isle, which name has since been applied to the entire group. Here,
upon a tiny islet in a large pond at the west end, the crew of Gosnold 's
ship, the " Concord," constructed a rude fort, and here they lived for a
period of three weeks. This settlement, short-lived though it was,
thus antedates the founding of Jamestown by five years and that of
Plymouth by eighteen years, a fact which was commemorated by the
erection and dedication of a monument to Gosnold on Cuttyhunk
upon the occasion of the tercentenary of the original landfall.^

Cuttyhunk is the only member of the Elizabeth Islands which has
been able to boast a permanent popidation. The little town of
Gosnold, named after its ilhistrious founder, has long existed as a fish-
ing village at the east end of the island and today has about one hun-
dred inhabitants. During the whaling days schooners bound for
New Bedford were accustomed to stop at Cuttyhunk to pick up their
pilots.

Another claim to fame on the part of one of the Elizabeth Islands
may be made for Penikese, the smallest and most desolate member of
the chain. Here, in the summer of 1873, Louis Agassiz founded his
school which, through the generosity of the New York merchant who

1 For further details concerning the history of Gosnold on Cuttyhunk and the exer-
cises which marked the dedication of the monument in 1903, see the Old Dartmouth
Historical Sketches, nos. 1 and 4. New Bedford, Mass. (1903).

IRREGULAR PAGINATION

122

Rhodora

[July

donated the island and funds for the construction of a laboratory,
became known as the Anderson School of Natural History. To this
summer school, the first of its kind in the country, came students from
all over the United States and the roll included names which later
became known as belonging to some of the foremost figures m Ameri-
can biology. Following Agassiz 's death in December, 1873, the school
was contmued for one summer by his son Alexander, but thereafter
was abandoned and the island reverted to the State of Massachusetts
later to be used as a leper colony, from which function it was released

only in 1921. . , .u

The island of Naushon has for several generations been the property
of the Forbes family, various members of which have summer homes
at the east end, near Hadley Harbor. To the Forbeses also belong
Nonamesset, Uncatena and Nashawena and it is only through the
generosity and hospitality of the owners that it has been possible to
carry on the botanical exploration which forms the basis for the
present survey.

PREVIOUS BOTANICAL WORK ON THE ELIZABETH ISLANDS

Mention has been made above of Gosnold's visit to the Elizabeth
Islands in 1602. With Gosnold on that expedition were Gilbert
\rcher and John Brereton, ''gentlemen and historians." The former
has left us a very readable and illuminating account of the voyage.

One of the chief objects which Gosnold had in visiting the New
World was to collect and carry home to England a cargo of native
Sassafras which was then much in demand because of its supposed
medicinal value. According to Archer, Elizabeth's Isle (Cuttyhunk)
was in 1602 overgrown with wood, a fact not without mterest m view
of the present treeless nature of this and of several other islands of the
chain Not only did Gosnold 's party find there the Sassafras which
they sought, but mention is also made of cedar, oak, beech and ash.
The very islet upon which the fort was built is spoken of as cedar-
covered Hills Hope (Penikese) was likewise overgrown with cedar,
and Naushon which was also visited is referred to as being forested, a
character which this island, almost alone of the Elizabeths, has re-
tained in large measure down to the present day.

References to the plant life of the Elizabeth Islands seem to be
lacking for a period of more than 250 years, but we may well suppose
that during that interval extensive deforestation was carried on and

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 123

the islands divested of their original tree growth. Certain it is that
within the memory of no living inhabitant have there been trees on
Cuttyhunk or Penikese, except the few which have been planted by the
hand of man. It is probable that this statement also holds for Pasque
and the greater part of Nonamesset and Uncatena. Nashawena
still has considerable areas which are more or less wooded, and Nau-
shon, as already mentioned, has apparently retained much of its
original forest.

The first published report, known to the writer, on the flora of any
of the Elizabeth Islands dates from the year 1874 and concerns the
island of Penikese. Among the students attracted to Agassiz 's
laboratory in the summer of 1873 was David Starr Jordan, who began
his scientific career with a botanical publication. The task assigned
to Jordan by Agassiz was an enumeration of the plants growing
upon the island and in the waters surrounding it. As the result
of this study Jordan published a list^ of the flora in which he included
not only flowering plants but cryptogams as well. So far as the
lower forms were concerned, since no attempt was made at micro-
scopic examination, only the most obvious species were included.
The list enumerates 83 species of algae, 2 mosses, 1 fern and 113 species
of flowering plants. Although the marine algae were preserved
and the original set is still in existence, the wTiter has it on the author-
ity of Dr. Jordan that no collection of specimens of the higher plants
was made, and we have therefore only the published names as records.
In 1923, on the occasion of the fiftieth anniversary of the founding
of the Anderson School, a botanical survey of Penikese was made by
the staff and students of the Marine Biological Laboratory at Woods
Hole, Massachusetts. The results of this survey were published in
Rhodora for 1924, and, insofar as they indicate the possible direction
of change in the elements of the flora of one of the Elizabeth Islands,
will be referred to later.

Since 1873 nearly a score of botanists have visited the Elizabeth
Islands and brought back specimens which are in one or more of our
eastern herbaria. Since these records have been incorporated into
the catalog which forms the second part of this study, a brief chrono-
logical account of these collectors is here presented.

Walter and C. E. Faxon, in 1873 and 1875 respectively, collected a
few specimens on Nashawena; these sheets are in the Gray Herbarium.

'Jordan, D. S. "The Flora of Penikese. " Am. Nat. vlii. 193 (1874).

124

Rhodora

[JULT

In 1890 a Miss Weir collected on Naushon a few sheets which are
now in the herbarium of the New England Botanical Club
Tn August, 1898. Dr. Arthur HoUick made a trip to the islands w. h
a view to studying the geological formations there presented In
the published account of this survey' the author makes mention o
ome of the plants which he found growing on the various member^ of
rchain! The few specimens which he collected are m the New
York Botanical Garden herbarium.

In S herbarium of the Marine Biological Laboratory at Woods
Hole at several plants collected on Naushon in 1901 Some of ^l'-
bear the name of S. B. Sipe while the others are merely signed K A.
S " Inquiry has shown that these initials refer to Miss Elizabeth A.
Simons (now Mrs. Eldred Jungerich).

ZiZ 1901 there dates the largest collection made on any o the
Eliabeths previous to that which forms the basis of the present re-
!!r In tJeL brary of the Gray Herbarium there is a manuscnpt
fist compiled by M^. Alice R. Northrop of the plants of Nasha-
wena Thl list contains the names of 335 species of flowermg plants
InHeri and therefore constitutes a substantial contribution to our
knowlX of the flora of this island. Mrs. Northrop spent the sum-
Z^ofml and a part of that of 1903 on Nashawena and was thus
admirably situated for making careful botanical observations. Her
SluLs many surprises: species which are either absent or very
locally known from southeastern Massachusetts, some of them con-
stating, indeed, notable extensions in range. To this class of ran-
t£ big such plants as Cyperus erylhrorhizos, Uvul^na perjoha^

Zerdandica, Ranuru^lu. reptaru> and Hydrocotyle Canbyr. Un-
for lately not a single one of these specialties appears to be corrobo-
rated by herbarium material. However, Mrs. Northrop did collect
orle spelens for permanent record, about one-fifth of the narn^
on her^ist being represented by sheets in the collections of the New
York Botanical Garden. And the fact that a few of her most interest-
S records, such as Habenaria blepharighttis, Lipans ileptorch^)
Zs^i fLlaria discolor and Asclepi^ vertieiUata are substantiated
by p n^e^and that others, such as Arisa^ma trphyUurn, Medeola
JrgSZru.. Ranunculi Mphinifolius, Myriophyllum p^nnatum,

.HoUlok. A. A Reconnaissance of the Elizabeth Islands. Cont. Geol. Dept.
Columbia Univ. xi. no. 72 (1901).

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 125

Hydrocotyle Canbyi, Comu^ florida and Trientalu horealis, have been
duplicated by the writer either for Nashawena or other islands of the
^oup, makes it necessary to give definite weight to the plants men-
tioned on Mrs. Northrop 's list.

In the herbarium of the New England Botanical Club is a sheet of
Habenaria orbicidata collected on Naushon by Lillian MacRae in
July, 1904. This is the only specimen bearing the name of this
collector seen by the writer.

A. H. Moore also visited the Elizabeth Islands in 1904 and several
sheets of his collecting from Naushon and Penikese are in the collec-
tions of the New England Botanical Club.

In 1906, J. A. Cushman paid two visits to the islands, collecting on
Nonamesset on July 27, and on Naushon, in company with Max
Morse, on August 25. Specimens from these trips are in the herbar-
ium of the Boston Society of Natural History.

Naushon and Nashawena were visited by E. F. Williams on July
10, 1911. The few plants collected on this occasion are in the New
England Botanical Club herbarium.

In 1911, also, F. W. Pennell made several collecting trips to the
Elizabeths, touching chiefly Nonamesset, Naushon, Nashawena and
Cuttyhunk. Dr. Pennell's specimens, numbering about fifty, were
distributed to the Marine Biological Laboratory and the University
of Pennsylvania.

Thhty sheets from Cuttyhunk, collected by S. N. F. Sanford in
1917, are now in the New England Botanical Club.

Scattered collections were made on various islands by W. R. Taylor
from 1917 to 1921. These specimens, with the exception of a sheet of
Liparis Loeselii which is now in the New England Club, are either at
the Marine Biological Laboratory or at the University of Pennsylvania.
Dr. H. K. Svenson visited Pasque on September 8, 1926 and collect-
ed a few specimens which are now in the herbarium of the New Eng-
land Botanical Club.

On August 10, 1927, Professor M. L. Fernald and the writer visited
Uncatena and Naushon, and the material collected upon that occasion
has been distributed to the New England Botanical Club and the
University of Pennsylvania.

A few specimens were collected by E. W Hervey on Cuttyhunk.
These bear no date and are now in the New England Botanical Club
herbarium.

126

Rhodora

[July

COLLECTIONS MADE DURING COURSE OF PRESENT STUDY

By far the largest number of records accumulated to form the basis
tor the present account of the flora of the Elizabeth Islands represent
collections made by the writer over a period of s.x years. From 1923
t^ 1928 Elusive, each island was visited many times and hundreds
S specimens were collected. This material has been worked over and
Idr^ens have been distributed to the following institutions: Gray
XrClum, New England Botanical Club. University of Pennsylvama
Souri Botanical Garden, Cornell University and Marme Biolog.cal
Laboratory, Woods Hole, Massachusetts. , j , j .„ i

In the Fiftieth Anniversary Survey of Penikese already referred to
the writer contributed the list of Spermatophytes -»-!^d m l^^^^
Subsequent visits have resulted in several additions to that list and
ladeTssible a more careful analysis of conditions on the island and
as stated earlier, such evidences of vegetational changes as are thus
afforded will be dealt with in a later section.

TOPOGRAPHY OF THE ELIZABETH ISLANDS

The general topography of the Elizabeth Islands is that o^^^^^^^^^^
undulating morainal hills with a maximum elevation of about 170
?eet All the features characteristic of typical morainal regions are
here presented, from the rounded hills and depressions, the latter
Xn occLp^^^ by ponds or peaty bogs, to the boulders, some the size
liTs^TLsl which are scattered everywhere. Nowhere except
a o^gThe beache; is there any considerable stretch of fiat land: a walk
across any of the islands necessitates repeated ascent and descent of

''llol?^^^^^^^^ sides of the islands, facing Vineyard Sound, the
shore tLs to be steep and precipitous, often presenting sheer sand
and gravel faces nearly a hundred feet high, rising abruptly from a
narrow cobble beach. In general the highest land is along this south
shore and the ground slopes away gradually to the opposite side of the
Ss where there are frequent coves and low brackish swamps or

'"SieTonds which occur in the hollows in the open, -n^ed ^^^^^^^^
inLe'cases, merely small pools which may form desiccated bog^^^^^
or disappear entirely in very dry seasons, or, m other cases, are

.The Flora of Penikese. Fifty Years Aft^r. Edited by I. F. Ix^wis. Rhooohx.
xxvi. 181-196. 211-219, 222-229 (1924).

1930] Fogg,— Flora of the Elizabeth Islands, Massachsuetts 127

sufficiently large to have been designated as lakes. The largest of
these latter, West End Pond on Naushon, is more than a quarter of a
mile in diameter.

The character of the beaches varies from those of the nigged
boulder type, liberally bestrewn with huge rounded stones, to tliose
which offer a smooth sandy shelf. The characteristic type lies be-
tween these extremes, and we find for the most part a shingle or cobble
beach with here and there piles of boulders and flat sandy patches.
Dunes of shifting sand are rare and occur extensively only at the east
end of Nashawena, and along the north shore of Naushon, west of
Kettle Cove.

In those islands which have been deprived of their trees the open
barren hills are covered with grasses, or other low growth, while the
dry hollows or protected lee slopes harbor dense patches of scrub
vegetation, made up mostly of Myrica carolinicnsis and species of
Gaylussacia or Vaccinium. Occasionally an extensive boggy hollow
will be densely wooded, the tops of the trees {Nyssa syhatica, Qucrcus
velutina, Acer rubrum, etc.) conforming to the height and contour of
the surrounding slopes.

The most conspicuous vegetational feature of the islands, aside from
the open grassy downs, is the dense growth of rather low beech
woods which clothes the greater part of Naushon and smaller areas on
some of the other islands. From a distance these woods are seen to
fit in closely with the general topography, due, doubtless, to the high
wind velocity which would tend to level forest growth to the existing
lines of the hills and ridges.

CHARACTERISTIC FEATURES OF THE SEPARATE ISLANDS

As the various members of the Elizabeth Islands exhibit some
diversity as regards general topographic and vegetational features,
and also in the influence which man has exerted upon the latter, a
brief description of each of the seven main divisions is here given.

NoNAMESSET. This island, the easternmost of the chain, is roughly
oblong with a length of 13^ miles and a greatest width of 3^ mile. Its
longitudinal axis lies east and west. The eastern three-quarters of
Nonamesset is essentially treeless, save for an occasional wooded depres-
sion, while the western quarter is heavily clothed with mixed beech
and oak woods. The rather sharp line between tliese two areas, a line
which follows, for the most part, an old stone wall, indicates that the

128

Rhodora

[July

treeless nature of the eastern portion is due to artificial denudation,
it probably having been cleared for purposes of agriculture or grazing.
Altogether there are about 15 ponds of more or less permanent char-
acter on Nonamesset and all but one of these lie in the exposed eastern
portion. Munsod Pond, as it appears on the charts, has now become
merely an arm of Lackey's Bay and the narrow bar which formerly
protected it has been submerged. Nonamesset is connected with
Naushon by three bridges which cross the narrow gates or "gutters"
separating Monohansett and Buck Islands. The East and West
Gutters are open and the rapid current flows through them as through
a mill race, but the Middle Gutter is " blind, " being closed by the stone
wall that forms the bridge.

Uncatena is a triangular-shaped island about J^ of a mile long and
Yl a mile wide. It lies to the west of Nonamesset across Hadley
Harbor and forms a wedge that juts northward into Buzzards Bay.
Uncatena is practically treeless, except for a small natural area in the
extreme southern corner, and here again the assumption is that active
deforestation has been carried on, for it was from the originally
densely wooded nature of Nonamesset and Uncatena that Woods Hole
is said to have derived its name. There are about a dozen small
fresh water ponds on Uncatena during the course of a moderately
rainy summer and a large inland brackish pond drains eastward into
Hadley Harbor by a sluice-way which makes of the northeastern part
of the island a peninsula. Dry, exposed, undulating grassland
characterizes the vegetation of nine-tenths of this island, with an in-
crease of scrubby thicket to the westward. The summer home of
Mr. Malcolm Forbes is located on the east side of Uncatena and a
large central area is fenced off for grazing. This island connects by a
bridge across the Northwest Gutter with Naushon.

Naushon is the largest of the Elizabeth Islands. It is 6H miles
long and averages 1 mile wide, with a width of about Ij^ miles at its
widest point. It extends from Hadley Harbor almost due southwest
to Robinson's Hole which separates it from Pasque. By far the
greater part (perhaps ^) of the surface of Naushon is covered with a
dense growth of trees. In some regions, like the area near French
Watering Place, these woods present an almost pure stand of beech,
in others there is considerable admixture of oak, hickory, hop
hornbeam, maple and black gum. Almost the only portions of Nau-
shon which are not wooded are those right along the shore or some of

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 129

the higher exposed ridges in the central part of the island. Along
the north shore, on one of these treeless stretches, Scotch Broom
{Cyiism scoparius) was introduced some years ago and has taken hold
so vigorously that it now solidly occupies an area of several acres. In
similar spots along this same shore, various conifers (notably larch,
Scotch pine and several spruces) have been set out as a windbreak
and these appear to be no more than holding their own. Naushon has
many ponds, nearly all of them fresh. West End Pond, Marv's
Lake and French Watering Place are the three largest, in the order
named, and there are about a score of smaller ones, depending upon
the degree of rainfall. The island is indented by two large coves,
Tarpaulin Cove and Kettle Cove, which are nearly across from each
other on opposite shores. It has been suggested tluit it was by the
approximation of similar indentations that Nashawena was separated
from Pasque in comparatively recent times and that the same process
may be going on here in Naushon. The gravel cliffs which in many
places form the south shore of the island are high and very precipitous.
East of Tarpaulin Cove they are usually barren on their crests, while
westward they are frequently wooded right to the very edge. Nau-
shon presents several areas of low brackish marshland, the most
extensive being near Job's Neck and at the west end near West End
Pond. At the east end of the island are the several residences of
members of the Forbes family. Here, also are tracts under cultivation
and scattered farmhouses and outhouses.

Pasque. Lying across Robinsons Hole from Naushon is the island
of Pasque, or Peskinese, as it was formerly called. Pasque is roughly
oval in form, with a long axis, lying east and west, of Ij^ miles, and a
short one about 1 mile. It is almost entirely destitute of trees, save
for a few protected hollows. The extreme eastern end is low and
marshy, and is drained by a sinuous tidal stream. Numerous fresh
water ponds are scattered around the rim of the island, for the central
part is high and arid. As on the other islands, the depressions among
the hills near the shore are often peaty and boggy and several ex-
tensive areas of this sort are to be found here. Pasque was formerly
the property of a fishing club. As evidences of this are a large frame
club-house, a landing wharf, a truck patch and outhouses, including
an ice-house on the edge of a small pond ; all at the eastern end of the
island. The ownership of Pasque has just recently passed into private
hands. Between Pasque and Nashawena lies Quicks Hole.

130

Rhodora

[July

Nashawena. The second largest island of the group is Nashawena
(formerly Nashuina), or "Little Naushon," with a length of 3 miles
and an average width of about a mile. Nashawena contains large
wooded areas, chiefly toward the east end, although in the troughs
between the long ridges of hills that traverse the island longitudinally
trees may be found, except in the western quarter. The shores and
the extreme western portion are barren and open, as are likewise the
higher hills throughout. Behind the line of dune hills, which forms
most of the eastern border of the island, lie two large ponds of fresh
water separated by a low swampy area: these are known as the
dune ponds. '^ The easternmost of these ponds is separated from the
sound by a low barrier beach, and evidence mdicates that inundations
of salt water probably occur occasionally durmg the severe storms of
winter. It seems also likely that these two ponds have from time to
time been connected as a single body of water. Here again, as on
Naushon, the highest land is along the southern side of the island
and the slope is to the north. Many of the hollows, especially around
the margin of the island, harbor small ponds, and in addition to the
dune ponds there are several good sized bodies of water the largest
of which is Choptauk Lake, two-thirds of the way to the west end
There are numerous swampy and boggy areas on Nashawena most
of them occurring in the central parts of the island m the wide de-
pressions between the lines of hills. In many cases these swampy
hollows are overgrown by tangles and thickets so dense as to be ^di
nigh impenetrable. It is supposed that, at one time, much of the
open land on Nashawena was under cultivation. The old stone farm
house, said to have been built in 1725, still stands, although a modem
wing has been added to it in recent years. An obscure gravestone
bears the date 1736. Today the island belongs to the Forbes famJy
and is used chiefly for the raising of sheep, of which there are thought
to be about 700. The caretaker and his wife, Captam and Mrs.
Mark Jamison, occupy the new wing of the old [armhouse, and to
their kind hospitality the writer owes a very enjoyable and botanically
profitable visit to the island in July, 1928.

CuTTYHUNK West of Nashawena, and separated from it by a very
narrow passage, Canapitsit Channel, lies the island which Gosnold in
1602 named Elizabeth's Isle but which has reverted, with some
modification, to the old Indian name of Cuttyhunk. Roughly oblong
in shape, the main body of Cuttyhunk is some 2 miles long, with a

1930] Fogg. — Flora of the Elizabeth Islands, Massachusetts 131

greatest width of about ^ of a mile, and lies along a NE-SW axis.
From its southeastern corner a narrow sand-spit runs due east for ^
of a mile toward Nashawena. On this strip is located U. S. Coast
Guard Station No. 50. The northeastern quarter of the island is
occupied by a large body of salt water known as Cuttyhunk Pond.
To the east this is dredged to Cuttyhunk Harbor and thus offers a
land-locked refuge to small vessels. West of Cuttyhunk Pond is the
little village of Gosnold, spread out upon the sheltered east-facing
slope which leads down toward the wharf. All of the central and
western portions of the island are open grassy downs, exposed to the
full blast of winds from the Atlantic. From the lookout station on
the highest hill, near the center of the island, the land slopes gently
away to the south and southwest. The low-lying western part of the
island is occupied by two bodies of water. One of these is fresh and
furnishes the supply of ice for the inhabitants; it has long been known
to visiting botanists as Ice-house Pond or Sheep Pond. The other is
the pond made known to fame by Gosnold and is variously termed
Gosnold Pond or West End Pond. Although originally mentioned
as a fresh-water pond, and still referred to as such, this body of water,
which is separated from the open sea by a very narrow cobble barrier
beach, is certainly frequently immdated. In 1927, Potaviogeton
bupleuroides and Ruppia maritlma, var. longipes, both reliable indica-
tors of a brackish medium, were found growing in it and in 1928
various species of marine algae were collected along its northern
shores. Toward the western end of this pond is the tiny island
upon which stands the monument to Gosnold already mention-
ed, an unpretentious structure of rough native stone. Not far from
this, but on the extreme western edge of the main island, stands the
Cuttyhunk lighthouse. Several smaller ponds and numerous boggy
hollows, some of them rather extensive, are scattered over the western
and southern portion of the island, but these tend to disappear late in
summer when the rainfall is less abimdant.

Penikese. The smallest of the main divisions of the Elizabeth
Islands lies a mile to the north of Cuttyhunk. Known also to Gos-
nold as a cedar covered isle, and called by him Hills Hope, this island
likewise goes today by its Indian name and is, if possible, even more
barren and treeless than Cuttyhimk. Penikese is about 3^ of a mile
long and J^ a mile wide, with a broadly spatulate peninsula rimning
eastward for 3^ mile from its northern end. Its contour resembles

132

Rhodora

[July

that of the other islands, the highest point, however, being only about
70 feet. Its few trees are mostly the result of an attempt at a planting
made around the building which, in the days of the leper colony,
served as the home of the resident physician, although several scrub
willows grow in one of the hollows around the margin of a small pond
on the east side. Of the former luxuriant forest growth Dr. Jordan,
writing in 1874, says, " there is now no trace left save the rotten roots
of a solitary beech stump and a few branches of red cedar and red
maple (?) found buried in the muck of a small swamp. " The status
of the ponds on Penikese appears to be even more precarious than of
those on the other islands. In favorable seasons six small ponds, two
of them brackish, may be found; during a dry summer the number has
been known to be reduced to half. Penikese, then, is dominated by
open, grassy downs with the exception of the narrow cobbly strip
which connects the two portions of the island. That part of the open
hillsides forming the northern slope of the island has been taken over
by the terns (common and roseate), thousands of which here find their
breeding ground. In fact, now that the island has reverted to its
wilder state, these birds show a tendency to usurp it altogether. It is
extremely difficult to walk across any of the grassland areas during
the nesting season without stepping upon the eggs or the young birds.
The handsome stone residence building, on the east side near the land-
ing, has been partly demolished, leaving only a portion of the structure
to house the caretaker who is still stationed there. The frame cottages
on the west side of the island, formerly occupied by the unfortunate
lepers, were destroyed in 1927, a single concrete structure being all
that remains. Thb, and the tiny graveyard at the extreme north end
of the island, bear mute testimony to the use to which Penikese was
put from 1905 to 1921. The remains of an old wooden reservoir cap
the highest hill on the island, while, set in a large boulder near by, is
a bronze tablet placed there in 1923 to commemorate the fiftieth
anniversary of the founding of the Anderson School of Natural History
by Jean Louis Rodolphe Agassiz in 1873.

{To he conlinued)

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 147

THE FLORA OF THE ELIZABETH ISLANDS,

MASSACHUSETTS

John M. Fogg, Jr.

(Continued from page 1S2)
HABITATS

Despite their almost uniformly bleak and arid nature, the Elizabeth
Islands offer a considerable diversity of habitats and the chief of these,
together with a few of the most characteristic plants of each, are here
described.

Beaches. These may be of boulders, cobbles or pure white sand.
In the first two cases, few plants may be sought for, although on the
shingle or cobble beaches Ammophila brcviligulata sometimes manages
to get a foothold in the loose stones. The sandy beaches, however,
offer a habitat for certain characteristic species, of which the following
may be noted :

Ammophila brcviligulata
Triplasis purpurea
Polygonum glaucum
Atriplex patula, var. hastata
Salsola Kali

Arenaria peploides, var. robusta
Cakile edentula

Lathyrus maritimus
Euphorbia polygonifolia
Convolvulus sepium, var. pubescens
Solanum nigrum
Solidago sempervirens
Xanthium echinatum
Sonchus oleraceus

On some of the beaches dead Eel Grass, Zostera marina, has been
piled up by the waves, forming dense mats often two or three feet
thick.

Salt Marsh. There are no extensive salt marshes on the islands,
but a few restricted areas of this nature do occur, such as those along
the southwestern shore of Nonamesset, the eastern side of Uncatena,
the northeastern end of Naushon facing Lackeys Bay, the regions on
the north shore at the west end of the same island, the southeastern
corner of Basque and the extreme west end of Nashawena. Other
similar patches, scattered throughout, are too limited or local to
merit enumeration. As typical of these low, brackish marshlands,
or the slightly elevated peaty areas bordering them, may be cited:

Typha angustifolia
Andropogon glomeratus
Echinochloa Walter!
Spartina alterniflora, var. pilosa
S. patens
Distichlis spicata

Cyperus ferax

C. strigosus

Eleocharis parvula

E. uniglumis, var. halophila

E. rostellata

Scirpus Olne3d

IRREGULAR PAGINATION

148 Rhodora [August

S. validus

S. campestris, var. paludosus

Carex hormathodes

Juncus buf onius

J. Gerardi

J. articulatus, var. obtusatus

Rumex maritimus, var. fueginus

On the brackish mud flats, like those bordering Cutty hunk Pond,
occur such characteristic things as :

Salicornia europaea
Suaeda linearis
Spergularia leiosperma
Hibiscus Moscheutos
Ptilimnium capillaceum
Pluchea camphorata

Suaeda linearis
Spergularia leiosperma
Plantago oliganthos

Puccinellia paupercula, var.

alaskana
Salicornia europaea
S. ambigua

Brackish Ponds. Either in the salt marsh areas themselves, or
near the shore and separated from the sea only by narrow shingle
barriers, occur several brackish ponds, in which may be found such
plants as :

Potamogeton bupleuroides
P. pectinatus

Ruppia maritima, var. subcapitata
R. maritima, var. longipes

Fresh Water Ponds. There are at least 65 fresh water ponds of a
more or less permanent nature on the Elizabeth Islands. Of these,
some are mere pools 20 or 30 yards across in low hollows or kettle-holes,
while others, such as West End Pond on Naushon, may have a width
of nearly one-third of a mile. With this wide difference in size there
goes a corresponding diversity of pond bottoms and the accompanying
floras. Some of the ponds have grassy bottoms and merely represent
hollows which have become filled with rain water; others have a sandy
or cobbly bottom; while the bottoms of still others are formed of a
thick grayish clay. As representative of the plants, floating or sub-
merged, of these fresh water ponds may be mentioned:

Elatine minima
Ludvigia palustris
Myriophyllum scabratum
M. humile

Potamogeton Oakesianus
P. pulcher
P. diversifoUus
Vallisneria americana
Glyceria acutiflora
Lemna minor

Nymphozanthos variegatus
Nymphaea odorata
Ranunculus delphinif olius
Callitriche heterophylla
C. palustris

M. tenellum
Proserpinaca palustris
Hydrocotyle umbellata
H. Canbyi
H. verticillata
Nymphoides lacunosum
Utricularia geminiscapa

Pond Margins. The margins of the fresh water ponds support, in

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 149

most cases, a rather characteristic flora, the components of which de-
pend usually upon the nature of the shore.

Around those ponds with a pure sandy border the following plants
may be listed as fairly typical :

Cyperus dentatus
Scirpus americanus
Mariscus mariscoides
Juncus pelocarpus
J. militaris
J. marginatus

Ranunculus Cymbalaria
Potentilla pacifica
Samolus floribundus
Limosella subulata
Ilysanthes inaequalis
Coreopsis rosea

Surrounding those ponds, however, which offer a peaty or boggy
border, a somewhat different series may be cited as representative:

Thelypteris palustris, var.

pubescens
Sagittaria latif olia
Sparganium americanum
Glyceria obtusa
G. pallida

Eleocharis acicularis
Scirpus cyperinus
Rynchospora alba
R. capitellata
Carex lurida
Eriocaulon septangulare
Xyris caroliniana
Juncus canadensis
J. acuminatus

Iris versicolor
Habenaria lacera
Drosera rotundifolia
D. intermedia
Spiraea tomentosa
Polygala cruciata
Hypericum boreale
H. virginicum
Viola lanceolata
Rhexia virginica
Scutellaria epilobiifolia
Lycopus uniflorus
Gratiola aurea
Bidens connata

It is not suggested that the two classes of plants above presented
be taken as mutually exclusive. In general, however, these species
exhibit a marked preference for the habitat under which they are
listed.

Swamps. Swampy areas occur near some of the larger ponds,
notably west of French Watering Place on Naushon and around
portions of the dune ponds on Nashawena. In addition, most of the
islands boast one or more swampy hollows and Naushon and Nasha-
wena each has several rather extensive swamps in low-lying depres-
sions near the shore. A few of the characteristic plants of this type
of habitat may be noted :

Typha latif olia
Spartina Michauxiana
Phragmites communis
Dulichium arundinaceum
Scirpus validus
S. cyperinus

Juncus efTusus, var. costulatus
Spiraea tomentosa
Rosa palustris
Impatiens biflora
Decodon verticillatus
Sium suave

Cephalanthus occidentalis
Eupatorium verticillatum

150 Rhodora [August

Asclepias incarnata, var. pulchra
Lysimachia terrestris

Bogs. In addition to the restricted boggy areas in the moist
hollows and those forming pond margins, there are several rather
extensive bogs of a permanent character. Chief among these may be
mentioned one chain of bogs at the east end of Pasque and another
series at the west end of Cuttyhunk. Many of the plants which
occur around the peaty borders of the small ponds grow also in these
open bogs, but certain other species reach their fullest development
only in the more extensive areas. The following is but a partial list
of some of the more conspicuous of these bog plants:

Woodwardia areolata
Thelypteris palustris, var.

pubescens
Osmunda regalis, var.

spectabilis
Lycopodium inundatum, var.

Bigelovii
Panicum longifolium
Glyceria obtusa
Eriophorum virginicum
Rynchospora f usea
Carex cephalantha
C. Howei

C. canescens, var. disjuncta
C. virescens
C. Umosa

Eriocaulon septangulare
Xyris caroliniana

Grassland. As already indicated, the larger part of the surface
of the Elizabeth Islands is dominated by open, undulating grassland.
The following list, incomplete though it is, will convey more adequate-
ly than could any description an impression of the character of these
bleak grassy downs:

Juncus effusus, var. costulatus

Habenaria clavellata

H. lacera

Pogonia ophioglossoides

Calopogon pulchellus

Drosera rotundifolia

D. intermedia

Rubus hispidus

Polygala cruciata

Viola lanceolata

Epilobium palustre, var. monticola

Clethra alnifolia

Rhododendron viscosum

Chamaedaphne calyculata

Vaccinium macrocarpon

Bartonia virginica

Menyanthes trifoliata, var. minor

Dennstaedtia punctilobula
Andropogon scoparius, var.

frequens
Paspalum pubescens
Panicum virgatum, var. spissum
P. depauperatum
P. Lindheimeri, var. fasciculatum
P. meridionale
P. oricola
P. sphaerocarpon
Anthoxanthum odoratum

Aristida purpurascens
Phleum pratense
Holcus lanatus
Poa pratensis
Festuca rubra
Agropyron repens
Cyperus filiculmis, var.

macilentus
Carex albolutescens
C. silicea
C. Muhlenbergii

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 151

C. Swanii
C. varia
Juncus tenuis
J. Greenei

Sisyrinchium angustifolium
Spiranthes gracilis
Rumex Acetosella
Spergularia rubra
Stellaria graminea
Cerastium vulgatum
Ranunculus acris
Lepidium virginicum
Potentilla pumila
P. argentea
Trif olium arvense
Polygala polygama
Euphorbia maculata
Hypericum perforatum
HeJjanthemum canadense
H. dumosum
H. Bicknellii
Hudsonia tomentosa
Lechea maritima

Viola fimbriatula

Daucus Carota

Trichostema dichotomum

Linaria canadensis

Eupatorium hyssopif olium

Chrysopsis falcata

Solidago suaveolens

S. nemoralis

S. graminifolia

S. tenuifolia

Aster patens

A. linariifohus

Sericocarpus asteroides

Antennaria neglecta

Anaphalis margaritacea

Gnaphalium obtusifolium

Rudbeckia hirta

Achillea Millefolium

Chrysanthemum Leucanthemum,

var. pinnatifidum
Krigia virginica
Leontodon autumnalis
Hieracium Gronovii

Scrub Growth. Under the protected lee of the hills, in dry
sheltered hollows or bordering the woods, where they form a transition
zone between the grassland and the woodland, occur open patches or
dense, scrubby thickets of low shrubs, of which the following may be
designated as characteristic:

Myrica Gale
M. caroliniensis
Betula populif olia
Pyrus arbutifolia
Amelanchier oblongifolia
Rubus Andrewsianus
Rosa virginiana
Prunus serotina
P. maritima
Rhus copallina

Ilex verticillata
Clethra alnifolia
Rhododendron viscosum
Leucothoe racemosa
Lyonia ligustrina
Gaylussacia frondosa
G. baccata

Vaccinium corymbosum
V. atrococcum
Viburnum dentatum

Woodland. In certain areas, especially near the center of Nau-
shon, tlie native woods are made up of almost pure stands of beech,
Fagiis grandifolia. These trees grow nowhere very tall, averaging,
perhaps, 30-40 feet, and their low, flat, leafy crowns meet overhead,
forming a thick roof through which a subdued light filters. This
climax beech forest may also be seen on a somewhat reduced scale in
portions of Nonamesset and Nashawena. Usually, however, the
wooded areas, although they may be dominated by beech, contain a

152

Rhodora

[August

liberal sprinkling of certain other species, most prevalent among which

are:

Sassafras officinale
Acer rubrum
Nyssa sylvatica

Ostrya virginiana
Quercus alba
Q. velutina
Hamamelis virginiana

In addition to these important constituents of the densely forested
portions, a few other trees occur scattered here and there, seldom
entering conspicuously into the formation of the heavy woods. As
such may be named :

Pinus rigida Prunus serotina

Chamaecyparis thyoides Rhus Vernix

Juniperus virginiana Hex opaca

Carya alba Cornus florida

Mention has already been made of the efforts which were carried on
to introduce certain trees either as a windbreak or for ornamental
purposes. Some of these, such as white poplar, ailanthus and catalpa,
have taken hold and are spreading, while others apparently just
manage to survive. A partial list of these introductions follows:

Pinus sylvestris
Larix decidua
Picea Abies
P. glauca
P. pungens
SaUx alba
Populus alba

The herbaceous flora which enjoys the protection of the native
woodland of the islands is for the most part a rather meagre one. The
dry, leaf-covered floor of the pure beech woods is almost uniformly
sterile, so far as vascular plants are concerned, although such an
habitat presents a rich and varied mycological flora, especially follow-
ing a heavy rain. In the more open mixed woods, however, several
characteristic species inhabit the shaded knolls. As examples may be

cited :

Pteridium aquilinum, var. Sanicula canadensis

latiusculum Monotropa uniflora

Thelypteris noveboracensis Epif agus virginiana

Panicum dichotomum Galium pilosum
Carex cephalophora

Certain of the low depressions or hollows in the woodland areas are
swampy and, in addition to high-bush blueberries {Vacciniwn corym-
bosum and V, atrococcum) , may harbor such plants as:

Betula pubescens
B. pendula
Gleditsia triacanthos
Robinia Pseudo-Acacia
Ailanthus glandulosa
Catalpa bignonioides

1930]

Pogg —Flora of the Elizabeth Islands, Massachusetts 153

Sparganium eurycarpum
S. americanum
Sagittaria latif olia
Glyceria striata
Carex lupulina

Juncus effusus, var. solutus
Decodon verticillatus
Siam suave
Lycopus uniflorus
Erechtites hieracifolia

Other similar depressions are moss-covered and boggy and in such
situations may be found:

Carex Howei

C. canescens, var. disjuiicta

C. brunnescens, var. sphaero-

stachya
Arisaema triphyllum

Oakesia sessilifolia
Maianthemum canadense
Medeola virginiana
Trientalis borealis

CHANGES IN THE FLORA OF THE ELIZABETH ISLANDS

Here it is proposed to indicate the possible direction and nature of
the changes in the flora of these islands. It has seemed advisable to
put on record certain facts which illustrate what has already taken
place in this respect and to point out others which may be of interest
to the future student of the islands in interpreting further changes.

The original wooded nat\ire of all of the Elizabeth Islands has
previously been alluded to, as has also the fact that the present treeless
condition of some members of the chain is apparently the direct
result of cutting by man. Right here the (juestion may very well be
asked, "Why have the islands thus denuded never regained their
fores/ growth?" In attempting to solve this problem two chief
factors must be taken into account and their relative importance

weighed.

In the first place, sheep have been raised more or less extensively on
the islands from time to time and the effects of these browsing animals
in cropping off the young vegetation must not be lost sight of. Despite
the numbers and activities of the sheep, however, they have not sue-
ceeded in keeping down completely the herbaceous growth in those
areas which they inhabit. Even on Xashawena, where their numbers
are greatest, the open grassy downs where they graze boast a large
number of species of grasses as well as other plants and one has no
difficulty in collecting perfect and unmutilated specimens of any plant
which he desires. While evidences of grazing are certainly not absent,
the region in general does not present the devastated appearance which
so often results where slieep have been allowed to nm wild; and the
fact that so many herbaceous and shrubby plants are able to make a
showing, especially in the protected hollows, would seem to indicate

154

Rhodora

[August

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 155

that it can scarcely have been the sheep alone which kept back the
developing growth so severely that the trees were unable to regain
their foothold.

The second factor which deserves serious consideration is a geologic
one. In an important paper on coastal subsidence in 1893,^ Dr.
Arthur Hollick called attention to the fact, already well established,
that the era of elevation which was active along the eastern borders of
the North American continent in late Tertiary times resulted in an
uplifted coastal plain, the eastern limits of which probably coincided
with the present 100-fathom contour (about 100 miles from shore).
This elevation is supposed to have reached its maximum shortly after
the advent of the Ice Age. Then, either previous to, or subsequent to
the period of greatest ice accumulation, an era of depression set in.
The rate of subsidence has been roughly calculated and Hollick
supposes that 6000 years ago the area included within the present 20-
fathom line would have been dry land.

That is, not only would the Elizabeth Islands, together with Long
Island, Block Island, Martha 's Vineyard and Nantucket, have formed
a portion of a continuous land surface, but they would have been some
miles inland from the actual coast line. The nearest approach of the
20-fathom line to Cuttyhunk is at a point almost due south, where it is
now about 20 miles (32 kilometers) out from the shore. To the south-
east this distance increases to about 90 miles (146 kilometers) as the
submerged contour swings out to sea to conform roughly to the out-
lines of Nantucket.

The part which this post-Pleistocene land shelf may have played
in the migration of plants to the Elizabeth Islands will be a matter for
consideration in the section of the Origin of the Flora which follows.
The point requiring present emphasis is this: if all the islands were
heavily wooded at a time when perhaps their inland location afforded
them some protection, it seems highly probable that later, when they
assumed their present position, the severe maritime conditions then
prevailing would be such as to discourage nature's attempts at re-
forestation, once the original woodland were removed. This would
suggest that those islands which have never been disturbed, such as
Naushon, have merely retained an original forest growth, the posses-
sion of which they owe to conditions previously more advantageous

» Hollick, A. Plant Distribution as a Factor in the Interpretation of Geological
Phenomena, with Special Reference to Long Island and Vicinity. Trans. New York
Acad. Sci. xii. 189-202 (1893).

'

than obtain at present, while those less fortunate have suffered
through their comparatively recent exposure to the unmitigated
forces of the Atlantic. That excessively high wind velocity is an
effective factor in retarding tree growth is nowhere better shown than
on the eastern side of Nantucket with its extensive scrub oak barrens.
This is further borne out on the Elizabeths by the fact that in the
open, unprotected areas scrub growth forms only in the more or less
sheltered hollows and the occasional isolated sapling which does get a
start elsewhere remains dwarfed and stunted.

It is true that on Martha's Vineyard the woods along certain sec-
tions of the north shore have been cut for their timber more than once,
and that new growth has been quickly made. But this slope enjoys
the protection of the high line of morainal hills, averaging 200-300
feet, which shelter it from the winds of the open sea. No such protec-
tion exists on the Elizabeth Islands and the lack of it, rather than the
presence of sheep, appears to be the determining factor in the failure
of natural reforestation. In the light of these facts, it would seem
futile to hope that the devastated areas can ever regain their former
wooded luxuriance.

Another phase of vegetational change which it seems worth while
putting on record is the behavior of certain introduced species on the
Elizabeth Islands.

Reference has already been made to the planting of Scotch Broom,
Cytisus scoparius, on Naushon. It is interesting to note that, in the
account of his reconnaissance made between August 10 and 16, 1898,
Dr. Hollick says of this species that it was "planted over extensive
areas" on Naushon but that it "did not appear to be in a very thriving
condition."^ Today Cytisus occupies solidly an area of many acres
along the north shore of the island, near Kettle Cove. On the 10th
of August, 1927, Professor Femald and the writer visited this locality
and had the unique experience of wandering through this exotic
plantation. The plants grow very close together, and are often 6-8
feet tall, and the tendency in attempting to traverse the area is to
lose completely one 's sense of direction. Unless checked in some way,
Cytisus bids fair to encroach even further upon the surrounding region
and to usurp in time a much larger area than that which it now domi-
nates. Although introduced also on Pasque and Nashawena, Scotch
Broom has nowhere else made the showing that it has upon Naushon.

1 Hollick, A. Cent. Geol. Dept. Columbia Univ. zi. no. 72. 391 (1901).

156

Rhodora

[August

f'* -«

Another leguminous plant that has been successful in establishing
itself is the Woad-waxen, Genista tinctoria. This species was intro-
duced at the extreme east end of Naushon, near Hadley Harbor. It
now occupies almost solidly a large field in this vicinity and occurs
scattered elsewhere over the open hillsides here as well as on Uncatena.
At several places the Tree of Heaven, Ailanthus glandulosa, has
become thoroughly naturalized and appears to be spreading rapidly.
This is especially true on Naushon, north of Tarpaulin Cove, where, in
at least one protected hollow, this tree has formed an extensive and
almost impenetrable thicket.

There remains to be considered in this connection such light as is
thrown on the nature of vegetational changes by an examination of
David Starr Jordan's account of the flora of Penikese as he found it
in 1873.^ The author states it as his hope that his list may have an
interest for future botanists, especially "as showing which plants
survive a prolonged struggle for existence against grass and sheep. "
And as this is the first published list of the flora of any of the Elizabeth
Islands, it forms our chief basis for a study of those changes which may
have occurred over a considerable period of years.

In the paper entitled " The Flora of Penikese, Fifty Years After,* '
which has already been mentioned. Dr. I. F. Lewis summarizes the
numerical differences berween Jordan 's list and that compiled as the
result of the survey made in 1923.^ It is not intended to duplicate
that summary here, but subsequent exploration by the present writer
has yielded so many additional records, and a closer scrutiny of the
terminology employed in the earlier list has resulted in a so much
.better understanding of the discrepancies involved, that it seems well
to consider, as briefly as possible, just how much significance, if any,
attaches to the marked difference in the superficial aspects of the two

lists.

The list for 1873, compiled by Dr. Jordan, contains 1 fern and 113
flowering plants, whereas the present list includes 3 ferns, 1 gymno-
sperm (introduced) and 178 flowering plants. Of the 114 species of
vascular plants tabulated by Jordan for Penikese, including Gull
Island, a considerable number (about 25) have not been found as the
result of recent investigations. On the other hand, of the 182 species
of vascular plants on the present list an even larger number (probably
100) were not enumerated in the earlier report.

1 Jordan 1. c. p. 193.

'Lewis, I. F. Rhodora, xxvi. 188 (1924).

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1930] Fogg, — Flora of the Elizabeth Islands, JVIassachusetts 157

In comparing these two lists the necessary allowances must be made
for the very natural changes in nomenclature which have come about
during the intervening 55 years. Dr. Jordan states, in a letter to the
writer, that the names he employed were those found in the edition of
Gray's Manual then most recent (Ed. 5). With this fact in mind, it
then becomes possible to reconcile a few of the disparities in the two
lists. In general, these discrepancies fall roughly into three categories
which may be briefly described as follows :

In the first place, there are those cases in which a difference of
names involves direct synonymy. Thus, the plant listed by Jordan as
Dicksonia pundilobula Kunze is surely the same as that which we are
today calling Dcnnstaedtia pvndilobula (Michx.) Moore. Similarly,
his Panicum Crusgalli L. corresponds to our Echinochloa Cru^galli
(L.) Beauv., his Triticum repens L. to our Agropyron repens (L.)
Beauv., his Scirpus pungens Vahl to our ^S. americanus Pers., his
Maruia Cotula DC. to our Anihemis Cotula L., and so on.

The second class of discrepancies includes cases involving mistaken
identity or in which an older species has become recognized as con-
sisting of two or more separate and distinct entities. For example,
Calamagrostis arenaria Roth of Jordan's list is certainly the plant
known today as Ammophila hreviUgulata Fernald. This is not at all
a case of direct synonymy, but merely an instance where an American
plant, as beautifully pointed out by Fernald, has proved upon study to
be entirely distinct from its Old World ally. Again, Jordan *s Spartina
stricta Roth is doubtless our 5. alterniflora Lx)isel. var. pilosa (Merrill)
Fernald, his Scirpu^ maritimvs L. our S. campestris Britton, var.
paludosus (A. Nels.) Fernald, his Sisyrinchium Bcrmudiana L. our
S. angustifolmm Mill. Spergularia salina Presl, of Jordan's list,
appears not to grow on Penikese at the present time, but 5. leiosperma
(Kindb.) F. Schmidt is fairly common and we are presumably warrant-
ed in applying a modern interpretation to the older name. Compar-
able to this are Ccrastium viscosum L. for which we find only C.
vulgatum L., Viola sagiitata Ait., which is represented only by V.
fimhriatula Sm., Scutellaria galericulata L. which is replaced by S.
epilobiifolia Hamilton, and a host of similar cases. In all such
instances, then, we are probably justified in assuming that a plant
bearing an unallowable name on the early list is represented today by
the name of a recently recognized segregate or a closely related species,
rather that that it has actually disappeared from the flora.

158

Rhodora

[August

Finally, there are several plants on the Jordan list concerning the
identity of which, in the complete absence of herbarium material, it is
futile even to hazard a guess. Panicum dichotomum L., for example,
which so far has not been collected, may ultimately be found still
growing on the island, or, since that name was applied in a very broad
sense in 1873, Dr. Jordan may really have had reference to P. oricola,
P. meridionale, P. Lindheimeri, var. fasciculatum or to something
still different not yet reported from the island. Likewise, Carex
straminea Schkuhr, a name used loosely before this group had received
critical study, may be equivalent to either C. Longii, C. hormathodes or
C. silicea or to all three. And again Polygonum Hydropiper L., not
known from Penikese today, may equal P. pundatum Ell., which is
ubiquitous, or some other species not yet found. Through the unfor-
tunate lack of preserved vouchers, therefore, all such ambiguous
references, when uncorroborated by subsequent collections, must be
discredited.

After having made all due allowances, however, for inequalities
arising from synonymy, modern revisions and ambiguous records, there
still remain slightly more than a score of plants of the 1873 list which
recent searches have failed to reveal. Most of these are species which
occur on the other islands and their absence from Penikese may be
merely an apparent one, to be remedied by further scrutiny. As
such may be mentioned: Poa annua, Juncu^ pchcarpus, Atriplex
arenaria, Salsola Kali, Euphorbia polygonifolia, E. maculata, Hyper'
icum mutilum, and Asclepias incarnata, var. pulchra. With the
exception of the last two species named, which may have disappeared
as the result of gradual drying up of the ponds, there is every reason
to expect that these plants will some time be collected on the island.
A few others, such as Ruppia maritima, Salicomia europaea (S.
herbacea) and Suaeda maritima, are plants of brackish situations and
their absence is confirmed by independent observations on the tend-
ency of the two saline areas on the island to become overgrown and to
lose their brackish character. Even during the six years from 1923
to 1928, South Pond has become distinctly less brackish, both as to
the nature of its margin and its algal flora. The same can apparently
be said for the ponds on the peninsula, and while these may be merely
local and transitory phenomena, they suggest the gradual diminution
of brackish areas formerly more extensive, a change worth noting.
Three of Jordan's plants which were listed from Gull Island only,

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 159

namely, Rhus Toxicodendron, Coelopleurum lucidum {Archangelica
Gmelini) and Limonium carolinianum (Statice Limonium), are not
only still missing from Penikese, but have completely disappeared
from Gull Island as well. The total absence of Poison Ivv from
Penikese, in the face of vigorous and repeated search for it, is one of
the queer and not altogether unpleasant surprises of this island.
Finally, Jordan lists three plants which are not only unknown from
Penikese but, so far, have been collected on none of the other Elizabeth
Islands: these are Puccinellia (Glyceria) maritima, Salix discolor, and
Iva oraria (I. frutescens). The first of these may well refer to P.
paupercula, var. alaskana, known only from Cutty hunk, the next
might easily have been an introduction which had died out, and Iva
probably is another of the diminishing salt marsh tribe, a species
which it would be interesting to add to the list of Elizabeth Islands
plants.

Turning now to the modem list of the flora of the island, we find,
after again making the necessary correction for synonymy, errors, etc.,
that of the 182 species of vascular plants which it includes, at least
96 (53)% can not in any way be identified with anything on the earlier
list. This large number of species not seen or listed by Dr. Jordan can
conveniently be divided into three groups, as follows: (1) Garden
escapes, about 20 species; (2) Cosmopolitan adventives, about 20
species; (3) Native plants, over 50 species.

Garden escapes. This includes a few ornamentals which may well
have been planted during the days of the leper occupation, some of
which have spread, while others have just barely managed to persist; a
few have escaped from the cultivated area near the site of the old
laboratory building which was destroyed by fire in 1896. A partial
list includes:

LiHum tigrinum
Asparagus officinalis
Gysophila paniculata
Dianthus barbatus
Rubus laciniatus
Rosa rugosa

Oenothera grandiflora
Ligustrum vulgare
Digitalis purpurea
Lonicera japonica
Helianthus annuus
Coreopsis lanceolata

Here also, since this is a class of plants the introduction of which
would appear to have been premeditated, should be mentioned a few
trees which were set out around the dwelling of the resident physician,
namely:

160

Pinus sylvestris
Salix pentandra
S. alba

Rhodora

[August

Populus alba

Acer Pseudo-Platanus

A. platanoides

Cosmopolitan adventives. These are the ever present European
introductions which occur more or less commonly in dry sterile soils
and cleared ground, especially near the haunts of man. It is rather
surprising that so many plants of this class should be lacking from
Jordan's list, but they have probably been brought in with fodder,
building materials and other supplies. A few of these may be cited :

Avena sativa
Dactylis glomerata
Bromus secalinus
B. hordeaceus
Carex contigua
Polygonum Convolvulus
Stellaria graminea
Sisymbrium altissimum
Ranunculus acris

Trifolium pratense
Hypericum perforatum
Daucus Carota
Convolvulus arvensis
Linaria vulgaris
Tanacetum vulgare
Sonchus arvensis
S. oleraceus
S. asper

Native plants. As noted above, more than one-half (96 species) of
the plants on the present list of the flora of Penikese appear to have
reached the island since 1873. Of these 96 species, about 40 have
received consideration in the two classes just dealt with ; their appear-
ance on the island since 1873 may be accounted for in the light of their
being introductions, either accidental or intentional. Permitting of
no such simple explanation, however, is the occurrence today on the
island of more than 50 species of native plants which were not recorded
as present in 1873 by Dr. Jordan. A few of these, such as Ranunculus
delphinifoliuSj Potentilla pumila, Callitriche hetcrophylla and Ilysanthes
inaequalis, are rather inconspicuous forms and might conceivably
have been overlooked in the preparation of the original report. Others
are late-blooming members of the Compositae and, as Jordan lists
only one Golden-rod, Solidago semperviren^, and not a single Aster, it
would seem that he had not remained on the island long enough to
obtain a fair sample of the flora of late summer and might thus have
completely missed Solidago juncea, S. rugosa, S. nemoralisy S. canaden'
sisy S. tenuifoliay Aster undulatus, A. multiflorus and A. vimincus, all
of which appear on the present list. In this group also, might be
placed such things as Gnaphalium obtu^ifolium, Rudbeckia hirta, and
Bidcns connata, although these plants are generally recognizable by the
first of August, as, indeed, are most of the Goldenrods and Asters
listed above. Incapable, however, of any such interpretations as

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 161

those just offered, is the present existence on Penikese of the following
plants, most of them conspicuous, some of them dominant elements
of the vegetation :

Athyrium angustum
Thelypteris palustris, var.

pubescens
Typha latif olia

Panicum virgatum, var. spissum
Danthonia spicata
Distichlis spicata
Scirpus validus
Carex hormathodes
C. silicea

Juncus dichotomus
J. Greenei

J. effusus, var. costulatus
J. acuminatus
J. articulatus

Smilax rotundifolia

Sisyrinchium graminoides

Myrica caroliniensis

Rumex maritimus, var. f uegiiius

Amelanchier oblongifoHa

Rubus pergratus

Rosa palustris

Prunus serotina

Rhus typhina

Parthenocissus quinquefolia

Oenothera biennis

Ligusticum scothicum

Asclepias syriaca

Galium Claytoni

Sambucus canadensis

It is difficult to believe that all these plants could have escaped the
attention of the compiler of the original list and we are rather forced to
the conclusion that they have made their advent to the island since
1873. By just what means they may ha\'e made their way to Penikese
and just how much significance may be attached to their occurrence
there today, are matters for conjecture. Certainly they are not
species preeminently adapted for wind dispersal, although Typha and
Asclepias constitute exceptions to this statement, and spores of the
two ferns may possibly have been transported by that agency. A
few of them, such as Scirpus validus^ Juncus acuminatus, J. articulatus
and the Rumex y which grow in or around ponds, may have been
brought in by birds, while a few others have fleshy seeds or fruits and
may also have been introduced in this way. On the other hand, the
presence of this large block of recent arrivals may be merely illustra-
tive of what takes place on these islands when the practice of raising
sheep is discontinued, although it is difficult at first sight to see just
why these particular species should have been kept down until recent
times when so many others were not only present in 1873 but have sur-
vived the "prolonged struggle for existence against grass and sheep."

{To he continued)

THE FLORA OF THE ELIZABETH LSLANDS,

MASSACHUSEITS

John M. Fogg, Jr.

(Continued from page 161)
THE ORIGIN OF THE FLORA

Any attempt to unravel the geographic origin of the chief elements
comprising the flora of the Elizabeth Islands must not only concern
itself with a close scrutiny of the vegetation of the immediately
adjacent regions, hut must also take into account supplementary
evidence from two main sources, namely, the history and nature of
the coastal plain and its flora, on the one hand, and the study of the
morainal deposits of which most of southeastern Massachusetts is
composed, on the other. In fact, so far as concerns the Elizabeth
Islands, these two problems are rather inextricably linked and one can
scarcely be considered independently of the other.

The Coastal Plain. Perhaps no geographic province in North
America has received greater botanical attention over a long period of
time than the Atlantic coastal plain. Occupying a narrow strip east
or southeast of the Piedmont Plateau, from which it is more or less
sharply marked off by the fall line, the coastal plain has been thought
of as extending from the Gulf States and Florida northeastward
through the Southern and Middle Atlantic States and reaching its
northern limit in northeastern New Jersey, near the Hackensack
Marshes, with a representation eastward on Staten Island, Long
Island, and the immediate coast district of southeastern Massachu-
setts.

The surface of the coastal plain presents in general a very gentle
slope to the southeast, which, in southern New Jersey for instance,

108

Rhodora

[September

averages 5 to 6 feet per mile and is seldom over 10 to 15 feet. East-
ward, beneath the waters of the Atlantic, the coastal plain continues
with the same gentle slope to the 100-fathom mark, where, about 100
miles from shore, it suddenly drops off to abysmal depths. In the
southern states the elevated portion of the coastal plain widens to
about 150 miles, while the submarine portion dwindles, finally, off the
east coast of Florida, to disappear almost entirely. Northward the
submerged portion increases in width, reaching 500 miles off the coast
of Newfoundland, while the subaerial portion diminishes, becoming a
mere fringe of islands and the peninsula of Cape Cod, and finally
disappears altogether. Throughout, the soils of the coastal plain are
of a recent nature, being largely Tertiary and Quaternary, and it
appears likely that the present fall line represents roughly the shore
line at the end of Cretaceous time.

The flora of the coastal plain has long been recognized as distinctly
southern in character, due partly to the nature of its constituent
soils (for the most part, sands, clays, gravels, etc.) which have per-
mitted a northeasterly extension of a Carolinian flora, and partly to
its climate, for the temperature is appreciably milder than that of the
only slightly more elevated continental mass to the westward.

In the state of New Jersey, three-fifths of which lies inside the
coastal plain province. Dr. Witmer Stone recorded in 1910^ the
presence of 479 species of austro-riparian affinities, plants ranging
from Florida, Georgia or the Carolinas north to southern New Jersey,
some of them reaching Long Island, Massachusetts or, as we now
know, even farther northward. The recorded number of such species
has been materially increased by recent study.

In 1911 there appeared in Rhodora a very significant paper by
Professor Fernald describing a botanical expedition to Newfoundland
and southern Labrador.^ In part II of this paper, the author discusses
the Geographic Origin of the Flora of Newfoundland. Analysis of the
constituent floral elements there represented reveals that 274 indigen-
ous plants (35% of the total flora) are southwestern types and that,
of these, 60 species (over 7% of the total flora) are Carolinian types,
being known from southern New Jersey (or even farther southward).
Long Island, southeastern Massachusetts, Nova Scotia and Newfound-

» stone, W. The Plants of Southern New Jersey with Especial Reference to the
Flora of the Pine Barrens. Ann. Rept. N. J. Mus. (1910).

» Fernald, M. L. A Botanical Expedition to Newfoundland and Southern Labrador.
Rhodora, xiii. 109-162 (1911).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 169

land, but not found inland or in continental eastern Canada. As typi-
cal of this Carolinian subclass are cited: Schizaea pusillaf Ammophila
breviligulatay Car ex hormathodes, C. silicea, Corema Conradii, Hudsonia
ericoideSf Myriophyllum tenellum, Utricularia geminiscapa and others.

In an endeavor to account for the presence in Newfoundland of this
coastal plain or Carolinian element, Professor Fernald, after consider-
ing the part which may have been played by birds, ocean currents,
floating ice and logs, and winds, and concluding that they are all
inadequate in explaining this distribution, turns to the question of a
post-glacial land bridge. Hollick 's paper on this subject has already
been referred to, and Professor Fernald, quick to see the phytogeo-
graphic significance of these conclusions, quotes at length from Rol-
lick's statements and appends corroborative data from other sources.

Evidence derived from a study of the conditions attending the last,
or Wisconsin, glaciation indicates that the amount of water then
withdrawn from the ocean may have been sufficient, in conjunction
with the tendency to uplift already noted, to leave uncovered a con-
siderable portion of the now submerged continental shelf from the
South Atlantic states to Nova Scotia and Newfoundland. Of course,
as Professor R. A. Daly has recently pointed out, the apparent
upward trend of the coastal shelf, resulting in part from the removal of
this vast volume of water from the sea, would have been counteracted
by the lowering pressure exerted by the tremendous weight of the ice
on the continental mass. Nevertheless, it appears that during the
Wisconsin advance, and for some time following it, a very considerable
portion of the coastal bench must have been above sea level, forming,
with the exception of shallow channels such as that draining the Gulf
of Maine or Cabot Strait, a nearly continuous platform for the migra-
tion of plants, and animals as well, northeastward from the southern
states to Nova Scotia and Newfoundland. This strip of silicious
soils probably offered a nearly imiform habitat for the advance of
species of austro-riparian affinities and their extension northward may
even have occurred at a time when the ice had not fully retreated from
the mainland of the continent.

Later, with the melting of the glacial ice and the liberation of vast
quantities of water to the ocean, and perhaps through the operation
of other factors as well, this continental shelf underwent a period of
submergence which resulted in the drowning of this coastal plain flora
except in those areas, higher than the rest and often widely separated.

170

Rhodora

[September

which suffered no such submergence. Hence we have today in the Pine
Barrens of New Jersey, on Cape Cod and the adjacent islands, in south-
western Nova Scotia and in certain parts of Newfoundland the relics
of this formerly continuous flora. That these plants are nearly all
species of sterile or silicious soils indicates that they were admirably
adapted for migration northeastward along this post-glacial land
bridge and probably explains why they have never subsequently ex-
tended their ranges to the better, richer soils immediately inland but
continue to exhibit the disrupted distribution so well typified by
Schizaea pusilla and Corema Conradii}

In treating this subject elsewhere, Professor Fernald says, "Of
greater interest are the coastal plain species, because they represent
in New England, eastern Canada and Newfoundland a relic of the
extensive flora which during the late Tertiary migrated northward
along the then highly elevated continental shelf and at the drowning
of the shelf were left as relics at isolated points. This isolated remnant
of the flora derived from the southern coastal plain is represented by
about 200 species north of New Jersey, and nearly every excursion to
southwestern Rhode Island, Cape Cod, Plymouth County (Massa-
chusetts), Nantucket, southern Nova Scotia, Cape Breton, eastern
New Brunswick, Prince Edward Island, the Magdalen Islands or
southeastern Newfoundland, adds to the number of thus isolated
species known to us or extends our knowledge of those already recog-
nized. "^

And in 1921, in discussing the results of botanical exploration in
Nova Scotia, the same author states that " if there were need of further
evidence that, since the Pleistocene glaciation the continental shelf
of eastern North America has been high in the air, affording an essen-
tially continuous line of migration across the mouth of the Gulf of
Maine to Nova Scotia, thence to Newfoundland, that evidence is now
abundantly at hand. A striking feature of this migration northward
of the southern coastal plain flora is the fact that several distinctive
species or genera, Schizaea pusilla, Lophiola, Hahenaria flava, and
perhaps Ccratiolay reached Nova Scotia without establishing colonies
on Long Island, Cape Cod or Nantucket. This would seem to
indicate that the uplifted shelf was a region of some complexity or
else some subtle qualities in the habitats of these plants. "^

> See Fernald. 1. c. Plato 90. opp. p. 140.

2 Fernald, M. L. The GeoKrai)hic Affinities of the Vascular Floras of New England,
the Maritime Provinces and Newfoundland. Amer. Jour. Bot., v. 224 (1918).

•Fernald, M. L. The Gray Herbarium Expedition to Nova Scotia. 1920. Rho-
dora, xxiii. 168 (1921).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 171

Enough has probably been said to indicate the overwhelming
amount of botanical evidence in favor of a post-Pleistocene land
connection permitting the northeastward extension of a southern
coastal plain flora. It now becomes imperative to inquire what
part this connection played in the migration of plants to the area
under immediate consideration. However, before discussing the
direct bearing of these findings upon the problem as presented by the
Elizabeth Islands, it will be found helpful to consider what has already
been learned concerning the adjacent areas, especially Nantucket,
Martha 's Vineyard and Cape Cod.

Nantucket. Nantucket, from its isolated position to the south-
east, might be expected to have caught more of these coastal migrants
than the areas to the west and northwest and is therefore considered
first. For our modern knowledge of the flora of this island we are in-
debted to the discerning and painstaking researches of the late Eugene
P. Bicknell, whose account of "The Ferns and Flowering Plants of
Nantucket" appeared in the Bulletin of the Torrey Botanical Club
from 1908 to 1919. The final section of this paper is devoted to a
consideration of the origin of Nantucket's flora.^ Omitting the
hybrids, Bicknell finds upon the island 1103 [1108] species of plants
of which 362 [31%] are listed as introduced and 746 as native. Of
the 746 indigenous species, "over one-half [373 +] • . . may fairly
be accounted as prevailingly more southern in their general distribu-
tion," while "something over 150 species ... are at least pre-
vailingly more northern in their general distribution. "

Turning first to the plants of southern affinities, we find that over a
hundred of them reach their northeastern limit of range in south-
eastern Massachusetts, others reach Vermont, New Hampshire or
Maine, others occur in the Maritime Provinces, while a small group
is found in Newfoundland. The author then gives a list of 38 plants
which appear not to have been found at any more northern or eastern
point than Nantucket. It is of interest to note, in passing, that only 8
of these are known from the Elizabeth Islands. More than 190 of the
prevailingly southern plants occur in the Pine Barrens of New Jersey,
over 300 are plants of the coastal plain elsewhere in that state, while
all of Nantucket's southern-ranging maritime plants, about 40 species,
also occur in New Jersey. Thus we have over 530 species [consider-
ably more than 50%] in the Nantucket flora which display this south-

Bicknell. E. P. Bull. Torrey Club. xlvl. 423 (1919.).

172

Rhodora

[September

ern relationship. To account for this large percentage of austral
forms Bicknell resorts to Fernald's views on the submerged coastal
shelf and sees isolated on Nantucket the remnants of an extensive flora
of southern derivation belonging to the New England seaboard of
Tertiary time, " a flora lost to our later day with these broad coastal
tracts which now lie beneath the sea. Yet not wholly lost. We find
it still, much of it, we may believe, in the less disturbed flora of our
more southern coastal plain, and we find its remnants persisting as the
merest fringe along the withdrawn more northern coast-lines of the
present day. And isolated on Nantucket it has been preserved to us
in that assemblage of southward ranging plants, now a primary
element in the general composition of the flora. "^

Turning now to the more northern element in the Nantucket flora
we find a group of over 150 species of plants. Of this number about
15 are found nowhere at a more southern point, while about 45 are
near the southern limit of their coastwise range; others reach south to
Long Island and a large number find their southern limit in New
Jersey. In this connection it is interesting to note that of the list of 59
species given by Dr. Stone as reaching from the Maritime Provinces
south to New Jersey, 18 are unknown from Nantucket. Stone's list,
as it appears in his Flora of Southern New Jersey ,* is here given. It
should be observed that the terminology has been revised so as to
correspond to that employed in the present Catalog of Elizabeth
Islands plants. The letter " N " following the name of the plant indi-
cates it is known fron Nantucket, "M" from Martha's Vineyard, and
"E" from the Elizabeth Islands.

Isoetes Braunii N
Lycopodium inundatum
Schizaea pusilla

Potamogeton Oakesianus NME
Scheuchzeria palustris, var.

americana
Triglochin maritima NME
Hierochloe odorata N
Spartina Michauxiana NME
Phalaris arundinacea
Glyceria canadensis NME
G. obtusa ME
G. grandis N
Scirpus subterminalis
S. campestris, var. paludosus NME

> Bicknell. 1. c. p. 434.
« Stone, W. 1. c. p. 49.

Eriophorum tenellum NME

E. gracile N

Carex lanuginosa NE

C. trichocarpa

C. exilis

C. livida

C. canescens, var. disjuncta NME

C. rostrata, var. utriculata N

C. limosa E

C.siUcea NME

Eriocaulon septangulare NME

Juncus articulatus NME

J. pelocarpus NME

Sisyrinchium angustifolium NE

Populus tremuloides NM

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 173

1

P. grandidentata NM

Salix Bebbiana N

S. lucida

Suaeda maritima N

Chenopodium rubrum N

Arenaria lateriflora NME

Nymphozanthus variegatus NME

Actaea rubra NM

Ranunculus Cymbalaria NME

Polanisia graveolens

Rosa virginiana NME

Dalibarda repens

Geum striatum

Lathyrus maritimus NME

Geranium Robertianum

Corema Conradii NM

Hypericum boreale NME

H. ellipticum

H. Ascyron

Myriophyllum tenellum NME

Arctostaphylos uva-ursi, var.

coactilis N M
Vacciiiium pennsylvanicum NM
Glaux maritima N
Menyanthcs trifoliata, var.

minor NE
Limosella subulata NME
Utricularia intermedia N
Plantago oliganthos NME
Solidago uniligulata N
Aster nemoraUs
Xanthium commune

As will be noted, of this list of essentially northern forms Nantucket
has 41 representatives, Martha's Vineyard 27 and the Elizabeth

Islands only 25.

Bicknell gives a list of northern plants which occur on Nantucket
but are unknown from the coastal plain of New Jersey, only a few of
them passing on to Long Island. The significant feature of this list
in the present connection is that, although it contains 44 plants, not
more than a half dozen of these are found on the Elizabeth Islands.

Mr. Bicknell endeavors to account for the presence of these northern
forms, especially those of a maritime character, on Nantucket and
elsewhere to the southwest by supposing that the same marginal land
connection which allowed the plants of the southern coastal plain
to reach Newfoundland would have permitted a counter extension of
northern species to the southwest, perhaps at a later date. The
author also points out that no farther away than Cape Cod there are
established others of these northern species which have not readied
Nantucket and likewise that Cape Cod possesses an extensive coastal
plain flora which is not represented on this seaward island only a few
miles to the southeast. These last two facts are significant because
they apply, even in more marked degree, to the Elizabeth Islands. Not
only, as indicated in speaking of the lists of northern species found on
Nantucket, are many of these boreal forms lacking from the Elizabeths,
but we also fail to find there that large element of southern coastal
plain types which is conspicuous on Nantucket and almost dominant
in certain regions on the Cape.

Martha's Vineyard. Unfortunately, far too little is known con-
cerning the flora of Martha's Vineyard to permit of drawing any

174

Rhodora

[September

conclusions as to the origin of the elements there represented. It is
rather surprising that this relatively large and very attractive island
should have escaped careful botanical treatment, but such is the case.
This is not to say that no botanist has ever visited the Vineyard for the
purpose of collecting specimens, for there have been over a score of
independent collections made. A little more than a century ago,
1829, William Oakes visited the island and recorded some interest-
ing finds. One of the most important collections made was that of
Sydney Harris, who from 1891 to 1904, and again later in 1911 and
1914, collected many sheets, mostly from around Chilmark. The
island was visited by C. A. Weatherby in 1900, by Professor Fernald
in 1901, by A. H. Moore in 1904, by J. A. Cushman in 1906 and 1911,
by E. P. Bicknell in 1909 (and again in 1912 and 1913), by F. W.
Pennell in 1911, by Miss Magaret Heatley, (now Mrs. C. E. Moss),
beginning in 1916; and all of these brought back material which has
been distributed to one or more of the large herbaria of the eastern
United States. Perhaps the largest collections made were those of
Frank C. Seymour in 1916 and 1917; Seymour's specimens have been
sent out by the Gray Herbarium. But, so far, no one has published
any coherent account of the flora of the island, and New England
botanists in general know less about its vegetation than about that of
many a more isolated area.

The writer has undertaken to draw together in a single list all the
records based upon specimens available from Martha's Vineyard. In
the course of this task a systematic census was taken of the material in
the New England Botanical Club and the Boston Society of Natural
History. This resulted in the compilation of a list which inchides
about 700 plants. In an effort to supplement this knowledge two
field trips were made to the island, one in August, 1927, and the other,
in company with Professor Fernald, in August, 1928. From the
information thus derived only one conclusion can be safely drawn,
namely that from our present insufficient knowledge of the island, the
surface only of which seems to have been touched, it is impossible to
speak intelligently of the origin of its flora. One or two general
statements, however, can probably be made with a fair degree of
assurance.

In the first place, it seems evident that the long line of high hills
which flanks the north shore from Menemsha to West Chop supports
a flora of a northern or continental nature. Several plants were

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 175

found here, in the richer soils of the wooded slope, which are either
absent from or far from common in southeastern Massachusetts, and
it seems very likely that careful search will disclose many more things
of this nature.

A second point which seems entitled to emphasis is, that, in general,
the flora of Martha 's Vineyard is far from being closely related to that
of the southern coastal plain. Further study may serve to disprove
this observation, but Professor Fernald and the writer, while exploring
the region aroimd Gay Head, were imable to escape the conviction
that the flora dealt with was continental rather than coastal in char-
acter. Time and again the impression was borne liome that the
coimtless southern plants which form the primary element in the flora
of the middle part of Cape Cod were conspicuously absent. An
exception to this general statement may, perhaps, be constituted by
the flora of the eastern part of the island, the region around Edgar-
town, where, apparently, there is a larger representation of austro-
riparian types than may be met with elsewhere on the Vineyard. If
these observations be justified, they will be found to fit in rather well
with the interpretation of the effects of glacial activities upon the
origin of the flora of southeastern Massachusetts.

Cape Cod. In speaking of the flora of Cape Cod it is necessary first
to have very clearly in mind the fact that botanically, as well as geo-
logically, this region is far from being a unit, but, rather, may be
divided, more or less distinctly, into three separate provinces, which,
for the sake of convenience, may be designated the " Upper, " " Mid-
dle" and "Lower" Capes.

" Upper'' Cape. This includes roughly Sandwich, Bourne, Mashpee,
Falmouth and the western half of Barnstable. The line of morainal
hills which traverses this province from north to south may be traced
southwest from Woods Hole, as it is of this same ridge that the
Elizabeth Islands are formed. On the mainland these hills are rather
heavily wooded and the superficial aspect of this part of the Cape is
that generally associated with an Alleghanian flora with a slight tinge
of the Canadian. This impression is borne out by a study of the
plants which occur here, many of which are either entirely lacking or
only very locally known elsewhere on the Cape. There are well over
150 such plants, constituting a list too lengthy for inclusion liere, the
following few species may, however, be cited as typical:

Rhodora

[September

B. papyrifera
Alnus noveboracensis
Ranunculus recurvatus
Thalictrum revolutum
Actaea rubra
Aquilegia canadensis
Chrysosplenium americanum
Potentilla tridentata
Rubus allegheniensis
Nemopanthus mucronata
Celastrus scandens
Circaea latifolia
Rhododendron canadense
Fraxinus americana
Scrophularia lanceolata
Pedicularis canadensis
Triosteum perf oliatum
Viburnum acerifolium
Lobelia inflata
Solidago ulmif olia
Aster nemoralis
A. acuminatus

176

Polypodium virginianum
Polystichum acrostichoides
Thelypteris Phegopteris
Osmunda Claytoniana
Botrychium virginianum
Lycopodium lucidulum
L. clavatum
Potamogeton Robbinsii
Panicum subvillosum
P. latifolium
Oryzopsis pungens
Cinna arundinacea
Glyceria grandis
G. acutiflora
Scirpus debilis
Carex tribuloides
C. scabrata
Juncus secundum
Smilax herbacea
Trillium cernuum
Habenaria dilatata
Malaxis unifolia
Betula lutea

Few, if any, of these plants, as will be seen, may be looked upon as
species characteristic of a southern coastal plam flora.

"Middle'' Cape. This province embraces the eastern part of
Barnstable, all of Yarmouth, Dennis, Brewster, Harwich, and, perhaps,
Chatham and Orleans. A ridge of morainal hills extendmg east and
west along the north shore from Sandwich to Dennis, forms the back-
bone" of this part of the Cape and is flanked to the south by a broad
outwa^h plain. This area is not without its trees, but m certam
parts scrub oaks predominate and the appearance of the vegetation
differs strikingly from that of the inner Cape. The most salient
botanical feature of this province resides in the plants of the numer-
ous ponds and pond margins, many of which are sandy or peaty and
offer an ideal habitat for an extensive flora of an austro-riparian
nature. Altogether there are over 200 species which are either
peculiar to this part of the Cape or which here find their greatest
development, being represented only casually in the other two pro-
vinces. A partial list of these plants follows :

Thelypteris simulata

Pteridium aquilinum, var. pseudo-

caudatum
Potamogeton pectinatus
Sagittaria Engelmanniana

S. teres

Andropogon virginicus
Paspalum psammophilum
Panicum verrucosum
P. Bicknellii

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 177

P. microcarpon

P. annulum

P. mattamuskeetense

P. mattamuskeetense, var.

Glutei
P. spretum
P. Wrightianum
P. albemarlense
P. auburne
P. Commonsianum
P. polyanthes
P. Ashei

Cenchrus pauciflorus
Spartina cynosuroides
Glyceria laxa
G. Fernaldii
Puccinellia f asciculata
Eleocharis rostellata
Psilocarya scirpoides
Scirpus Smithii, var. setosus
S. campestris, var. novae-angliae
S. Eriophorum
Fuirena squarrosa
Hemicarpha micrantha
Rynchospora Torreyana
R. capitellata, var. discutiens
Carex straminea
C. alata
C. annectens
C. laevi vagina ta
C. MitcheUiana
Arisaema Stewardsonii
Peltandra virginica
Juncus effusus, var. compactus
J. pervetus
J. subcaudatus
J. aristulatus

Luzula campestris, var. echinata
Lilium superbum
Lachnanthes tinctoria
Salix sericea
Myrica asplenifolia
Fagus grandifolia, var.

caroUniana
Quercus stellata
Q. Margaretta
Q. prinoides, var. rufescens
Comandra umbellata
Polygonum Careyi
P. puritanorum
P. setaceum
Chenopodium leptophyllum

Acnida cannabina

Spergularia canadensis

Crataegus rotundifolia

Rubus tardatus

R. Enslenii

R. multispinus

Lespedeza Brittonii

Amphicarpa Pitcheri

Linum striatum

L. floridanum, var. intercursum

Callitriche palustris

Ilex laevigata

Ceanothus americanus, var. inter-

medius
Parthenocissus vitacea
Hypericum dissimulatum
Helianthemum dumosum
H. Bicknellii
H. propinquum
Viola emarginata
V. incognita, var. Forbcsii
Lythrum hyssopifolium
Rhexia mariana
Epilobium moUe
Oenothera linearis
O. longipedicellata
Proserpinaca intermedia
Cicuta bulbifera
Lilaeopsis chinensis
Vaccinium stamineura
Sabatia Kennedyana
S. campanulata
Apocynum medium
Cuscuta pentagona
C. compacta
Stachys hyssopifolia
Lycopus virginicus
Limosella subulata
Aureolaria pedicularia, var.

caesariensis
Utricularia biflora
U. resupinata
U. subulata
Galium tinctorium
Eupatorium hyssopifolium
Mikania scandens
Chrysopsis falcata
Solidago erecta
S. puberula
S. EUiottii

Sericocarpus linifolius
Antennaria petaloidea

178

Rhodora

[September

A. fallax

Gnaphalium obtusifolium, var.
micradenium

Bidens coronata
Prenanthes serpentaria

As will be seen at a glance, the overwhelming majority of species
in this list are plants of prevailingly southern affinities. Many of
them are common in the Pine Barrens of New Jersey and a consider-
able number were not known to occur north of there, or perhaps
Long Island, when the last revision was made of Gray's Manual in
1908, but have been added to the Massachusetts flora only as the
result of recent investigation. The foregoing list is not in any sense
an exhaustive one, nor could it hope to be, for nearly every summer 's
exploration adds to the already large number of southern coastal
plain species which are known to occur on the central part of Cape Cod.

** Lower" Cape. At Orleans the Cape makes a right-angle turn
and continues almost due north through Eastham, Wellfleet and
Truro to Provincetown, which is at the extreme tip. This "fore-
arm" or outer portion of the Cape is characterized by rather high,
undulating hills, the axes of which for the most part run east and
west. It is thought that the troughs between these hills may have
formed the delta of a glacial river which drained into Lake Agassiz,
a large body of water impounded by the ice and today represented
by Cape Cod Bay. From Truro northward this part of the Cape
consists of a wave-built sand spit. A few of the plants peculiar to
the Outer Cape, such as Andropogon scopariuSy var. polycladoSf
Muhlenbergia mexicana, Cyperus filicinus, var. microdontv^, Orontium
aquaticum, Opuntia vulgaris, Aureolaria pedicularia and Baccharis
halimifolia, are far-ranging southern species, but the significant
feature of the flora of this province is that most of its specialties are
forms which display a northern or at least a continental affinity.
The following are a few of the plants belonging to this category:

Potamogeton natans

Elymus arenarius, var. villosus

Scirpus atrocinctus

Eriophorum spissum

Carex Muhlenbergii, var. enervis

C. limosa

C. oligosperma

C. lasiocarpa

C. Pseudo-Cyperus

C. buUata

Juncus effusus, var. Pylaei

J. articulatus, var. obtusatus

Allium canadense
Liparis Loeselii
Salix lucida
Ranunculus sceleratus
Cardamine parviflora, var.

arenicola
Pyrus melanocarpa
Potentilla tridentata
Rubus idaeus, var. strigosus
R. orarius
R. amnicola
R. recurvicaulis

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 170

R. arcuans
Prunus virginiana
P. pennsylvanica
Corema Conradii
Ilex verticillata, var. fastigiata
Hudsonia tomentosa, var, inter-
media
Circaea alpina
Cornus canadensis

Arctostaphylos Uva-ursi, var.

coactilis
Vaccinium pcnnsylvanicum, var.

niyrtilloides
V. Oxycoccus

Galium trifiduni, var. lialophiluin
Liiniaea borealis, var. americana
Bidens cernua
Lactuca Morssii
Hieracium marianum

Most of the species listed above are of a prevailingly northern
distribution; their affinities are with the widely dispersed Canadian
flora to the west and northwest. A few of them are here at, or near,
the southern limit of their ranges.

Thus, it will be seen, each of the three natural divisions of Cape
Cod possesses a rather distinctive flora: that of the Inner Cape is
essentially of a continental Alleghanian-Canadian character; that of
the Middle Cape is colored by the presence of a considera})le number
of. Carolinian or even Louisianian types; while that of tlie Outer
Cape is rendered striking by the occurrence of so many species of
Canadian, or in some cases even Hudsonian, affinities. An attempt
to explain the underlying reasons which account for this differentia-
tion will be made in summing up the evidence for the origin of the
flora of the Elizabeth Islands themselves.

Elizabeth Islands. Turning now to the region under immediate
consideration, we find that the total number of species, varieties
and forms of vascular plants known to occur upon the Elizabeth
Islands is 686. Of these, 128 (183^2%) ^^re introduced, while o^S
(81 3^%) may safely be classed as indigenous.

Introductions. The subject of those species of plants introduced on
the Elizabeth Islands was dealt with at such length in the section
on Changes in the Flora that it seems scarcely necessary to develop
it further here. It need merely be pointed out that by far the larger
part of these introductions is comprised of those ubiquitous European
and Asiatic adventives which ever^-^here throughout eastern North
.\merica have taken possession of recently cleared ground or dis-
turbed sandy areas, often completely dominating our native flora.
The rest of the exotics are either garden escapes, as exemplified in
the discussion of the foreign elements in the flora of Penikese, or
species, such as the Cytisus or the various Spruces, which have been
deliberately planted by man for a special purpose. A few plants

180

Rhodora

[September

native to North America are certainly not of indigenous occurrence
on the islands, but have been introduced either accidentally or inten-
tionally. As examples of such may be mentioned Glaucium flavum
and Solanum triflorum, which grow on the beach near Tarpaulin
Cove, Naushon, and Juniperus communis, var. depressa, which
appears to have been planted along the north shore at the west end of
the same island.

{To be continued)

THE FLORA OF THE ELIZABETH ISLANDS,

MASSACHUSETTS

John M. Fogg, Jr.
(Continued from page 180)

Native plants. The 558 species of indigenous plants on the Eliza-
beth Islands fall rather clearly into three fairly well differentiated
categories. In the first place, there is the southern element— plants
of the southern coastal plain which range north from Florida or the
Gulf States to achieve their northern limit in southeastern Massa-
chusetts, some of them passing on to Nova Scotia or Newfoundland;
then there is a group of species of northern affinities, many of which
have already been mentioned in speaking of Nantucket and Cape
Cod, which range south or southwest to Massachusetts or, at most,
New Jersey. And, finally, there is a large and very important
block of plants which fall into neither of the two preceding classes,
but belong rather to a continental upland flora than to that which
characterizes the lowlying reaches of most of Cape Cod and the ad-
jacent islands. It will be well to analyze briefly the constituents of
these three groupings before proceeding further.

The Southern Element. In rather striking contrast to the situa-
tion found on Nantucket, where, it will be remembered, over 50%
of the indigenous flora is prevailingly more southern in its hue, as well
as on the middle part of Cape Cod, where, as has been seen, the na-
tive flora is preeminently that of the southern coastal plain, is the

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 209

fact that on the Elizabeth Islands this southern element finds ex-
pression in something less than 20% of the total indigenous flora.

It is true that there are a few species of plants of the southern
coastal plain which, on the Elizabeth Islands are near the very north-
eastern limit of their distribution. Paspalum seiaccum, for example,
is known in Massachusetts only from the Elizabeths and Nantucket.^
Panicum longifolium is found nowhere east of Pasque Island, at which
place it is abundant in the peaty bog hollows,^ although it is repre-
sented in Nova Scotia by var. tuskctense.^ Tipularia discolor is near
the northeastern limit of its range on one of the Elizabeth Islands
(Nashawena) and on Martha's Vineyard. Rinnex vcrticillatus, long
known from Block Island, but otherwise rare in New England, has
recently been collected on the Elizabeths. Hydrocoiijlc Canbi/i^ and
H. verticillata, both known on the basis of old records from Woods
Hole (for years their only known station in New England) have,
during the course of the present survey, been discovered on the
Elizabeth Islands as well. Solidago minor ranges from Alabama
and Florida to Virginia, then ''jumps" to Nantucket, where it was
reported by Bicknell, and is now known to occur on Naushon, the
largest of the Elizabeths. Thus it will be seen, that these islands,
as is true of nearly every other locality along the coast from New
Jersey northward, are not totally lacking in records which represent
interesting, or even spectacular, northern extensions of plants which
are essentially southern in their affinities.

In general, however, the flora of the Elizabeth Islands far from sug-
gests that of the coastal plain. The following enumeration, which in-
cludes the seven species just mentioned, constitutes a nearly complete
list of the plants known from these islands which are also characteristic
species of the coastal strip, ranging from the Gulf States, Florida or
Georgia northeastward. Many of them, of course, continue farther
north and east, being known from Nova Scotia, New Bnmswick or
even Newfoundland, but they are, for the most part, plants of a
pronounced austro-riparian origin. It would be difficult to make
such a list comprehensive, for, in the absence of adequate data
concerning the complete ranges of every species, it is not always easy
to state categorically whether a plant belongs exclusively to the
southern coastal plain, or whether it enjoys an Alleghanian-Carolin-

»See Weatherby. Rhodora, xxx. 133 (1928).
« See Fogg, Rhodora, xxxi. 39 (1929).
»See Femald, Rhodora, xxiii. 192 (1921).

IRREGULAR PAGINATION

210

Rhodora

[October

ian distribution. All of the species listed below, with the exception
of those marked with an asterisk, are known also from Cape Cod.

Woodwardia virginica
Thelypteris simulata
Lycopodium inundatum, var.

Bigelovii
Chamaecyparis thyoides
Sparganium eurycarpum
Potamogeton Oakesianus
P. pulcher
Andropogon scoparius, var.

polyclados
A. virginicus
*Paspalum setaceum
P. pubescens
Panicum meridionale
P. albemarlense
P. oricola
*P. longifolium
P. Commonsianum
Setaria geniculata
Cenchrus paucifiorus
Stipa avenacea
Aristida purpurascens
Ammophila breviligulata
Spartina alterniflora, var. pilosa
*Diplachne maritima
Cyperus erythrorhizos
C. ferax

Eleocharis rostellata
Fimbristylis autumnalis
Scirpus Olneyi
S. robustus

Rynchospora capitellata
Carex Longii
C. straminea
C. alata
C. Howei
C. Mitchelliana
Xyris caroliniana
Juncus effusus, var. costulatus
Luzula campestris, var. echinata
Smilax rotundifolia
Iris prismatica
Sisyrinchium graminoides
Pogonia ophioglossoides
Calopogon pulchellus
*Tipularia discolor
Myrica caroliniensis
Spergularia rubra

Boehmeria cylindrica, var.

Drummondiana
*Rumex verticillatus
Polygonum glaucum
P. punctatum
Drosera intermedia
Pyrus arbutif olia
Rubus Andrewsianus
Rosa palustris
Prunus maritima
Desmodium obtusum
D. marilandicum
Lespedeza capitata
Polygala cruciata
Euphorbia polygonifolia
Ilex opaca
I. glabra

Hibiscus Moscheutos
Hypericum virginicum
Helianthemum canadense
H. Bicknellii
Lechea villosa
L. maritima
Decodon verticillatus
Rhexia virginica
MyriophyUum scabratum
Proserpinaca palustris
*Sanicula canadensis
Hydrocotyle umbellata
H. Canbyi
H. verticillata
Ptilimnium capillaceum
Clethra alnif olia
Rhododendron viscosum, var.

glaucum
Leucothoe racemosa
Samolus floribundus
Bartonia virginica
B. paniculata
Nymphoides lacunosum
Asclepias verticillata
Teucrium canadense, var.

littorale
Ilysanthes inaequalis
Gratiola aurea
Agalinis maritima
Utricularia gibba
*Plantago virginica

1930] Fogg,--Flora of the Elizabeth Islands. Massachusetts

211

Eupatorium hyssopifolium

SoUdago EUiottii

*S. minor

S. tenuifolia

Aster dumosus

A. vimineus

Pluchea camphorata
Gnaphalium purpureum
Coreopsis rosea
Krigia virginica
Lactuca hirsuta
Hieracium Gronovii

A point requiring particular emphasis is, that many of these species
are by no means common on the islands. Indeed, a few of them,
such as Thelypteris simulata, Paspalum setaceum, Panicum longi-
folium, P. Commonsianum, Cenchrus paucifiorus, Diplachne mari-
tima, Eleocharis rostellata, Carex straminea, C. alata, C. Mitchelliana,
Luzula campestris, var. echinata, Tipularia discolor, Rumex verticil-
latus, Prunus maritima. Ilex glabra, Sanicula canadensis, Hydro-
cotyle Canbyi, Rhododendron viscosum, var. glaucum, Triosteum per-
foliatum, SoUdago minor and Coreopsis rosea, are known only from a
single locality, while certain others, though less restricted, are never-
theless rare and local. And seldom, if ever, are these coastal plain
plants present in sufficient abundance to create the impression,
inescapable on Cape Cod, of a southern flora transplanted almost

en masse.

Pursuing this last idea further, it will be found interesting to
contrast with the list just given a list of some of the southern coastal
plain plants which are known to occur on Cape Cod (most of them
from the Middle Cape), but which have not yet been found on the
Elizabeth Islands:

Pteridium aquilinum, var.

pseudocaudatum
Sagittaria Engelmanniana
S. graminea
S. teres

Paspalum psammophilum
Panicum verrucosum
P. Bicknellii
P. microcarpon
P. annulum
P. mattamuskeetense
P. spretum
P. Wrightianum
P. auburne
P. tsugetorum
P. columbianum
P. polyanthes
P. Ashei
P. scoparium

Aristida dichotoma
A. gracilis

Spartina cynosuroides
Tridens flavus
Cyperus filicinus, var.

microdontus
C. Grayii

Eleocharis Robbinsii
E. melanocarpa
Psilocarya scirpoides
Scirpus atrovirens, var.

georgianus
S. Eriophorum
Fuirena squarrosa
Hemicarpha micrantha
Rynchospora macrostachya
R. inundata
R. Torreyana
R. capitellata, var. discutiens

212

Rhodora

[October

Scleria reticularis
Carex annectens
C. intumescens

C. buUata, var. Greenei
Arisaema Stewardsonii
Peltandra virginica
Orontium aquaticum
Xyris Smalliana
Juncus subcaudatus

J. aristulatus
Lilium superbum
Aletris farinosa
Lachnanthes tinctoria
Myrica asplenifolia
Quercus stellata
Q. prinoides
Q. ilicifolia
Comandra umbellata
Polygonum setaceum
Polygonella articulata
Acnida cannabina
Drosera filiformis
Cassia Chamaecrista
Crotalaria sagittalis
Lupinus perennis
Tephrosia virginiana
Desmodium rotundifolium

D. marilandicum
Lespedeza procumbens
L. Stuvei

L. angustifolia
Strophostyles helvola
Linum floridanum, var.

intercursum
Polygala Nuttallii
Corema Conradii

Acer rul^rum, var. tridens
Ceanothus americanus, var.

intermedius
Vitis cordifolia
Hypericum adpressum
Hudsonia ericoides
Viola emarginata
V. primulifolia
Opuntia vulgaris
Rhexia mariana
Oenothera linearis
O. longipedicellata
Proserpinaca pectinata
P. intermedia
Lilaeopsis chinensis
Sabatia campanulata
Cuscuta compacta
Onosmodium virginianum
Stachys hyssopifolia
Lycopus sessilifolius
Agalinis purpurea
Aureolaria pedicularia, var.

caesariensis
Utricularia infiata
U. subulata
Viburnum pubescens
Eupatorium verbenaefolium
Mikania scandens
Solidago erecta
S. puberula
Aster spectabilis
A. subulatus
A. tenuifolius
Baccharis halimifolia
Bidens coronata
Lactuca floridana

Thus it will be seen that, while there occur on the Elizabeth Islands
something like 100 species belonging to a wideranging southwestern
flora. Cape Cod not only has practically every one of these same plants,
but boasts in addition at least an equal number of species of the same
class which, so far as is known, are totally lacking from the islands.

It may be worth while to note, in passing, that, while an overwhelm-
ingly large proportion of the more than 200 prevailingly southern
species which occur on Cape Cod occur likewise on Nantucket (and a
considerably smaller proportion on Martha's Vineyard), that island
has caught a number of these southern migrants which appear not to
have succeeded in reaching the Cape. Several of these may be listed:

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 2\:\

Eleocharis tricostata Ascyron hyi)ericoides

Scleria triglomerata Lespedeza virguuca

Carex Walteriana Lechea Leggettii

Habenaria ciliaris Ludvigia alterniflora

Quercus pagodaefoUa Pyenanthemum verticillatuni

Polygonum robustius Schwalbea americana

Amaranthus pumilus Aster concolor
Ranunculus laxicaulis

Sufficient evidence has probably been adduced to bear out the
contention that the relations of the flora of the Elizabeth Islands to
that of the southern coastal plain are anything but prominently
marked, and that this southern, or southwestern, element is nuich
more strongly represented in the closely adjacent regions, especially
Cape Cod and Nantucket. An attempt to determine the causes
which account for this break in distribution will shortly be made.

The Northern Elevient. Although lacking numy of the northern
types which distinguish the floras of parts of Nantucket and the
"Lower" Cape, the Elizabeth Islands are not entirely without their
representation of plants whose affinities are prevailingly boreal. In
all, about 50 such species, constituting nearly 9% of the total native
flora, may be considered as belonging to this class. It is significant to
contrast this number with the 150 northern plants (over 20%) listed
by Bicknell for Nantucket.

In general, these northern species which occur on the Elizabeth
Islands are plants which range from Labrador and Newfoundland
south to Massachusetts and New Jersey or, in a few cases, Delaware
or Maryland. Many of them range south of New England along
the mountains but reach the coastwise southern limit of their distri-
bution in Massachusetts, Long Island or New Jersey. In the list
which follows those species marked with an asterisk are to be looked
upon as essentially maritime.

♦Ruppia maritima, var. subcapitata

*Triglochin maritima

♦Agrostis stolonifera, var. compacta

Spartina Michauxiana

Glyceria obtusa

G. canadensis

*Puccinellia paupercula, var.
alaskana

Eleocharis uniglumis

Scirpus campestris, var. jialutlosus

Eriophorum tenellum

Rynchospora f usca

Carex hormathodes

C. silicea

C. canescens, var. disjuiicta

C. limosa

C. lanuginosa

Eriocauloii septangulare

Juncus pelocarpus

J. militaris

J. articulatus

Sisyrinchium aiigustifoliuni

Liparis Loeselii
Betula populifolia

214

Rhodora

(October

*Rumex maritimus, var. fueginus
Arenaria lateriflora
Sagina procumbens

*Ranunculus Cymbalaria
Drosera rotundifolia
Fragaria virginiana, var.
terrae-novae

*Potentilla pacifica

*Lathyrus maritimus
Rhus glabra, var. borealis
Ilex verticillata, var. fastigiata

Hypericum boreale

Epilobium palustre, var. monticola

Myriophyllum tenellum
*Ligusticum scothicum
*Coelopleurum lucidum

Menyanthes trif oliata, var. minor

Chamaedaphne calyculata

Vaccinium macrocarpon

Limosella subulata
*Plantago juncoides, var. decipiens

Anaphalis margaritacea

The Elizabeth Islands, then, appear to have received their share
of those far-ranging northern types which probably owe their existence
in coastal New England, and southwestward, to the former presence
of the broad continental shelf, already referred to, which permitted
of their extension to the southwest and then, following its submerg-
ence, left them stranded at isolated localities along the coast. This
would also explain why these islands possess fewer such plants than
Martha's Vineyard (see p. 173) and still fewer than either Nantucket
or the outer portion of Cape Cod. For, if these boreal species reached
southeastern New England from off the elevated coastal bench to the
eastward, then it seems logical to assume that a greater number of
them would have found a refuge on Nantucket and the "Lower"
Cape and that a smaller proportion would have succeeded in finding
their way to the areas inland to the west, especially if, as may well
have been the case, the retreat of the glacial ice from the latter region
lagged appreciably behind its retreat from Nantucket, Martha's
Vineyard and Cape Cod.

The Continental Element. It is only when we come to consider
the continental element as it appears on the Elizabeth Islands that
we find ourselves dealing with the type of vegetation which lends a
dominating color to their flora. Probably more than 400 plants
(about 70% of the total indigenous flora) from these islands are
neither prevailingly southern nor northern in their distributional
affinities but belong, rather, to a widespread continental flora which
might be characterized, somewhat arbitrarily, as Canadian-Alleghan-
ian in nature. This, it will be recognized immediately, is an attribute
which the Elizabeths share in common with the upper or inner part
of Cape Cod, and, indeed, there is every reason to suppose that the
flora of these islands may, until comparatively recent geologic times,
have been continuous with that of the line of hills which runs from

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 215

Falmouth to Boume, and even north nearly to Plymouth. This
entire ridge represents a terminal moraine which, beyond Cuttyhunk,
dips below sea level reappearing, as some geologists believe, to form
Block Island and, farther west, a portion of Long Island. The
separation of the Elizabeth Islands from the mainland and their
division into the seven present members of the chain are, as has
already been indicated, relatively modern events. So it is entirely
in keeping with the past history of this region that so many of the
types which are abundant throughout the Falmouth area should
likewise be common on the islands.

The relation of the flora of the Elizabeth Islands to that of the
mainland comes out most clearly upon an examination of the forest
types. The trees have already been listed (p. 151), but it seems
entirely justifiable to repeat in this place that the native woods of
the islands are made up not only of beech, Fagua grandifolia (surely
not a coastal plain type), but also contain Carya alba, Ostrya mrgin-
iana, Quercus alba, Q. velutina, Sassafras officinale, Hamamelis mrgin-
iana, Acer rubrum, Cornus florida and Nyssa syhatica. Now these
trees, while they may and do occur on the coastal plain, are neverthe-
less more common and more clearly at home on the richer soils of the
Piedmont and the areas inland, often reaching their finest develop-
ment on wet wooded slopes and the alluvia of river valleys.

Under the trees listed above, on the wetter parts of the forest
floor, occur such plants as Carex lupulina, Arisaema tnphyllum,
Oakesia sessilifolia, Maianthemum canaden^e, Medeola mrginiana
and Trientalis borealis. These again are types more commonly
associated with an Alleghanian woodland flora. The beech drops,
Epifagus mrginiana, a rare plant in southeastern Massachusetts,
occurs everywhere in the wooded parts of Naushon and Nashawena,
and many other cases of this sort might be cited.

It would be superfluous to list here all of the species of Canadian-
AUeghanian affinities which occur on the Elizabeth Islands. An
enumeration of them would include most of the names of native
plants in the Catalog that follows which have not been listed above in
dealing either with the Southern or Northern Elements. A few of
the most typical, however, not including the few trees mentioned
above, may be given for the sake of comparison:

Polypodium virginiauum
Asplenium platyneuron

Athjrrium angustum
Osmunda cinnamomea

216 Rhodora

Ophioglossum vulgatum

Isoetes Engelmanni

Sparganium americanum

Sagittaria latifolia

Andropogon furcatus

Glyceria striata

G. paUida

Elymus virginicus

Cyperus diandrus

C. rivularis

Scirpus cyperinus

Eriophorum virginicum

Carex rosea, var. radiata

C. cephalophora

C, crinita

C. virescens

C. communis

C. pennsylvanica, var. separans

C. digitalis

C. debilis, var. Rudgei

C. lupulina

Arisaema triphyllum

Symplocarpus foetidus

Acorus Calamus

Juncus effusus, var. solutus

Oakesia scssilifolia

Lilium philadclphicum

Maianthemum canadense

Medeola virginiana

Habenaria bracteata

H. clavellata

H. orbiculata

Arethusa bulbosa

Boehmeria cylindrica

Polygonum scandens

Phytolacca americana

Ranunculus delphinifolius

Anemone virginiana

Coptis groenlandica

Spiraea tomentosa

[October

Amelanchier oblongifolia
Geum canadense
Geranium maculatum
Acalypha virginica
A. digyneia

Callitriche heterophylla
Rhus typhina
R. Vernix
R. Toxicodendron
Ilex verticillata
Impatiens biflora
Vitis labrusca
V. aestivalis
Viola papilionacea
V. pallens
Ludvigia palustris
Cicuta maculata
Slum suave
Heracleum lanatum
Monotropa uniflora
M. Hypopithys
Epigaea repens
Gaultheria procumbens
Lysimachia quadrifolia
L. terrestris
Trientalis borealis
Apocynum androsaemifolium
Verbena hastata
Scutellaria galericulata
Pycnanthemum muticum
P. flexuosum
Epifagus virginiana
Cephalanthus occidentalis
Triosteum perfoliatum
Sambucus canadensis
Lobelia cardinalis
Solidago juncea
S. canadensis
Aster divaricatus
Cirsium discolor

The great bulk of these plants are species primarily of the interior;
they attain their fullest development in the Piedmont and the Up-
lands and their occurrence on the coastal plain may be regarded, in
most cases, as rather casual.

In summing up, it need merely be pointed out that the Elizabeth
Islands, while serving, as does every other locality along the Atlantic
coast, as a meeting ground for both northern and southern species
of plants, exhibit both qualitatively and quantitatively a very strong

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 217

relationship with a widely dispersed flora of a continental nature, a
fact which seems readily explicable \ipon the basis of the close connec-
tion existing between these islands and the inner, hilly part of Cape
Cod, both as regards geologic history and general topography.

Finally, there remains to be considered, as briefly as may be, the
subject of the last glacial advance over this region and its possible
effects in influencing the present-day distribution of the flora.

Glacial History. Probably the greatest student of the geology
of southeastern Massachusetts since the days of N. S. Shaler, was
the late J. B. Woodworth of Harvard University. Professor Wood-
worth had prepared, shortly before his death, an exhaustive treatment
of the glacial history of the Cape Cod region. This manuscript,
unfortunately, still awaits publication and the details which it em-
bodies are not yet available. Happily, however, Woodworth had
related his broader conclusions to A. P. Brigham, geologist to C(>lgate
University, and the main arguments are set forth by Brigham in his
popular book entitled " Cape Cod and The Old Colony. "

According to Woodworth, the advance of the last or Wisconsin ice
over southeastern Massachusetts took place not as a solid sheet,
but in the form of three tongues or lobes. One of these, the " Buzz-
ards Bay Lobe," came down over the region now occupied by Buzz-
ards Bay and deposited as a frontal moraine much of the nuiterial
which now forms the line of high hills along the northwest shore of
Martha's Vineyard, from :\Ienemsha to West Chop. Then, following
an interval which represented a retreat and a second advance of the
ice, this lobe laid down, as a secondary moraine, the ridge which made
the' Elizabeth Islands and the "Upper Cape." The line of the
islands, as may be seen from a map, almost exactly parallels the line
of the morainal hills on the northwest shore of the Vineyard.

The second lobe, which lay to the east of the " Buzzards Bay Lobe,"
advanced southward over what is now the middle section of Cape Cod
and laid down, as a terminal moraine, the sand, gravel and boulders
which form the northeast shore of Martha's Vineyard and the higher,
crescent-shaped portion of Nantucket. This Woodworth terms the
"Cape Cod Lobe." This lobe then retreated, as did the Buzzards
Bay Lobe and, as its secondary moraine, deposited the till which com-
poses the "backbone" of Cape Cod from Sandwich to Brewster and
Orleans. Thus, Martha's Vineyard was built by the combined
action of two lobes and the central and southern parts of the island

218

Rhodora

!■■*

I

represent an outwash or apron plain derived from two separate
moraines. The southern and southeastern parts of Nantucket and
almost the entire south shore of the Cape likewise represent outwash
plains, both formed from the materials deposited by the Cape Cod

Still further to the eastward, the third, or "South Channel Lobe"
advanced over the area now submerged and known as Georges Banks.
With the deposits of this lobe we are not so much concerned, as they
now lie mostly beneath the sea, save for such materials as may have
contributed to the building of the outer or lower part of Cape Cod.
How far this lobe may have extended eastward over the then elevated
continental shelf is apparently not definitely known.

It now becomes pertinent to inquire into the relative ages of these
deposits and as to whether any evidence is forthcoming to indicate at
what time and in what manner the various lobes retreated. Probably
Woodworth's report, when it becomes available, will throw much
light on this question. However, the writer has it on the authority
of Dr. Wigglesworth of the Boston Society of Natural History, who
is conversant with Woodworth's views, and who is himself a student
of the geology of Martha's Vineyard, that in all probability the middle
or Cape Cod lobe was the first one to retreat. If this was the case,
it then means that Nantucket, the eastern part of Martha's Vmeyard
and the central part of Cape Cod were free of ice at a time when the
regions to the east and to the west were still covered by the South
Channel and Buzzards Bay lobes respectively. Remembermg that
the coastal shelf was probably considerably higher at that time than
it is today, and that the Vineyard, Nantucket and the Cape may well
have been contmuous dry land, it at once becomes apparent that
there was thus opened up an area which soon became available as a
refuge for that migration of southern coastal plain species of plants
which probably began as soon as the ice commenced to retreat.

It is necessary to point out here that there is a lack of complete
agreement as to the exact period of subsidence of the continental
shelf and as to whether this migration might have occurred previous
to the advent of the Wisconsin ice or whether it could not possibly
have taken place until after the glacier had receded.

Douglas Johnson, the eminent student of coastal phenomena, m
discussing the date of submergence of the Banks cuesta (i. e., the
New England-Acadian portion of the outer coastal shelf) states that

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 219

"we should expect the subsidence to be at least post-Miocene and
more probably post-Pliocene."^ And further, "It seems probable
that the date of submergence of the drowned topography must be
post-Tertiary. "2 Johnson, then, is inclined to view the elevation
of the continental shelf as pre-glacial rather than post-glacial, a
condition which would have necessitated the plant migration having
antedated the advent of the Wisconsin ice. And, indeed, Femald
sees no reason why these plants should not have moved northeast-
ward along the exposed shelf before the coming of the Wisconsin
glaciation and "have persisted outside the subsequently glaciated
area, finally taking possession of their present isolated habitats on

the receding of the ice. "^

In connection with the present study, however, it matters little
whether these species of the southern coastal plain reached the New
England area before or after the last glaciation. In either case they
must have moved inland from off the broad shelf to the eastward to
take the places left vacant for them by the recession of the ice, and
if we are justified in assuming that it was the Cape Cod Lobe of the
glacier which receded first, then we are in a position to understand
why so many of these species are to be found upon Nantucket and
the "Middle" Cape, and, to a lesser degree, upon the eastern portion
of Martha's Vineyard and are so generally lacking from the Elizabeth
Islands and inner Cape Cod. Even if the western half of the Vmeyard,
the Elizabeth Islands and the "Upper" Cape were free from ice at
the time when this migration was operative, it seems likely that they
offered a type of habitat which was less attractive to these coastal
plain plants than the low-lying silicious areas of Nantucket and the
"Middle" Cape which they must have reached first and where they
today abound, seldom exhibiting a tendency to widen their ranges
into the neighboring regions. And although, as Femald suggests,
these plants may have persisted upon the outer shelf while the ice
still covered the area inland, it is nevertheless probable that the
Nantucket-" Middle" Cape region would have been the first to wit-

ness their return. • j j f _

Ccmcltmon: In summing up, it may be said that, consKlered from

the viewpoint of broad, geographic origins, the native flora of the

1 Johnson. D The New England-Acadian Shon-lino. New York. 302 (1925).

1 F^JLw^'m I A PrelinUnary Statement of Results of Studies on the Northeast-
ward Disiribution of the coastal Plain Flora. Amer. Jour. Sci. 4th Ser. xl. 18 (1915).

220

Rhodora

[October

Elizabeth Islands is seen to consist of three distinct elements. In
the first place, there are those species (less than 20%) which exhibit a
relationship with the flora of the southern coastal plain. Their
presence upon these islands is to be explained upon the basis of a
former land connection with New Jersey and southward which took
the form of an elevation of the outer coastal bench, now submerged,
and which, either prior to or following the Wisconsin glaciation,
permitted of the migration of plants from the southwest to the New
England area and even farther north and east. That so many more
of these southern plants occur upon Cape Cod and Nantucket than
upon the Elizabeth Islands is probably to be explained by the be-
havior of the glacial lobes which covered this area and which, by their
differential recession, seem to have rendered the former areas access-
ible to occupation by plants at an earlier date. Secondly, there is a
small percentage (less than 9%) of plants displaying a boreal affinity,
the occurrence of which may be attributed to a counter extension
southward along this same uplifted shelf. And the fact that the Eliza-
beth Islands ha^ e received a smaller number of these northern repre-
sentatives than either Cape Cod or Nantucket is probably to be
accounted for on the basis of their inland position and the character
of the habitat which they oft'er, which is, in general, less favorable
for these northern plants than the situations which they occupy on
the "Lower" Cape. And, finally, there is the overwhelming majority
(over 70%) of plants occurring on the Elizabeth Islands which show
an essential relationship with the flora of the mainland and which
give to the islands the dominating character of a Canadian- Alleghan-
ian region. The prevalence of this continental element is doubtless
due to the close geologic and physiographic similarity existing between
these islands and the "Upper'' Cape. Thus it will be seen that the
evidence derived from a study of the geographic origin of the flora of
the Elizabeth Islands fits rather well into what is already known
concerning the history of the neighboring territory and that these
islands take their place botanically as an extension of the adjacent
mainland rather than as a link in that chain of outposts of a formerly
continuous but now highly disrupted coastal plain flora extending
from the South Atlantic States to Newfoundland.

Acknowledgements. The writer desires to express his profound
and grateful appreciation of the stimulating genius of Professor M. L.
Femald under whose guidance this study has been conducted and
without whose inspiration it could never have been completed.

Fernald,— Gentiana proccra

221

19301

Mr C A Weatherbv and Mr. Bayard Long have both rendered
invaluable assistance in their willingness to aid in the determination
of critical material. To them the writer 's deepest thanks are due.

He is also indebted to Professor L. H. Bailey, who has kmdly
examined several specimens of Ruhus, to Professor K. ^L Wiegand,
who has looked over some of the Amdanchier material, and to Mrs.
Agnes Chase who has given her opinion on a few sheets of Panicum.

{To he continued)

THE FLORA OF THE ELIZABETH ISLANDS,

MASSACHUSETTS

John M. Fogg, Jr.

(Continued from page 221)

Part II. Annotated List of the Vascular Flora

In the Catalog of Vascular Plants of the Elizabeth Islands which
follows several conventions and abbreviations have, for the sake of
convenience, been adopted ; these are here explained :

NOMENCLATURE. Th^ International Rules of Botanical Nomen-
clature have been followed.

ELEMENTS IN THE FLORA. The various elements constituting the
flora have been differentiated thus :

Indigenous species appear in capital letters.
Introduced species are given in italics.
Discredited records are included in brackets.

CITATIONS AND SYNONYMY. Synonyms are given in italics and,
in general, are included only when they represent names which have
been superseded since the last edition of Gray's Manual (Ed. 7, 1908).
Usually, in such cases, a reference is given to the place where the
new name was published or its status discussed.

LOCALITIES. The seven main islands are indicated by taking the
first three letters of the name of each ; thus :

NON: Nonamesset, including Pine Island.

UNC: Uncatena

NAU: Naushon, including Captain's Island, Ram's Head, Mono-

hansett. Buck, East Buck, West Buck and the Weepeckets.
PAS: Pasque.
NAS: Nashawena.
CUT: Cuttyhunk.
PEN: Penikese, including Gull Island.

19301 Fogg.-Flora of the Elizabeth Islands. Massachusetts 227
COLLECTORS Those who have collected specimens from the

where the collector has employed a mmibermg system. ot^«^^'«^ ^^
t da* A list of the chief collectors is here gn-en m alphabetical

order:

Cushman, J. A. (1906)
Duggar, B. M. (1911)
Faxon, C. E. (1875)
Faxon, Walter (1873)
Fernald, M. L. (1927)

Hervey, E. W. (no date)
HoUick, Arthur (1898)
Jordan, David S. (1873)
MacRue, LilUan (1904)
Moore, A. H. (1904)

Northrop, Alice R. (1901, 1903)

Pennell,F.W.(1911)
Sanford,S.N.F (1917)
Simons, EUzabeth A. (1901)

Sipe, S. B. (1901)
Svenson,H.K.(1926)
Taylor, W.R. (1919-1921)

Weir, "Miss" (1890)
Williams, E.F. (1911)

OOre, A. n. VA»vrt; 1- X i ,.

Further information concerning these collectors and the distribu-
tion of thei^material may be found in the section on Previous Botan-
ical Work on the Islands (Pages 123-12o). ^

N B Serial numbers unpreceded by a name are lo
rep;esenting material collected by the present^.iter.

HFRBAMA The following system of initials has been eu p j
to desrate the herbaria in which specimens have been seen or to
which they are known to have been distributed:

(A) Academy of Natural Sciences of Philadelphia

(B) Boston Society of Natural History

(C) Cornell University .

G Gray Herbarium, Harvard University
J) Arnold Arboretum, Jamaica Plain, Mass.
(M) Missouri Botanical Garden
(N) New England Botanical Club
(P) University of Pennsylvania
(U) United States National Museum
W) Marine Biological Laboratory, Woods Hole, Mass.
(Y) New York Botanical Garden
(o) no specimen seen

POLYPODIACEAE

,r*, T P TiulaaTe of eastern American

POLYPODIUM VIRGINIANUM L. r. ?^*^"^ . ^.^x j^ ggen

'Z«A).2489(N);NAS:^ortArop(o) ^^^,^,,„,„„ (pesv.)

Pteridium aquilinum (I^:) f^"!""' . Apparently not common;

IRREGULAR PAGINATION

228

Rhodora

[NOVBMBBR

WooDWARDiA viBGiNiCA (L.) Sm. Locally abundant; forms
considerable growth around the dune ponds «" N^l^ton wV^ 1 772
WiUiams, July 10, 1911 (N); NAS: NoHhrop. July, 1901 (Y), 1772

^^whREOLATA (L.) Moore. Locally abundant; with rM^/p^em
palmtris, var. pubescens occupying boggy hollows m open hills 1 Ab
Lrwon, Sept. 8, 1926 (N) ; NAS: Northrop, Aug. 1901 (Y), 3492 (N,P) ,

"^ aIpSuS'SLon (L.) Oakes. Reported by Mrs. North-
roD from Nashawena; no specimen seen. ^ , . ^,. . • ^ ^f

Athyeium angustum (WiUd.) Presl. Aspkmum fil^^-^^^^Sf
American authors in part See Gutters Rhodora. xu. 190 0917).
Boggy woods and open hillsides. NAS: 3515 (P) , CU 1 . ^bib (.in;.

A ANGUSTUM, var. ELATius (Link) Butters^ Rhodora, x.x. 191
(1917) Dry, exposed hillside. PEN: 459 (N,P,W).

Thelypteris palustris (Salisb.) Schott, var. pubescens (Lawson)
FeSd ™di«m Thelypteris of Manual. See Fernald, RhodoRA
xxx" 34 (1929). Common in low, boggy ground and around borders
of ponds NON: 2238 (N); UNC: 2993 (N); NAU: Stp., Aug. 901
HNyTaS- Northrop (o), 2349 (N,P); CUT: Sanjord, Aug. 15, 1917

'i^'^Sl^r^k^ £3'%W,-„« sim.latu.^ Dav. See
Weatherby Rhodora, xxi. 174, 178 (1919). Apparently rare »" the
Elands NAU : Sipe, July, 1901 (W) : PAS: Svenson, Sept. 8, 1926 (N)
T NOVEBORACENSIS (L.) Nieuwl. Aspidium noveboracerise (L.)
Sw Mdst wo'odi:;nd NON: 2907 (G.N,P,M,C); NAS: Northrop

^%^lltJ^VLOSJ. (O. F. MuUer) Nieuwl. Aspidium spinvlosum (O. F.
Miller) Sw Reported by Mrs. Northrop from Nashawena; no

'PdeTstTedtia punctilobula (Michx.) .foor^^,f^^ZtrM
tilobula (Michx.) Gray. Rocky open h^'^'^es. NAS. Northrop (o),
3509 (P) ; CUT : 2534 (N,P) ; PEN : 46 (N,P,\\ ,M)

Onoclea sensibilis L. Apparently not abundant. NAS. North
rop (o); CUT: Sanford, .\ug. 15, 1917 (N).

OSMUNDACEAE

OsMUNDA REGALis L., var. sPECTABius (WiUd.) Gray. Occasional
in open bogs. NAS: AW< Arop (o), 1777 (P). ^mN- 2881 fNV

O ciNNAMOMEA L. Bogs and ^V^tholows NON. 2881 (N),
UNC: 3029 (N); NAS: Northrop (o), 3513 (P); CUT: 3450 (P).

OPHIO GLOSS ACEAE

Ophioglossum vulgatum L. Found only in y^^dy Jeld at the
east end of Cuttyhunk, near Cuttyhunk Pond. NAS. Northrop (o).
CUT : 3582 (N,P).

1930] Fogg.— Flora of the Elizabeth Islands, Massachusetts 229

[BoTRVCHiuM MATRICARIAE (Schrank) Sprcug. is reported by Mrs.
Northrop for Nashawena. This may refer to B. dusedum Spreng.
or one of its allies, but in the absence of herbarnim material it seems
best to disregard the record entirely.]

EQUISETACEAE
Equisetum arvense L. Seen only in a boggy clearing in the woods
at the east end of Naushon. NAU: 701 (P,\V).

LYCOPODIACEAE
Lycopodium inu.ndatum L., var. Bioelovii Tuckerm. Peaty mar-
gins of ponds. NON: 3381 (N); N.\U: 3865 (P).

ISOETACEAE

IsoETES Engelman.ni A. Br. A single sheet of this species collected
on Nashawena by C. E. Faxon, but bearing no date, is m the Gray

Herbarium.

PINACEAE

PiNUS Strobus L. Reported by Mrs. Northrop from Nashawena,
where probably introduced ; no specimen seen.

P R^GiDA Mill. A small group of these trees grows at the extreme
eaJt end of Naushon near the West Gutter Seen nowliere else,
althoueh perhaps more common. JN AU : 6-iZo y^y\J' ,

Ps%Lis L. Scotch Pine has been introduced on sev-e^al o^ tl^
islands but seems nowhere to be spreading. N AU : 38 a (N ) , M Ab .
Northrop (o). 3482 (P) ; PEN: 402 (W).

Larixdccidua Mill. Planted at several localities along the north
shore of Naushon. NAU: 3870 (N P,M,C).

Picea Abies (L.) Karst. P. cxcelsa Link. This and the next two
have been planted extensively along the north shore of Naushon.

""f'-gfZ^vIss. P. canadensis B. S. P. N.-VU: 3690 (J.P), 369.

(J,P), 3869 (J,N). ^j, TT •isrsr TP^

P nun a ens ^nzeim, JNAU : oc!soo i,J,r;.

bHAMSpARfs THYOIDES (L.) B. S. P. Of infrequent occurrence
on the islands, although plcntifi.1 around a large pond "e^r Tarpa.din
Cove on Naushon. NAU: » dl.ams, July 10^ 19 1 (^)',f;7j'^'„Yv on

JUNIPERUS COMMU.MS L., var. depressa Pursh Collected onlv on
the north shore of Naushon, where it may possibly have been intro-

'^TviRG^^iANfL' ^Ktiful in the woods near the East Gutter on
Nona™et; not teen elsewhere. NON: 2668 (N, P); NAS: Sorthrop

(")• TYPHACEAE

Typh A LATiFOLi A L. Swamps and pond borders. NON : 267a (N ) ;
NAS: A'or<Arop (o);PEN:463 (N,W).

230

Rhodora

[November

T. ANGUSTIFOLIA L. Mostly in brackish areas near the shore.
NON: 31C5 (N); NAU: 3887 (P); NAS: Northrop (o), 1766 CI').

SFARGANIACEAE

Sparganium eurycarpum Engelm. Found at two stations ; both
of them swampy hollows in woods at east end of Naushon. NAU.

^T i ScSi^(&im.) Morong. S. lucidum Fernald & Eames.
See FrnXm-nLAf xxivS7 (1922)^ ^^o.-^^.'^^^^^^ --
at east end of Naushon near Hadley Harbor. NAU. 2469 (N.r)

S. AMEBiCANUM Nutt. Including var. «"*««'«''"™ °f.*'of (N P )•
Swampy woods and Pond'i°les; rather common. N^^^^^
UNC: 3016 (N,P), 3113 (P); NAU: 2382 (P,C), 2383 ^JN.i-.mj, i-^^o.
3527 (N.P.M).

NAJADACEAE

POTAMOGETON Oakesianus Robbins. In a small pool bordered by
Vceodon verticillatvs along the north shore of Naushon, near the east

'"p PULCHER TucSnf The commonest Pondweed on the islands;
me^t wTtlXuIntly in the small pond holes, where it of ten conUnues

to grow upon the n.ud after the Pf"'!^, l^^f "'"^fy '" W (G N P)'
This soecies produces f niit rather abimdantly. UNC . 2997 (tr^ ,n,
3013 (PM); NAU: 711 (P); PAS: 710 (P,M); NAS: Faxort, C. E.,

"V.^r.SuRoLf^^rnald. Found growing abundant,, i„Gosnold
or West End Pond on Cuttyhunk on July 15, 1925. Ihis pond,
LmerS fresh, had evidently been inundated by the ^a 1- precedmg
winter- it has remained brackish ever smce. CU 1 : 1001 (.r, w;.

Tp PUS Llus L- i« r«P°^t«> by Mrs. Northrop for Nashawena but
no specimen has been seen. Inasmucli as P. pv.dha has been a
source of considerable confusion and there is no way of dec.dmg the
correct application of this name without material th.s record can not

'"p"Dm^RLFOUUS Raf. P. hybrld.s of the Manual, in part. Col-
lecLTnly from the sandy borders of the du.. ponds on N .Uwena,
hilt nrobablv more w dcspread. NAS: 1083 (F,W j, 1 / /o U^^^A
'"p.TeSatus L. AlJlmdant in a somew-l.at -ck.sh pond abng
the south shore of Nonamessef, should be found elsewhere. NON.

''RuPPifMARiriMA L., var. ^^-ONGiPEsHagstrSm. See Fernald &
Wiegand, Rhodora, xvi. 125 (1914). NON: 2246 (N.P.M.C). 28/6
(N) N.\U: rcnndl 3151 (W); CUT: 2,333 (G.N.P.M.C).

All of the material of R. mariiima with long podogynes (1-6 cm.)
seems to have, at least in maturity, peduncles which are well over 3
cm in length and which are often much longer and usually spu-alled.

19301 Fogg.— Flora of the Elizabeth Islands. Massachusetts 231

If var. rostrata Agardh, with mature peduncles 0.5-3 cm. long, occurs
on these islands it has still to be collected.

R. MARITIMA L., var. suBCAPir.vr.. Fernald & Wiegand. liHO'^ouA.
xvi 126 (1914). NON: 2251 (N.P); N.\U: Duggar, J«'l-V. l^H ((')•
PenneU 3152 (P,W); PAS: 712 (N,P,W).

This variety with very short podogynes (2-6 nun.) was origmally
described from material collected on Naushon. Its range has smce
been extended from Qi.ebec and Prince Edward Island to Block

Island. Rhode Island.

ZosTERA MARINA L. Frequent in the shallow coves ""'1 sniall
tidal Streams. NON: 2285 (G.N.P); NAS: horthrop (o); PEN: 464
(W), 1702 (N,P).

Material collected in Sheep Pen Harbor on Nonamesset and from
near the landing on Penikese is remarkable for the copiousness with
which it produces flowers and fruits; this fact has been observc<l
without fail for several seasons.

JUNCAaiNACEAE

Triglochin MARITIMA L. Apparently rare on the KIbabeth
Islands, although common in brackish marshes on the adjacent main-
land. NAS: Northrop, .\ug. 1901 (\ ).

ALISMACEAE

Sagitt.^ria latifolia WiUd. Not common; swampy hollow in
woods near Tarpaulin Cove. N.\U:2384 (N).

'°S .AxtFOUA^Willd., forma "astata (Pi^sh) Robinson, l-orin
with very narrow, acute leaves. U M U : 3UU.i U> )■

HYDROCHARITACEAE

VaLLIS.NER.A AMERICANA MicllX. ^ ^^'^ ''■^ »f., ^Y''"""' ^ Jf.

Fernald. Rhodora. xx. 108 O^IS). -^bundant in Sheep Pond
near west end of Cuttyhunk. CUT : 2o32 (G.N ,P,M).

GRAMINEAE

Andropogon SCOPARIU8 Michx., var. frequens Hubbard See
Rhodora, xix. 103 (1917). Abundant eyeryw^iere «" "P^" ' J^ /^"^
sandv lowlands. NON : 2585 (P.C) . 291 1 (N.P.M) : UN C . 31 09 (N ,
NAU: 3924 (N.P.M); PAS: Sv,„so„, Sept. 8, 1926 (N), 3<o3 (I).
CUT:San/orrf, .\iig. 15, 1917(N). i;tioral!'<

\ srnPAnnis Michx., var. polyclados Scribn. & Hall. A.litiorau.i
Nash T.'o;i"t^ li"oralls (Nash) Hitchc See Rhodora

xix 103 (1917). Collected on the white sand beach bordering \\est
End Pond on Naushon. NAU: 2940 (G.N.P).

232

Rhodora

[November

A. GLOMEEATUS (Walt.) B. S. p. Seen only in a low brackish
mareh near the east end of Naushon; probably more abundant. NAU .

^^f. ^^faimcus L. Open grassland and por.d borders: material
collected on Nonamesset has conspicuously hairy lower leaves and
sheaths. NON: 3915 (N.P.M.C); UNC: 3118 (N).

A FURCATUS Muhl. Sandy fields and clearings; not common.
UNC- 3011 (N,P); NAU: 2603 (N.P).

Digitaria sanguinalis (L.) Scop. Disturbed sandy areas. UNC:
3107 (P,M). 3053 (N,P); PEN: Jorrfore(o). ,_ . ,, ..

PaspIlum setaceum Michx. Found growmg abundantly on the
exposed grassy slopes at the extreme east end of Nashawena. NAb.
3549 (N,P).

This grass, rare in New England, is known elsewhere in Massa-
chusetts only from Nantucket. See Weatherby, Rhodora, xxx. 133

fl928)

P PUBESCENS Muhl. Including P. Muhlenbergii Nash. See
Weatherby, Rhodora, xxx. 134 (1928) Abundant everywhere o^^
open grassy hillsides. UNC: 2967 (N.P), 3642 (P); NAU: 2599 (N).
3885 (P,C); CUT: Sanford, Aug. 15. 1917 (N).2538 (N,P,M).

Panicum capillare L. Seen growing only along border of salt
marsh at west end of Nonamesset. NON : 2899 (N,P,M).

P. DiCHOTOMiFLORUM Michx. Occasional m moist grassy hollows.

NON: 3197 (N,P). . . , „ • ia nQ99^

P viRGATUM L., var. SPissuM Lmder. Rhodora, xxiv. 14 (1922).
This variety with densely tufted culms seems entirely to replace the
snecies on the Elizabeth Islands. Common in open grassland^ ^e*
hollows and sandy clearings. NON: 2632 (P); UNC: 3030 (N,P,M);
NAU73908 (N,P); PAS: 3763 (P,C), 3786 (N,P,M); NAS: 2360

^V^lo^gSI^ouum ToTt This species, known elsewhere jn Massa-
chusetts only from Marion. Plymouth County, was f "e<=tf in the
peaty and Jggy ^oUowsat the east end of Pasque on Au^st 28, 1928.
See Fogg, Rhodora, xxxi. 39 (1929). PAb. .J7S/ U>.J^.*»^

P dIIIuperatum Muhl.. var. psilophvllum Fernald. Rhodora
xxiii. 193 (1921). This species is represented on the islands only by
the smooth-leaved variety. Grassy and sandy slopes. NON . 2579

(N) ; NAU: 2491 (N,P), 3319 (P).
P dichotomum L. Mostly in open sfndy woods. NON. 2650

(N). 3423 (P.M) ; NAU: 2478 (N,W), 2483 (P). 3906 (N.P.C).

^ P. LiNDHEiMERi Nash. var. fasciculatum (Torr.) Fernald.

P. huachv^ae .\she. P. huachu^ae y>iT^ sihicola Hitchc & Chase.

P tm-npiirense Ashe See Fernald. Rhodora. xxiii. 223 (19^U.

NON 2295TNPW) 2301 (W.C). 2589 (P). 2633 (P.M). 2648 (P.W .

Vfifin rKH -iiVfi /f^ 3374 (PU) 3375 (P). 3376 (P,U), 3420 P.U);

nTuSs? (N.P?3S (PUh CUT: I2& (N.P.M), 3433 (P.U.C).

3456 (P,U) ; PEN : 478 (N,P,W) , 3405 (P) .

19301 Fogg.— Flora of the Elizabeth Islands. Massachusetts 233

This is the most ubiquitous and the most variable Panicum on the
islands. Of the specimens cited Mrs. Agnes Chase, who has kindly
examined some of the material from the Elizabeths, has designated
no 3374 as P. implkatum Scribn.. no. 3420 as typical P. huachucae
Ashe, and nos. 3376. 3433. 3456 and 3903 as P. huachucae var. s.h,-
cola Hitchc. & Chase. In speaking of no. 3420 Mrs. Chase says.
"The pubescence on upper surface of blades varies from very scant
or almost wanting to fairly copious." Thus, degree of pubescence
varies, not only as between separate plants, but in different parU o
the same plant and. in view of the many intermediates which make it
impossible here to draw sharp specific or even varietal lines, it seems
best to follow the more conservative treatment of this species pro-
posed by Fernald, 1. c. p. 226. Viewed in this light, the Elizabeth
Islands material with spikelets 1.5-2.1 mm. long and leaf-blades
short-pilose to glabrous above, falls into the y^xriety fasaculatum

P. MERiDio.NALE Ashe. Abundant on «=''P°^«^. 8'"f?>:N sSS"^
Often very slender with leaves not over 1.5 mm. wide NON. 258-
(P)- NAU: Taylor, July 24. 1917 (P); NAS: 2452 (N). 35.>0 (:^,f),

'^"p'^SMlYElsSe'-'^'Ti'^southern species, found abundantly
inLent years on the "Middle" Cape, is known from two stations on
the ElSabeth Islands, both of them exposed sandy areas. LNC : 3082

borders. NAU: 718 (W), 2492 (P). 2714 (N). 3081 (P). N.Vb. -35J
(N) . 3466 (P) ; PEN : 479 (P.W).

The lines between this and P. meridionale are not always sharp;
in fact, the two behave more like varieties of one polymorphic species_
Differences in size of spikelets are trivial, pubescence cannot be relied
upon, and habit varies with the situation in which these plants grow.
To be sure, there is a well marked meridionale extreme and an equally
characteristic oricola one, but between them an almost complete

series of integradations.

P. COMMONSIANUM .\she. Met with Only on an open sandy bank
along the north shore of Naushon. west of Kettle Cove. NAU. 387.->

^YiHAEROCARPON Ell. Occasional on open grassy downs. NON:
'"? 'S^i^i^^i^tS^s and clearings; not common.

UNC: 3063 (N); NAS: Northrop (o); PEN: 1092 (F, W).

234

Rhodora

[November

E Walteri (Pursh) Nash. Moist, usually brackish, situations.
NON: 3188 (N); UNC: 2992 (N); NAU: 3156 (C), 3888 (P,M,C).

Setaria geniculata (Lam.) Beauv. See Hitchc. Contr. U. S.
Nat. Herb. xxii. 168 (1920). Open grassland and around borders of
brackish areas. NON: 2695 (N,P); UNC: 3033 (N,P), 3103 (N,P).

This species is in urgent need of further study, for even a casual
examination of a large series of specimens is sufficient to indicate
that our perennial Foxtail of the salt marshes of New England is
distinct from the far-ranging southern plant which occurs in tropical
and subtropical America.

S. glauca (L.) Beauv. See Rhodora, xxxi. 109, 110 (1929). Dis-
turbed sandy area near beach. UNC : 3098 (N).

S. mridis (L.) Beauv. Reported by Jordan from Penikese; no

specimen. ^

Cenchrus pauciflorus Benth. C. carolinianus of Manual, bee

Chase, Contrib. U. S. Nat. Herb., xxii. 67 (1920). Seen only in sandy

field near east end of Basque. PAS: 3766 (N,P,M,C).

[C. TRIBULOIDES L., reported by Mrs. Northrop from Nashawena,

probably refers to the preceding. C. tribnloides is unknown east of

Long Island.] i n t^ c

Leersia oryzoides (L.) Sw., forma inclusa (Wiesb.) I^ogg. bee
Rhodora, xxx. 81 (1928). The only material of the Rice Cut-grass
seen on the islands grew on the sandy margin of Mary's Lake and had
the panicles enclosed within the upper leaf-sheaths. NAU: 313o

(G,N,P,M). , ^. . , ,

Phalaris canaricnsis L. Sandy area near landmg at east end ot

Pasque. PAS: 1794 (P). .

Anthoxanthum odoratum L. Everywhere m open grassland. JNUiN :
2250 (N,P,M); NAU: 3315 (P,C); PEN: 468 (N,P,W,M)

Stipa avenacea L. Apparently rare and local. NAS: Northrop

(o), 3560 (P). , J .... . .

Aristida purpurascens Poir. Abundant on exposed hillsides,
especially on Naushon. NAU: 1010 (P,W), 3140 (N), 3683 (P).

Phleum matense L. Seemingly only on outer islands, where abund-
ant on open slopes. CUT: 2280 (N); PEN: 480 (W).

Agrostis stolonifera L. a. alba of most American authors, not
L. See Make, Canada Dept. Mines, Bull. no. 50. (1926). Grassland
and sandy fields. NON: 2653 (N), 3418 (P); PAS: 3772 (P); NAS:
Pennell 2914 (P) ; PEN: 3390 (N,P).

A. STOLONIFERA L., var. coMPACTA Hartm. A. alba, var. mantima
(Lam) Mey. See Malte, 1. c. Beaches and brackish situations.
NON: 2721 (P), 3417 (N); NAU: 2713 (N); PEN: 3397 (N,P,M,C).

A TENUIS Sibth. A. vulgaris With. A. alba, var. mlgans (With.)
Thurb. Doubtfully A. cavillaris L. See Malte, 1. c. Dry hillsides
and hollows. NON: 3352 (P); PEN: 466 (P,W), 3391 (P).

1930] Fogg,— Flora of the Elizabeth Islands, ]Massachusetts 235

A HYEMALis (Walt.) B. S. P, Sandy woods; occasional. NAU:

Pennell 3124 (W,P), 3918 (N). , , , •

A pkrennan's (Walt.) Tuckerm. Sandy .x,odsa„dc^^^^^^^^^^^
more frequent than the preceding. NAU: 26/9 (P), 3881 {!S,f),

^^JiULoKOSTls CANADENSIS (Michx.) Beauv. Wet hollows and
border^ of ponds. NON: 3359 (N); NAS: 1007 (P.W), l^dhams,
Julv 10. 1911 (N); CUT: 3453 (F.M).

C cmNOiDES (Muhl.) Barton. Seen only in a boggy thicket near
Job's Neck. NAU: 2604 (N.P).

Ammophila breviligulata Fernald. A. armaria of Manual.
See Fernald. Rhodora, xxii. 70 (1920) Cobble and sandy beaches.
NON : 2698 (N,P,M.C) ; UNC : 3058 (N) ; PEN : 407 (W)

Hnlrv., lanaius L Ubiquitous in grassland and sandy fields and
cleaS. NON: 2291 (h); NAU: 2709 (N.P): ^NAS: f oHArop
(o) • CUT: Tayhr 2408 (P , 3455 P.C) ; PEN: 4/7 (N.F.W.U)-
^ Deschampsia flexuosa (L.) Trin. Dry woods near Tarpaulin

Cove. NAU: 1009 (N,P,W) PFN.4r.qa\n

ATcna saliva L. Sandy soil near cultivated area. PEN • 409 (VV )•
Danthonia spicata (L.) Beauv. Abundant on dry knolls and
opSi Tearing. NON: 2249 (N.P). 2658 (P); PEN: 473 (P.W).

Several trends are evident in material of this species from these

islands and elsewhere in eastern North America. Specimens collected

from the beach near French Watering Place. Naushon for example,

have lemmas with the aristate teeth of D. sericea, although lacking

the pubescence of that species. This material can be satisfactorily

referred to nothing included in our existing treatments ot the genus.

It is hoped that the intensive study of our American material already

embarked upon by the wTiter may lead to a better undersUnding of

these puzzling forms. A large series of collections « being held for

critical examination and comparison with types, but the preceding

may be referred to as fairly typical of D. spicata as it is commonly

understood. ,

(P) 2910 (N,P,M); UNC: 3095 (N,P); NAU: 36So (N), PAb. bvenson,

236

Rhodora

[November

ing only on sandy upper beach on Uncatena. NON: Cushman 126
(B);UNC:3049(N).

S. PATENS (Ait.) Muhl., var. caespitosa (A. A. Eaton) Hitchc.
This variety, characterized by its densely tufted habit, was collected
in a brackish marsh near Tarpaulin Cove, where it formed a dense
turf. NAU: 2377 (N,P).

DiPLACHNE MARiTiMA Bicku. Bull. Torr. Bot. CI. xxxv. 195 (1908.)
Found only along a brackish ditch near Tarpaulin Cove. NAU:
3889 (N,P,M).

Phragmites communis Trin. Occasional in swampy situations.
NAU; 1803 (N,P,W), 2608 (P); NAS: W. Faxon, Aug. 1873 (G),
Northrop, Aug. 1901 (Y).

Triplasis purpurea (Walt.) Chapm. Abundant on beaches and
low sandy stretches. NON : 2871 (N,P,W,M,C) ; UNC : 3009 (N,P,M) ;
NAU: 2510 (N), 2732 (P); PAS: 3754 (P).

Eragrostis pectinacea (Michx.) Steud. Common everywhere
on open grassland and in sandy clearings, except on the outer islands.
NON: 2580 (P,W), 3167 (N); UNC: 3651 (P); NAU: 3919 (N,P,C);
PAS: 3765 (P,M); NAS: Northrop, Aug. 1901 (Y).

An intensive search has failed completely to reveal var. spectabilis
Gray with smooth sheaths, which is abundant on Martha's Vine-
yard and Nantucket.

DiSTiCHLis SPiCATA (L.) Greene. With Spartina patens in brackish
marshes. NON: 2901 (P), 3208 (N); UNC: 2694 (P), 3097 (N);
PAS: 3737 (P,M); PEN: 1091 (P,W).

Dactylis glomerata L. Sandy soil near cultivated area on Penikese;
otherwise not seen, although probably more general. PEN: 472
(N,P,W,M).

Poa annua L. Not common. Disturbed sandy areas on two of
the islands. N AU : 33 1 1 (P) ; PEN : 3389 (P) .

[P. SEROTiNA Ehrh. Reported by Jordan from Penikese. This
may refer to P. palustris L. (P. triflora Gilib.), but, in the absence of
material, it seems best to disregard the record.]

P. PRATENSis L. Fairly common on open downs. NON: 3249
(N,P), 3351 (P); PEN: 481 (P,W), 3388 (N,P,M).

Glyceria obtusa (Muhl.) Trin. Pond borders and moist hollows.
NAU: 2934 (N,P); NAS: Northrop, Aug. 1901 (Y), 2348 (N,P,C).

G. CANADENSIS (Michx.) Trin. Border of Decodon swamp near
Tarpaulin Cove. NAU: 2396 (N,P).

G. STRIATA (Lam.) Hitchc. G. ncrvata (Willd.) Trin. Boggy
woods; apparently local. NAU: 2479 (N).

G. PALLIDA (Torr.) Trin. Swampy woods and wet hollows. NON:
2575 (N,P), 3358 (P), 3371 (P); NAU: 2470 (P); NAS: 3525 (N,P,M,
C).

G. ACUTiFLORA Torr. Found almost completely submerged in
several small ponds on Nonamesset and one on Nashawena. NON:
2243 (G,N), 3344 (P), 3380 (P,C); NAS: 3524 (N,P,M).

11

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 237

[PucciNELLiA MARITIMA (Huds.) Pari. Reported by Jordan from
Penikese but probably refers to the following which, though unknown
from Penikese, grows on the neighboring island of Cuttyhunk.]

P. PAUPERCULA (Holm) Femald & Weatherby, var. alaskana
(Scribn. & Merr.) Femald & Weatherby. Rhodora, xviii. 18 (1916).
Abundant on mud flats bordering Cuttyhunk Pond. CUT: 3572
(N,P,W,M,C).

Festuca Myuros L. Edge of sandy road leading to Tarpaulin Cove.
NAU: 2509 (N,P).

F. octoflora Walt. Apparently not general. NON: Pennell
2847 (W).

F. RUBRA L. One of the commonest grasses on the islands. Gravel
banks, borders of beaches, open downs, sandv clearings, etc. NON:
2845 (W,P), 3349 (P), 3350 (N,P), 3370 (P^, 3416 (P); NAU: 2712
(N,P,M), 3877 (N,P,C); CUT: 2279 (N,P); PEN: 476 (N,P,W).

Extremely variable as to habit, texture of foliage and degree of
glaucousness.

F. oviNA L. Reported by Mrs. Northrop from Nashawena and by
Jordan from Penikese. No material seen, but this record is allowed
to stand for the present.

F, elatior L. Found only on Penikese, where abundant in certain
areas. PEN: 475 (P,W).

Bromiis secalinus L. Growing in dry sandy ground along east
shore of Penikese. PEN : 470 ( W) .

B. hordeaceus L. Disturbed sandy areas on Nonamesset and along
path near landing on Penikese. NON: Pennell 2839 (W), 3304
(N,P), 3347 (P,M,C); PEN: 3412 (N,P).

Agropyron repens L. Apparently not abundant except on outer
islands, where it takes possession of large sandy areas. CUT: 2278
(N,P,M); PEN: 465 (W), 3387 (N,P).

Elymus virginicus L. Collected only on Gull Island, Penikese,
where the plant is unusually robust. Reported by Jordan from
Penikese proper, but not seen there recently. PEN: 474 (N,P,W).

E. virginicus L., var. halophilus (Bickn.) Wiegand. Rhodora,
XX. 83 (1918). Dry sandy banks facing the sea at west end of Un-
catena. UNC: 3119 (N), 3648 (P).

CYPERACEAE

Cyperus diandrus Torr. Wet depressions and pond borders.
NON: 3198 (N); UNC: 3021 (N); NAU: 2737 (N,P,W).

C. rivularis Kunth. Seen only growing with the preceding on
grassy bank bordering French Watering Place. NAU: 2739 (N);
NAS: Northrop (o).

C. FiLiciNUS Vahl. C. Nuttalln Eddv. Borders of fresh ponds and
low brackish areas. NON: 2700 (N,P), 2889 (P); UNC: 3007 (N,P),
3048 (P); NAU: Cushman 313 (B), 2725 (N), 3864 (P).

238

Rhodora

[November

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 239

I'

C. DENTATUS ToFF. Not common; sandy margins of several ponds
on Naushon. NAU: 2467 (C), 2920 (P,M), 2935 (N,P,M).

C. ERYTHRORHizos Muhl. Reported by Mrs. Northrop for Nasha-
wena; no specimen seen.

C FERAX Rich. Borders of brackish marshes and low swales; not
common. NON: 2897 (N); NAU: 2369 (G,P,C), 3155 (N,P,W,M).

C. STRIGOSUS L. Of rather frequent occurrence in pond-holes,
borders of salt marshes, upper beaches, and moist clearings. NON:
2665 (P); UNC: 2982 (P), 2991 (N,P), 3057 (W); PAS: 3781 (P).

C. STRIGOSUS L., var. robustior Kunth. Moist depression in
open hills. UNC: 3083 (N,P,W).

C. STRIGOSUS L., var. compositus Britton. Small pond-hole at
west end of island. UNC : 3020 (N,P,W).

C. FiLicuLMis Vahl, var. macilentus Fernald. Dry open ground
and sandy clearings everywhere. NON: 2587 (N); UNC: 3061 (N);
NAU: 742 (P,W); PAS: 3769 (N); NAS: Northrop, as "C. filicuhms''
(o), 3553 (P); CUT: Sanford, Aug. 15, 1917 (N), 2265 (P,M).

DuLiCHiUM ARUNDiNACEUM (L.) Britton. Swampy hollows.
NAU: 745 (W); PAS: 744 (W); NAS: Northrop, Aug. 1901 (Y), 2343
(N,P); CUT: Sanford, Aug. 15, 1917 (N).

Eleocharis parvula (R. & S.) Link. Scirpus nanm Spreng.
See Svenson, Rhodora, xxxi. 168 (1929). Brackish ponds and salt
marshes. NON: 2287 (P,M), 2708 (N,P); UNC: 3050 (P); NAU:
1804 (N); NAS: Pennell 2924 (W).

E. OBTUSA (Willd.) Schultes. Moist hollows and pond margms.
NON: 2574 (P,M), 3204 (N,P); NAS: Northrop, Aug. 1901, labelled
"E. ovata'' (Y); CUT: Sanford, Aug. 15, 1917 (N); PEN: 485 (W).

E. ACicuLARis (L.) R. & S. Peaty and sandy pond borders:
apparently not common. NON: 2913 (N,P); NAU: 2507 (N,P,M);
PEN: Jordan (o).

E. Smallii Britton. See Fernald & Brackett, Rhodora, xxxi.
57-77 (1929). Peaty and boggy pond margins and low swampy
areas. UNC: 3002 (N,P); NAS: 3498 (N,P,M,C); CUT: 746 (P,W);
PEN: 849 (P,W).

E. UNiGLUMis (Link.) Schultes. See Fernald & Brackett, 1. c. 71.
This essentially boreal sedge is near the extreme southern limit of its
range on Cuttyhunk where, on August 11, 1927, it was collected from
the peaty margin of Sheep Pond. CUT: 2526 (N).

E. UNIGLUMIS (Link) Schultes, var. halophila Fernald & Brackett,
1 c. 72. Sandy or peaty margins of brackish ponds; rather frequent.
NON: 2703 (N,P,M,C), 2912 (P,M); NAU: 1028 (N,P,W), 3316 (P);
NAS: Williams, July 10, 1911 (N), 1029 (P,W).

E. ROSTELLATA Torr. Apparently infrequent. Seen only from a
low brackish marsh along the north shore of Naushon near Kettle
Cove. NAU: 3878 (N,P,M,C).

FiMBRiSTYLis AUTUMNALis (L.) R. & S. F. FrankH Steud. See
Blake, Rhodora, xx. 25 (1918). Grassy pond borders. NON: 3199
(N,P); NAU: 2738 (N,P),

SciRPUS AMERICANUS Pers. Everywhere bordering ponds, both
fresh and saline. NON: 2239 (P), 2888 (M); UNC: 2998 (P), 3025
(N); NAU: 2375 (P), 2921 (N), 2923 (P), 2723 (P,C); NAS: Northrop,
Aug. 1901 (Y), Fenncll 2910 (W), Williams, Julv 10, 1911 (N), 2355
(P,M); CUT: Sanford, Aug. 15, 1917 (N), 2277 (P,M); PEN: 486
(P,W).

S. Olneyi Gray. Seen growing at edge of salt marsh along south
shore of Nonamesset; probably more general. NON: 2898 (N,P);
NAS: Northrop, Aug. 1901 (Y).

S. VALIDUS Vahl. Common in wet depressions and swampy
borders of ponds. NON: 2244 (N,P); UNC: 2999 (N,P); NAU:
1033 (P,W); NAS: Northrop, as " S. lacustris'' (o); CUT: 3573 (P);
PEN: 488 (N,P,W,M).

S. ROBUSTUS Pursh. Not common. NAS: Northrop, Aug. 1901 (Y)

S. CAMPESTRis Britton, var. paludosus (A. Nelson) Fernald.
Occasional in brackish marshes. NON: 2288 (P,M), 3209 (N,P);
PEN:487(N,P,W,M,C).

S. CAMPESTRIS Britton, var. \ovae-a\gliae (Britton) Fernald.
Collected only from swampy clearing in woods near Job's Neck.
NAU: 3154 (N,P).

S. CYPERINUS (L.) Kunth. Everywhere in wet depressions and
around ponds. NON: 2566 (N,P), 3169 (P,M); UNC: 2990 (N,P);
NAU: 2922 (N,P,M), 3149 (P); PAS: Svcnsoji, Sept. 8, 1926 (N),
3791 (C); NAS: Northrop (o), 2340 (N,P); CUT: 2308 (N,P).

S. CYPERINUS (L.) Kunth, var. pelius Fernald. Found only along
the edge of a swamp near Tarpaulin Cove. NAU : 2395 (N).

Eriophorum tenellum Nutt. Occasional in boggv situations.
NAU: Pennell 2905 (P); NAS: Northrop, Aug. 1901, 'labelled "£.
gracile " (Y) ; CUT: 1030 (N).

E. virginicum L. Rather frequent in open bogs and wet peaty
hollows. NON: 2879 (N,P,M), 3180 (P); PAS: 3792 (N,P), Svenson,
Sept. 8, 1926 (N); NAS: Northrop (o), 3497 (P); CUT: Sanford, Aug.
15, 1917 (N), 1014 (P,W), 2517 (P,C).

Rynchospora fusca (L.) Ait. f. Apparently rare; seen only in a
low, boggy hollow near Tarpaulin Cove. NAU: 2512 (P,W).

R. ALBA (L.) Vahl. Open bogs and peaty depressions; seemingly
not abundant, although plentiful in an extensive bog at the west end of
Cuttyhunk. NAS: Northrop, Aug. 1901 (Y); CUT: 2319 (N,P).

R. CAPITELLATA (Michx.) Vahl. R. glomerata of Manual in part.
See Blake, Rhodora, xx. 25 (1918). Much more common than the
preceding. Bogs, pond-holes and pond borders. NON: 2644 (N);
UNC: 3014 (N,P), 3116 (P,M,C); NAU: 2393 (N); PAS: 1782 (N,P);
NAS: 3471 (P).

Mariscus mariscoides (Muhl.) Kuntze. Cladiinn mariscoides
Torr. See Rhodora, xxv. 49 (1923). Mostlv on sandy beaches
bordering ponds. NAU: 2924 (N); PAS: 3776 (P); NAS: Pennell
2917 (W), 2339 (N); CUT: 740 (P,W).

240

Rhodora

[November

Carex scoparia Schkuhr. Not common on the Elizabeth Islands.
Dryish border of small pond. NON: 3384 (N,P,M,C) ; PEN: Jordan

(o).

C. SCOPARIA Schkuhr, var. subturbinata Fernald & Wiegand.
Rhodora xiv. 115 (1912). Seen around margin of Dccodon swamp
near Tarpaulin Cove. N AU : 2399 (N) .

C. LoNGii Mackenzie. C. alholutescens of Manual. See Macken-
zie, Bull. Torrey Bot. Club, xlix. 372 (1922). Frequent in open
grassland. UNC : 3078 (N) , 3099 (?) ; N AS : 3469 (P) ; CUT : Sanjord,
Aug. 15, 1917 (N), 1013 (N,P,W,M); PEN: 848 (N,P,W).

C. STRAMINEA Willd. This is the plant v^^ith obovate perigynia
selected by Mackenzie as true C. straminea; see Bull. Torrey Bot.
Club, xlii. 603 (1915). It has been found growing only on the edge
of a bog at the west end of Cuttyhunk. CUT: 2327 (N,P).

C. HORMATHODES Fernald. Moist, occasionally brackish, situa-
tions as well as dry slopes near shores. NON: 2297 (N); UNC:
3644 (P); NAS: Williams, July 10, 1911 (N); CUT: 726 (P,W); PEN:
1754 (N,P).

C. SILICEA Olney. Dry, sandy banks and hillsides. PAS: 1780
(N,P); CUT: 2258 (N), 2267 (P,M,C); PEN: 1497 (P), 1498 (N,P,W).

C. ALATA Torr. Plentiful around edge of an extensive boggy area
near the center of Nashawena. NAS: 2486 (N,P).

C. FESTUCACEA Schkuhr. This is the small-fruited sedge retained
by Mackenzie as genuine C. festucacea. See Bull. Torrey Bot. Club,
xlii. 604 (1915). Apparently rare and local on the islands. NAU:
736 (N,P).

C. HowEi Mackenzie. C. scirpoidcs, var. capillacca (Bailey)
Fernald. See Mackenzie, Bull. Torrey Bot. Club, xxxvii. 245 (1910).
Abundant in boggy woods and pond borders. NON: 2654 (N,P,M),
2659 (P), 3377 (C), 3299 (P); NAU: 728 (W), 1018 (P,W), 1088 (P,W);
NAS: Pennell 2922 (W), 3542 (N,P) ; CUT: 727 (P,W).

C. INCOMPERTA Bickuell. Bull. Torrey Bot. Club, xxxv. 494 (1908).
NAU : Pennell 2907 (G,W) .

C. SEORSA E. C. Howe. Seen growing only in a boggy wooded
hollow on the south shore of Nonamesset. NON: 3379 (N,P).

C. CEPHALANTHA (Bailey) Bicknell. C. stellulata, var. cephalantha
(Bailey) Fern. SeeBull.Torrey Bot. Club, 1. 346(1923). Boggy woods
at east end of Nashawena. NAS: 3643 (P).

C. CANESCENS L., var. DisJUNCTA Femald. Boggy woods and wet
borders of ponds. NON: 3378 (N), 3398 (P); NAU: Williams, July
10, 1911 (N), 1019 (P,W).

C. BRUNNESCENS Poir., var. sphaerostachya (Tuckerm.) Kiikent.
See Fernald, Rhodora, xxviii. 162 (1926). Rare. Boggy hollow in
woods at east end of Naushon, NAU : 2480 (P, W) .

C. ROSEA Schkuhr, var. radiata Dewey. Apparently local.
Collected in dry sandy woods at east end of Naushon. NAU: 2475
(P,W).

V

' 11 ^

4 > k

ft

/-V V

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 241

C. contigua Hoppe. C. muricata of most American authors. See
Mackenzie, Bull. Torrey Bot. Club, 1. 236 (1923). Dry, open,
sandv ground. NAU: 734 (P,\V) ; CUT: 22S3 (N,P), Taylor 2413 (P) ;
PEN:484(P,W).

C MuHLENBERGii Schkuhr. Rather frecjucnt on exposed downs
and in sandv clearings. NON: Pennell 2838 (W), 2245 (N), 3361
(P,M);NAU:733(W). ^ i i

C. CEPHALOPHORA Muhl. Sandy oak and beech woods; local.
NON: 2594 (N,P,W); NAU: 2463 (P).

C. vulpixoidea Michx. Wet depressions. NAU: 2394 (N,P,M,

C); NAS: 3485 (N). ^ ^

C. LAEVIVAGINATA (Kukeut.) Mackenzie. See Bntton & Brown,
111. Fl. ed. 2, i. 371. (1913). Low, wet hollows; local. NAS: 3473
(P); CUT: 2281 (N). .

C. CRiNiTA Lam. Swampy and boggy depressions. INAU: U,i
(W) ; 2398 (P) , 2936 (N) ; NAS : 3475 (P) .

C. MiTCHELLiANA Curtis. See Weatherby, Rhodora, xxv. 17
(1923). Rare on the Elizabeths. NAS: Pennell 2921, labelled "C.
crinita'' (W).

A species of the southern coastal plain ranging north to Cape Cod
and Plymouth County.

C. viRESCENS Muhl. Boggy woods; rare and local. NAU: 1020

(P W) 3141 (N,P,M).

C. SwANii (Fernald) Mackenzie. Bull. Torrey Bot. Club, xxxvii.
246 (1910). C. virescens, var. Swanii Fernald. Dry, open ground
and sandy woods. NON: 2588 (N,P). 3354 (P,C); NAU: Pennell
3132 (W), 3680 (N); CUT: 1012 (N.P,W,M), 3452 (P).

C. COMMUNIS Bailey. Dry, sandy beech woods; rare. JNAU:

3324 (N,P).

C. VARIA Muhl. Sandy clearings, exposed knolls and hillsides;
widespread. Some of the material approaches the scarcely separable
forma co/ora<a (Bailev) Kukent. NON: 3284 (N,P.C), 3300 (N,P),
3301 (P), 3285 (P); NAU: 3322 (N,P); NAS: 2361 (N,P,W,M).

C PENNSYLVANICA Lam. Sandy banks and open grassland. AUiN :
3283 (N,P), 3353 (P,C); NAU: 3321 (N,P,M). ^ . ^ ^ „ ^ , ,

C. PENNSYLVANICA Lam., var. separans Peck in L C. Ho^^^, IN. 1 .
State Mus. 48 Ann. Rep. 174 (1896). Var. ^"^^/""^ (^^"^^^.^f "^^^^^^
Seen only in dry beech woods at east end of Naushon. NAU: 24/b

(P W M C).

C LIMOSA L. A far northern sedge which reaches the southern
limit of its coastwise distribution in a bog at the west end of Cuttyhunk.

CUT* 2311 (N,P,W).

C DIGITALIS Willd. Apparently not common; sandy beech woods
near Hadley Harbor. NAU: 2461 (P), 3323 (N,P).

C DEBiLis Michx., var. Rudgei Bailey. Collected only m a moist
hollow on the hillsides of Cuttyhunk. CUT: 101 1 (P).

242

Rhodora

[November

C. LANUGINOSA Miclix. Peaty margin of a slightly brackish pond
at the extreme west end of Nashawena. NAS: 3500 (N,P).

C. COMOSA Boott. Swampy clearings and wet depressions; locally
abundant. NON: 2905 (N,P); CUT: Sanford, Aug. 15, 1917 (N), 722
(W).

C. LURIDA Wahlenb. Pond borders and moist hollows. UNC:
2P89 (N,P), 3646 (P,C) ; NAU: 2385 (N,P,M) ; NAS: 3474 (N,P).

C. LUPULINA Muhl. Boggy clearings and low moist situations.
NAU: 2486 (N,P), 3882 (P,M,C); PAS: 519 (P,W); CUT: 731 (P,W).

ARACEAE

Arisaema triphyllum (L.) Schott. This seems to be the only
Arisaema on the islands. It is abundant in the boggy and swampy
woods at the east end of Naushon, and in similar situations on Nasha-
wena. NAU: 752 (N), 3133 (W), 3317 (P,M,C), 3331 (P); NAS
Northrop (o), 3528 (N,P).

Symflocarfus foetidus (L.) Nutt. Seen only in a low swampy
area at the extreme west end of Nashawena. NAS: Northrop (o), 3496
(N,P).

AcoRUS Calamus L. Wet grassy hollows and swampy pond bor-
ders; locally abundant. NAS: Northrop (o), 3565 (P); CUT: 2282
(N).

LEMNAGEAE

Lemna minor L. Occasional in small ponds in clearings. NAU:
2376 (N,P).

ERIOCAULACEAE

Eriocaulon septangulare With. K. articidatum (Huds.) Morong.
Sandy and peaty pond margins. NAU: Pcnnell 3118 (P), 2508
(P,M), 2942 (N); NAS: Northrop, Aug. 1901 (Y).

XYRIDACEAE

Xyris caroliniana Walt. Apparently not sommon. Sandy
beach bordering West End Pond, Naushon. NAU: 2919 (N,P);
NAS: Northrop, Aug. 1901 (Y).

X. TORTA Smith. X. flcxnosa Muhl. Reported by Mrs. Northrop
from Nashawena; no specimen seen.

PONTEDERIACEAE

Pontederia cordata L. Collected only in swampy border of
dune ponds on Nashawena. NAS: Northrop (o), 2368 (N,P), 3547
(P).

JUNCACEAE

Juncus bufonius L. Moist sandy or peaty soil. NAS: Northrop,
Aug. 1901 (Y), Williams 1279 (N), Penncll 2918 (W), 3467 (P,M);
CUT: 3575 (N,P).

V

* > •

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 243

J. Gerardi Loisel. Brackish areas; general. NON: 2877 (N);
PAS: 1779 (P); NAS: Northrop, Aug. 1901 (Y); CUT: 2276 (N.P,M,
C); PEN: 1094 (P.W), 1753 (N,P).

J. tenuis Willd. Common everywhere; dry woods, gravelly
shores, sandy paths and open fields. NOX: 3191 (N); UNC: 3(>45
(P); NAU: 3136 (N); NAS: Northrop, Aug. 1901 (Y), 3468 (P); CUT:
3441 (N,P,M); PEN: 492 (P,W), 3400 (P).

Much of the material which has been collected can not satisfactorily
be referred here, but is being held pending a critical comparison of
our American J. tenuis with Old World collections, a comparison
which may necessitate a careful revision of the entire species. The
preceding few numbers are, however, cited as fairly typical.

J. tenuis Willd., var. Williamsii Fernald. This phase, \\\i\\
strongly divergent floriferous branchlets, has been met with occasion-
allv. NAU: 2400 (N), 2710 (P,M).

J. DiCHOTOMUS Ell. NAU: 1800 (P,W), 3162 (P); PEN: 1756
(N,P,W).

Under this heading is groui)etl a large series of variations, of which
the variety platyphijUus, i)erhaps specifically distinct, represents
onlv one trend. This matter is under consideration at present, and,
until the species can be completely studied and revised, a few numbers
only, out of an extensive series of collections, are cited.

J. DICHOTOMUS Ell., var. platyphyllus Wiegand. PAS: 17S1 (P);

CUT: 1039 (P).

J. Greenei Oakes & Tuckerm. This is one of the commonest
plants of the dry open hillsides, being found on all of the islands.
NON: 2247 (N,P,M,C), UNC: 3130 (P); NAU: 766 (P,W), 1802 (P);
PAS: 765 (W); NAS: Northrop, Aug. 1901 (Y), 2362 (N); CUT:
Sanford, Aug. 15, 1917 (N); PEN: 491 (N,P,W,M).

J. effusus L., var. costulatus Fernald. Rhodora, xxiii. 239
(1921). Abundant in wet hollows and around ponds. NON: 2303
(P,M,C), 2567 (N,P); NAU: 2397 (P); PAS: Svenson, Sept. 8, 1926
(N); NAS: Northrop, Aug. 1901 (Y), Williams, July 10, 1911 (N),
1767 (N,P), 2347 (P); CUT: 2320 (N,P,M); PEN: 852 (P,W).

J. EFFUSUS L., var. solutus Fernald & Wiegand. Rhodora, xii. 81
(1910). Collected bv the writer only in a boggy hollow in woods near
Tarpaulin Cove. NAU: 2883 (N,P,M,C); NAS: Northrop, Aug. 1901 ,
on sheet with var. costulatus (Y).

J CANADENSIS J. Gav. Abundant in moist depressions and around
ponds. UNC: 2994 (P); 3114 (N,P,C); NAU: 2918 (N,P,M); PAS:
3778 (P); NAS: Northrop, Aug. 1901 (Y); CUT: Sanford, Aug. 15.

1917 (N). ,

J. PELOCARPUS Mey. Sandy shores and sandy or peaty pond

borders. NON: 2638 (P), 3203 (N); NAU: 1042 (W), 2379 (P),

^ -^ ▼

244

Rhodora

'■^

(November

2724 (M,C), 2925 (N,P); PAS: 3780 (P); NAS: Northrop, Aug. 1901
{Y) ; FEN: Jordan {o),

J. MiLiTARis Bigel. Sandy and swampy pond margins. NAU:
1038 (P), 2917 (N,P); NAS: Northrop, Aug. 1901 (Y), Williams, July
10, 1911 (N), Pennell 2915 (P,W), 1037 (P,W).

J. ACUMiNATUS Michx. Abundant in wet depressions and on boggy
or peaty borders of small ponds in open hills. NON: 2242 (P.C),
2704 (N), 3163 (N,P,M); UNC: 2995 (N,P); NAU: 1041 (P,W), 2391
(N,P); PAS: 757 (W); NAS: Northrop, Aug. 1901 (Y), Pennell 2909
(W); CUT: 3442 (P); PEN: 489 (N,P,W,M), 1757 (P,W).

J. ARTicuLATUS L. Occasional in boggy depressions. PEN: 490^
(P), 1499 (W).

J. ARTICULATUS L., var. OBTUSATUS Engelm. Peaty, usually
brackish, hollows. NAU: 3866 (N) ; CUT: 3574 (N,P).

J. MARGIN ATUS Rostk. Muddy borders of ponds. UNC : 31 17 (N) ;
NAU: 2390 (N,P,M), 2731 (P); PAS: 3779 (P); NAS: Northrop,
Aug. 1901 (Y), 3562 (N).

LuzuLA CAMPESTRis (L.) DC, var. MULTiFLORA (Ehrh.) Celak.
Sandy bank at east end of Nonamesset. NON: 3369 (N).

L. CAMPESTRIS (L.) DC, var. echinata (Small) Fernald & Wiegand.
Rhodora, xv. 38 (1913). Collected only in sandy beech woods on
Naushon. NAU: 3325 (N,P).

LILIACEAE

UvuLARiA perfoliata L. Reported by Mrs. Northrop for Nasha-
wena; no specimen seen.

Oakesia sessilifolia (L.) Wats. Seen in a boggy, wooded
hollow along the south shore of Nonamesset. NON: 3295 (N.P):
NAS: Northrop (o).

LiLiUM PHiLADELPHicuM L. Reported by Mrs. Northrop from
Nashawena and said by wife of caretaker there to grow abundantly at
certain localities on the island today; no specimens seen.

L. tigrinum Ker. Found persisting around leper cottages on
Penikese. PEN : 494 (P, W) .

Asparagus officinalis L. In sandy soil near areas under cultivation.
NAS: Northrop (o); PEN: 493 (N,P,W).

Maianthemum canadense Desf. Abundant in boggy wooded
hollow along south shore of Nonamesset. NON: 3296 (N); NAS:
Northrop (o).

Medeola virginiana L. Boggy woodland on Nonamesset and
edge of brackish marsh at east end of Naushon. NON: 3337 (P);
NAU: 3332 (N); NAS: Northrop (o).

Smilax rotundifolia L. Rather abundant in thickets and wet,
overgrown hollows. NON: 2296 (N,P); UNC: 3044 (N); NAS:
Northrop (o) ; PEN: 495 (W).

» This Is the material erroniously reported from Penilcese as J. debilis Gray in
Rhodora. xxvl. 223 (1924).

V

/TV

f •»

1930J Fogg, — Flora of the Elizabeth Islands, Massachusetts 245

S. HERBACEA L. Reported by Mrs. Northrop from Nashawena;
no specimen seen.

S. GLAUCA Walt., var. leurophylla Blake. See Rhodora, xx.
78 (1918). Known only on basis of Mrs. Northrop's collection from
Nashawena. NAS: Northrop, " July-Aug. " 1901 (Y).

AMARYLLIDACEAE

Hypoxis hirsuta (L.) Coville. Abundant on open, sandy slopes
and knolls at west end of Cuttyhunk. NAS: Northrop (o); CUT:
Sanford, Aug. 15, 1917 (N); 2310 (P).

IRIDACEAE

Iris versicolor L. Wet depressions and pond borders; abundant.
NON: 2678 (N); UNC: 3031 (P), 3121 (N); NAS: Northrop (o),
2357 (N); PEN: 496 (P,W).

I. PRISMATICA Pursh. Apparently not common; reported and
collected by Mrs. Northrop from Nashawena. NAS: Northrop,
"July-Aug." 1901 (Y).

SiSYRiNCHiUM ANGUSTiFOLiuM Mill. Seen growing only in dry
sand along east shore of Penikese, where fairly plentiful. NAS:
Northrop (o); PEN: 497 (P,W).

S. GRAMixoiDES Bickucll. S. graminvxim Curtis, not Lam. Grassy
slopes and pond borders. NON: 2300 (P), 3360 (N,P); CUT: 3449
(N,P,C); PEN: 3408 (N,P,M), 3415 (P).

S. ATLANTICUM Bickn. Wet hollows, open slopes and bogs. UNC:
3081 (P); CUT: 771 (W), 2318 (N,P,M,C), Hcrxey (N); PEN: 3401

(P).

ORCHIDACEAE

Habenaria bracteata (Willd.) R. Br. Reported by Mrs. North-
rop as "common in meadow'* on Nashawena; no specimen seen.
NAS: Northrop (o).

This is one of the surprises of the Northrop list. It is hard to be-
lieve that the compiler could have confused this species of the damp
upland woods with H. clavcllata, II. hie phari glottis or H. lacera, all of
which she likewise reported and the first two of which are represented
by specimens at the New York Botanical Garden. It is greatly to
be regretted that she did not also collect H. bracteata. As it is,
however, there seems no course but to let this record stand.

H. clavellata (Michx.) Spreng. Open bogs, boggy pond borders
and wet, peaty depressions. NAS: Northrop, Julv 1901 (Y), 3540
(N,P); CUT: Sanford, Aug. 15, 1917 (N), 2527 (P.) '

H. ORBicuLATA Torr. NAU: MacRae, July 25, 1904 (N).

Collected by Lillian J. MacRae on Naushon, July 25, 1904.
Specimen originally in the local herbarium at Woods Hole but since

-^n

246

Rhodora

[November

transferred to the collections of the New England Botanical Club.
This, like H. bradcata, constitutes another remarkable addition to
the flora of southeastern Massachusetts, for this plant has not pre-
viously been reported from nearer than Norfolk County.

H. BLEPHARiGLOTTis (Willd.) Torr. Apparently rare; known only
from Mrs. Northrop 's collection on Nashawena. NAS: Northrop,
Aug. 1901 (Y). , „ ^^^„

H. LACERA (Michx.) R. Br. Swampy and boggy hollows. NAU:
2606 (N); PAS: 1787 (P); NAS: Northrop (o); CUT: 2315 (P), 3576

(N). , ^

PoGONiA OPHIOGLOSSOIDES (L.) Ker. Locally abundant m open
peaty and boggy depressions. PAS: 1789 (P); NAS: Northrop (o),
Williams, July 10, 1911 (N) ; CUT: 524 (W). .

Calopogon pulchellus (Sw.) R. Br. Not infreciuent; usually with
the preceding. PAS : 1 788 (N,P) ; NAS : Northrop (o) ; CUT : 523 (W) .

Arethusa bulbosa L. Mrs. Northrop reports this from Nasha-
wena with the comment "quite common in some places." Unfortu-
nately, no specimen is extant, but there seems to be no good reason for
questioning this record. NAS: Northrop (o).

Spiranthes Beckii Lindl. Rare; seen only on dry hillside west
of French Watering Place. NAU : 2946 (N) .

S. GRACILIS (Bigel.) Beck. Dry, grassy, open slopes; fairly abund-
ant. NON: 2622 (N); NAU: 2601 (N); NAS: Northrop, Aug. 1901

(Y).

S. CERNUA (L.) Richard. Local; wet, peaty hollows. UNC:
3024 (N,P); NAU: Cushman & Morse, Aug. 25, 1906 (B).

LiPARis LoESELii (L.) Richard. Local; Mrs. Northrop 's comment
is "moist hollow in face southern cliffs." Taylor's sheet is from
border of Sheep Pond. NAS : Northrop, Aug. 1 901 (Y) ; CUT : Taylor,

July 15, 1919 (N). ^ . , , . ^

TiPULARiA DISCOLOR (Pursh) Nutt. This rare orchid, hitherto
known on the islands of Massachusetts^ only from an old collection on
Martha's Vineyard, is listed by Mrs. Northrop from Nashawena as
"quite common in 3 or 4 localities, c. & ne. part island." Fortunate-
ly, in this instance a specimen was preserved and is now in the herba-
rium of the New York Botanical Garden. NAS: Northrop, Aug. 1901

(Y).

SALICAGEAE

[Salix amygdaloides Andersons. Appears on Mrs. Northrop 's
Nashawena list. This record probably refers to S. alba or some other
willow and, in the absence of material, can scarcely be admitted.]

S. pentandra L. Introduced on Naushon and Penikese; apparently
thriving on latter island. NAU: 3873 (P) ; CUT: 590 (N,P,W,M,C).

S. alba L. X? This hybrid, probably S. alba X fragihs, is abund-

»See Fogg, Rhodora, xxxii, 114 (1930).

* ^m \

i\

^ ^-

^M" >«

.♦

1930J Fogg, — Flora of the Elizabeth Islands, Massachusetts 247

ant around a small freshwater pond on Penikese. PEN: 591 (N,P,W,
M).

S. discolor Muhl. Reported by Jordan from Penikese; no speci-
men.

Populvs alba L. In boggy woods near farmhouse on Nashawena
and around dwellings on Penikese. NAS: Northrop (o), 3511 (P);
PEN:499(N,P,W).

P. Tacamahacca Mill. P. balsamijera of most American authors.
See Sargent, Jour. Am. Arb. i. Gl (1919). Reported by Mrs. North-
rop for Nashawena; no specimen seen.

P. balsamifera L. p. ddtoides jVIarsh. See Sargent, Jour. Am.
Arb., i. 02 (1919). Small clump near landing on Penikese. PEN:
500 (P,\V).

MTRICACEAE

Myrica Gale L. Only locallv abundant, as around dune ponds on
Nashawena. NAU: Williams, July 10, 1911 (N); NAS: Northrop (o),
1773 (N,P,W,M).

M. CAROLiMENSis Mill.^ A commou shrub on the Elizabeths;
thickets, pond margins, and sheltered hollows in open hills. NON:
2237 (N,P); UNC: 2951 (N); PAS: 1783 (N,P,W); NAS: Northrop (o),
2342 (N); CUT: Saiiford, Aug. 15, 1917 (N); PEN: 498 (N,P,W).

JUGLANDACEAE

JuGLANS ciNEREA L. On Mrs. Northrop 's list for Nashawena; no
specimen seen.

Carya alba (L.) Koch. Occasional in mixed woods. UNC: 2454

(N,P); NAU: 777 (P,W); NAS: Northrop (o).

BETULAGEAE

CoRYLUS CORNUTA Marsh. C. rostrata Ait. Reported by Mrs.
Northrop for Nashawena; no specimen seen.

OsTRYA VIRGIMANA (Mill.) K. Koch. An important constituent of
the native woods, especiallv on Nonamesset and Naushon. NON:
2593 (N) ; NAU: 779 (P,\V) ; NAS: Northrop (o).

Betula lutea Michx. Mrs. Northrop reports two specimens near
Choptauk Lake, Nashawena; no material seen.

B. POPULIFOLIA Marsh. Bordering woods and in protected hollows,
where it often forms dense thickets. NON: 2G35 (N,P,M); UNC:
3043 (N,P); NAS: Northrop (o); PEN: Jordan (o).

B. pendula Roth. Planted and established near eastern end of
Naushon. NAU: 2471 (N,P).

B. pubcsccns Ehrh. B. alba, var. pubesccns Spach. With the last.
NAU: 2472 (N,P,M).

1 This was the original spelling employed hy Miller, Ganl. Diet. ed. 8 (1768) and not
carolinensis, as usually given.

248

Rhodora

[November

1930]

Pogg^_Flora of the Elizabeth Ishmds, Massachusetts 249

FAGACEAE

Fagus grandifolia Ehrh. The most important tree on the islands.
Forms a pure stand over large areas on Naushon and Nonamesset, and
occurs mixed with other trees on these two islands as well as parts of
Uncatena and Nashawena. When growing in the heavier woods this
tree apparently does not form fruit, but an occasional isolated speci-
men in the open will be found laden with nuts. NON: 2592 (N);
NAU: 3147 (N) ; NAS: Northrop (o), 3532 (P,M).

F. GRANDIFOLIA Ehrh., forma pubescens Fernald & Rehder. See
Rehder, Rhodora, ix. Ill (1907). Occasional with the species.
UNC: 3125 (N); NAU: 3137 (P), 3693 (N,P).

QuERCUS ALBA L. Abundant; dry sandy woods. NON: 2657 (P);
UNC: 3127 (N); NAS: Northrop (o), 3531 (P).

Q. RUBRA L. A single tree near landing on Penikese, where possibly
introduced. PEN: 592 (N,P,W).

Q. VELUTiNA Lam. Next to the Beech the most important constit-
uent of the wooded areas. NON: 2656 (N), 3309 (P,M), 3916 (P);
UNC: 3126 (P); NAU: 780 (P,W), 3880 (P); NAS: Northrop (o),
3530 (N,P).

[Q. MARiLANDiCA Mocnch is reported from Nashawena by Mrs.
Northrop but is not represented by a specimen. The Black Jack Oak
is unknown northeast of Long Island and it seems probable that this
record refers to some other species, possibly a sapling of Q. velutina.]

URTICACEAE

Ulmus AMERICANA L. Reported by Mrs. Northrop from Nasha-
wena, no specimen seen.

Urtica urens L. Listed by Mrs. Northrop as growing "about
houses" on Nashawena; no specimen seen.

BoEHMERiA CYLiNDRiCA (L.) Sw. Moist woods and pond borders.
Extremely variable and its relation to the variety Drummondiana not
clearly understood. An extensive series has been collected and is
being held for further study. The following is typical. NAU: 3151
(N,P,M,C); NAS: Northrop (o).

B. CYLINDRICA (L.) Sw., var. Drummondiana Weddell. Var.
scabra Porter. See Fernald, Rhodora, xii. 10 (1910). Borders of
salt marshes; perhaps distinct from var. Drummondiana of the south
Atlantic states. CUT: 2523 (N).

POLYGONACEAE

RuMEX Britannica L. Reported by Mrs. Northrop from Nashaw-
ena; no si>ecimen seen.

R. crispus L. Waste places and pond margins; often appearing
indigenous. NAS: Northrop (o); PEN: 598 (N,P,W,M), 3399 (P).

R. obtuMfolivs L. Occasional; the Nashawena collection was made
from the swampy margin of thick woods. NAS: Northrop (o), 3566
{N); FEN '.Jordan (o).

. 'f%

i

.1

R. VERTiciLLATUS L. Swampy margin of woods at east end of
Nashawena with the last. NAS: Northrop (o), 3529 (N).

This and Block Island are the only stations for the plant in south-
eastern New England.

R. MARITIMUS L., var. fueginus (Phil.) Dus^n. R. pcr^icaroidcs,
in part, of the Manual. See St. John, Rhodora, xvii. 73 (1915).
Brackish situations or nmddy pond borders. NAU: 2372 (X,P);
PEN: 3/oorc 1917 (N), 450 (P,W). .

R. Acetosdla L. Exposed knolls, slopes and sandy clearings. ^Ab:
Northrop (o); CUT: 2262 (N); PEN: 597 (N, P, W).

Polygonum glaucum Nutt. P. mantimuvi of Manual. See her-
nald, Rhodora, xv. 69 (1913). Sand and cobble beaches. NON:
2868 (P); UNC: 3051 (N); NAU: C ashman iSc Morse, Aug. 25, 1906
(B), Williams, July 10, 1911 (N); NAS: Northrop, Aug. 1901 (\).

P AVICULARE L. Sandy patches; not common. NAS: i\orthrop

(o);PEN:594(P,W). . , . „ , iivr

P. PENSYLVANICUM L. Not common; borders of small ponds. UI\<^:

2987 (N). , , T^ •

P. PENSYLVANICUM L., var. LAEViGATUM Femald. Uhodora, XIX.
70 (1917). Moist situations; local. PAS: Svnison, Sept. 8, 1926 (N).

P. Hydropiper L., var. projectum Stanford. Rhodora, xxix. 86
(1927). Seen only from margin of small pond near east end of Nona-
messet. NON: 3202 (N).

P. puNCTATUM Ell. P. acre HBK. See Stanford, Rhodora, xxlx.
77 (1927), where, in a footnote, reason is given for attributing prionty
to P. vundatum over P. acre. Pond borders, margins of swamps and
wet situations generally. NON: 2893 (N,P,M), 2699(P,M,C): UNC:
2986 (N,P), 3018 (P); NAU: 2371 (P), 2460 (P,C), 3150 (N,P,M);
CUT: Sanford, Aug. 15, 1917 (N); PEN: 593 (N,P,W).

P Persicaria L. Pond margins and moist depressions. LAC : .501 /
(N); PAS: Svemon, Sept. 8, 1926 (N); NAS: Northrop (o); PEN: 596

(N,P,W,M).

P. ' hydropiperoides Michx. Apparently not common. PAS:

Sven^on, Sept. 8, 1926 (N). . , ^ o

P OPELOUSANUM Riddell, var. adenocalyx Stanford, bee KHO-
DORA, xxviii. 28 (1926). Pond borders. NON: 3207 (P); PAS: 3747

(N,P). ., , y

P Convolvulus L. Trailing over thicket and low growth surrounding

pond. PEN:595(N,P,W,M,C).

P SCANDENSL. Covering scrub growth m swampy hollows. iNUiM :

3172 (N,P); UNC: 3038 (N,P). „ . • t .,

P DUMETORUM L. This species, with smaller fruit than the pre-
ceding, was found only on Pine Island, Nonamesset, ^ ^^^!;5/^V^^"^^^
a dense tangle over the other vegetation on the beach. NON: 3J1-

(N,P).

250

Rhodora

(November

CHENOPODIACEAE

Chenopodium album L. Beaches and exposed sandy areas. UNC:
2956 (N); NAS: Northrop (o); CUT: Penncll 2993 (P); PEN: 1401
(P,W).

Atriplex arEiVARIa Nutt. Sea beaches; not abundant. UNC:
3106 (N); PAS: 1793 (N,P), 3756 (P,M); NAS: Northrop (o); PEN:
Jordan (o).

A. PATULA L., var. hastata (L.) Gray. One of the commonest
plants of the sea beaches and disturbed sandy areas near the shore.
UNC: 2978 (N,P); PAS: 1791 (N,P); NAS: Northrop (o), 3502 (P);
CUT: Pennell 2996 (P), 2272 (N); PEN: 1400 (P,W), 3386 (M,C).

Salicornia mucronata Bigel. Salt marsh areas; less frequent than
the two following. NAU: Pennell 3140 (W); PAS: Svenson, Sept. 8,
1926 (N).

S. EUROPAEA L. Salt marshes and brackish mud flats. NON: 2892
(N); UNC: 3094 (N,P); PAS: Svcnson, Sept. 8, 1926 (N); 3742 (P);
NAS: Northrop (o); CUT: 3570 (N,P); PEN: Jordan (o).

S. AMBiGUA Michx. Same situations as last. NAU: Pennell 3112
(P), Penncll 3145 (W); PAS: Svcnson, Sept. 8, 1926 (N), 3741 (N,P);
CUT: 3571 (N,P,M,C).

SuAEDA MARiTiMA (L.) Dumort. Reported by Jordan from Peni-
kese; no specimen.

S. LINEARIS (Ell.) Moq. Salt marshes and brackish muds. UNC:
2963 (P,W), 3076 (N); PAS: 3739 (N,P); CUT: 3567 (P,M,C).

Salsola Kali L. Beaches and brackish marshes. NON: 2286
(N,P); NAU: Cushman & Morse, Aug. 25, 1906 (B), 3886 (P); NAS:
Northrop (o), 2364 (N); CUT: 2273 (P); PEN: Jordan (o).

S. Kali L., var. caroliniana (Walt.) Nutt. With the last; less
abundant. UNC: 2961 (P), 3075 (N,P); PAS: 1792 (N,P).

AMARANTHACEAE

Amaranthus retroflexus L. Dry sandy soil near cultivated area.
PEN: 1402 (N,P,W,M).

PHYTOLACCACEAE

Phytolacca americana L. P. decandra L. See Rhodora, xvii.
180 (1915). Apparently rare; seen growing only along edge of boggy
depression at east end of Pasque. NAU: Petmell 3174 (W)- PAS*
3785 (N).

AIZOACEAE

Mollugo verticillata L. Bare, grassy slopes and sandy patches.
NON: 2891 (N); UNC: 3052 (N,P); NAS: Northrop (o), 3470 (P).

CARYOPHYLLACEAE

Spergularia rubra (L.) J. & C. Presl. Dry exposed slopes and
sandy clearings. NAS: Northrop (o), Williams, July 10, 1911 (N)
3465 (P,M); CUT: 3459 (P); PEN: 1410 (P,W).

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 251

S salina J. & C. Presl. Sand and cobble beach along Northwest
Gutter. Rare ; represented on these islands usually by the next species.

NAU: 3132 (N, W). ^ . , _ . , .

S. LEiosPERMA (Kindb.) F. Schmidt. S. rnarmnof Manual m
part. See Fernald & Wiegand, Rhodora, xu. 5/ (19 0)^ ^^00^,^,^
cobbly beaches, mud flats and brackish n^a^shes NON : ^fOiN);
UNC- 2962 (N,P,M); NAU: 2455 (N); PAS: 3738 (N,P,C); CUT
3569 (N,P); PEN: 1409 (P,W).

Spergula arve7isis L. Not common. NAS: ^orthrop (o), PEN

"^"sSna^procumbens L. Upper beaches and dry, gravelly banks
NON: 2692 (N,P); NAU: 2728 (N); NAS: .\orthrop (o), 3464 (P),

PEN* 1408 (W)

Arenaria lateriflora L. Seen growing only >" sandy thicket
along north shore of Naushon near the east end. NAU: 66lt, (,<.m;,
NAS: A'ortArop,"July-Aug." 1901 (Y). ,, ^ ^ . ,„

A. PEPLOiDES L., var. robusta Fernald. See Rhodora x. 109
(1909). Occasional on sandy or cobbly beaches. CUl. ^^74 (,lN,r;,

PEN: 1403 (W). „ . , ^^^.

A. serpyllifolia L. Sandy and gravelly areas; occasional. NUM.
PennM2M0 (W); N.\U: 3424 (N); NAS: A'^rMrop (o)

StcUaria graminca L. Grassland on Penikese. PbN . 1411 ir,v\;,

^1° meia(L.) Cyrill. Dry, sterile soil and grassland on Penikese;
probablv more widespread. PEN: 855 (V,), 339o (N. P . ,

CeraMium vulgatum L. Abundant on open, grassy hillsides and
sandy clearings NON: 3367 (N,P.M), 3280 (P); NAS: Northrop
(o); CUT: 3438 (P); PEN: 1404 (N,P,W).

Agrostemma Githago L. Reported by Mrs. Northrop from Nasha-

wena; no specimen. . , . , j t> -i ^o« PPM-

Lychnis alba Mill. Occasional m grassland on Penikese. i't.IS

^"^^psophila paniculata L. Persistent around leper cottages on Pen-

ikese. PEN: 1406 (P,W). ^

Diayithus barbatus L. Same situation as the preceding. PEiN . 14Uo

^^' Armeria L. Seen only on edge of wet hollow at west end of
Cuttyhunk. CUT: 3445 (N,P).

PORTULACACEAE

Portulaca oleracea L. Sandy ground, usually near cultivated areas.
NAS: Northrop (o); PEN: 1412 (W).

NYMPHAEACEAE

Nymphozanthos variegatus (Engelm.) Fernald. ^:2/7|_^«^« «^
vena, var. vanegaia Fernald. See Fernald, Rhodora, xxi. 183 (1919).

252

Rhodora

[November

Dune ponds on Nashawena and Sheep Pond, Cuttyhunk; possibly
more general. NAS: Northrop (o), 2366 (N,P); CUT: 786 (W), 2531
(N).

Nymphaea odorata Ait. Castalia odorata Woodv. & Wood. See
Rhodora, xviii. 161 (1916). Rather general in smaller ponds as well
as Sheep Pond, Cuttyhunk. UNC: 3004 (P); NAU: 2734 (N); NAS:
Northrop (o), 2367 (N,P); CUT: 2530 (N).

RANUNCULACEAE

Ranunculus Cymbalaria Pursh. Sandy and gravelly shores;
rather frequent. NON : 2302 (N,P) ; NAU : 528 (W,P) ; CUT : Hervcy,
no date, (N); PEN: Northrop (o), 1414 (W).

R. DELPHiNiFOLius ToFF. Small, gFassy-bottomed ponds in open
hills. NAS: Northrop (o), 3522 (P) ; PEN: 451 (P,W), 1759 (N,P,M).

[R. REPTANS L. appears on Mfs. NoFthFop's list. It is hard to say
what this may refeF to in the absence of a specimen. R. reptans, var.
ovalis (R. Flammula, vaF. reptans of the Manual) is unknown fFom
southeasteFn Massachusetts and it seems haFdly justifiable to FepoFt
it on the basis of such an ambiguous recoFd.]

R. hulbosus L. Seen only on a steep gFavel bank at the extFeme
east end of Nonamesset; possibly moFe abundant. NON: 3282 (N).

R. acris L. RatheF common in open gFassland, especially on outeF
islands. NAS: Northrop (o); CUT: 2325 (N); PEN: 1413 (N,P,W).

Thalictrum revolutum DC. RepoFted by Mfs. NoFthFop as
T. purpurascc7is, but had pFobably best be FefeFFed hcFe. NAS:
Northrop (o).

Anemone virginiana L. Collected in dFy, sandy woods on " Cap-
tain's Island," Naushon. NAU: 3907 (P).

A. quinquefolia L. RepoFted by Mfs. NoFthFop foF Nashawena;
no specimen seen.

CoPTis groenlandica (OedeF) FcFuald. C. trifolia of the Manual.
See FeFnald, Rhodora, xxxi. 136 (1929). Included in Mfs. NoFthFop's
list with the comment "in one filled up pond under fcFu Ivs. VeFy
small, no fFuit." NAS: Northrop (o).

This is an interesting addition to the floFa of the islands. Coptis
is unknown fFom BaFnstable County, although repFCsented in both
Plymouth and BFistol Counties. MoFeover, it is FepoFted by Bick-
nell for Nantucket, where, as on Nashawena, it may be a Femnant of a
foFmerly moFe dominant continental flora. There seems to be no
real reason, even in the absence of herbaFium matcFial, to discredit
this recoFd.

BERBERIDACEAE

Berheris Thunbergii DC. Occufs scattcFcd oveF the open hills at
the east end of Uncatena. UNC: 2975 (N).

19301 Fogg, — Flora of the Elizabeth Islands, Massachusetts 253

LAURACEAE

Sassafras officinale Nees & Eberni. N. varufolium (Salisb.)
Ktze. See Blake, Rhodora, xx. 9S (1918). ScattcFed abundantly
thFOUghout all of the wooded areas. NAS: Northrop (o), 3533 (P).

S. officinale Nees & EbcFm. var. albidum (Nutt.) Blake. Rho-
dora, XX. 98 (1918). Occasional with the species. UNC: 3123 (N).

Benzoin aestivale (L.) Nees. RepoFted by Mfs. NoFthFop fFom
Nashawena; no specimen.

PAPAVERACEAE

Glaudum flavum CFantz. Established on beach neaF TaFpaulin
Cove. NAU: 789 (P,W), 1798 (N).

CRUCIFERAE

Bcrteroa incana (L.) DC. Dfv sandv soil neaF landing at east end
of Pasque. PAS: 3798 (N,P,M).

Lepidium virginicum L. Open gFassland and low sandv stretches.
UNC: 3056 (N,P); PAS: 3762 (P); NAS: Northrop (o); CUT: 2259
(N,P); PEN: 1417 (N,P,\V,M,C).

Capsclla Bursa-pastoris (L.) Medic. Occasional in disturbed situ-
ations. NAS: Northrop (o); PEN: 854 (P,W).

Cakile edentula (Bigel.) Hook. Sandy and cobbly sea beaches,
especiallv outeF islands. NAS: Northrop (o); CUT: 2271 (N); 3431
(P); PEN: 1416 (P,\V).

Raphanus Raphanistnnn L. DistuFbed sandv soil neaF cultivated
aFeas. NAS: Northrop (o); CUT: 2268 (N,P,M,C); PEN: 1418 (W).

R. sativus L. Same situations as last. PEN: 1419 (N,P,W).

Brassica juncca (L.) Cosson. On peninsula, east end of Penikese.
PEN: 1415 (P,W).

B. nigra L. DFy, sandy slope. NAU: 790 (\V); FE^^: Jordan (o).

Sisymbrium officinale (L.) Scop., vaF. leiocarpum DC. BaFFen,
sandy gFound; waste places. PEN: (N,P,W,M).

S. altissimum L. Same situations as the pFeceding. PEN: 1420
(W).

Rorippa Nasturtiiun-aquaticum (L.) Schinz & Ihel. Radicida ^as'
turtium-aqiiaticum BFitt. & Rendle. Seen gFowing along a moist rill
dFaining into Sheep Pond, Cuttvhunk. NAS: Northrop (o); CUT:
1054 (P,W).

DROSERACEAE

Drosera rotundifolia L. Boggv situations and boFdcFs of ponds.
NON: 2652 (N), 3362 (P); PAS: 1786 (N,P); NAS: Northrop (o),
3472 (P).

D. intermedia Hayne. D. longifolia of Manual. AppaFently not
common; boFdcFS of dune ponds on Nashawena. NAS: 1778 (N,P),
Northrop (o), Williams, July 10, 1911 (N).

254

Rhodora

[November

SAXIFRAGACEAE

RiBES HiRTELLUM Michx., var. CALCicoLA Femald. See Fernald,
Rhodora, xiii. 76 (1911). Seemingly rare. NON: 2844 (P).

HAMAMELIDACEAE

Hamamelis virginiana L. Dry to moist woodland and thickets.
Some of this material approaches the variety parvifolia Nutt. NON:
2591 (N,P); NAU: 3152 (N,P); NAS: Northrop (o), 3512 (P).

ROSACEAE

Spiraea tomentosa L. Pond borders and moist hollows; abundant.
NON: 2564 (N); PAS: 1784 (N,P); NAS: Northrop (o), 2345 (N,P).

Pyrus arbutifolia (L.) L. f. Thickets and wet depressions; of
frequent occurrence. NON: 2885 (N,P); UNC: 3046 (N,P); NAS:

1774 (N,P), 3495 (P). ^^ . ^ ^ ^.

P. ARBUTIFOLIA (L.) L. f., var. ATROPURPUREA (Britton) Kobmson.
Same situations as the last but less common. NON: 3175 (N,P);
UNC: 3124 (N,P); NAS: Northrop, Aug. 1901 (Y).

Amelanchier oblongifolia (T. & G.) Roem. Dense thickets and
sheltered hollows; abundant. NON: 2630 (P), 2883 (N,P), 3205 (P),
3202 (P,C), 3308 (P,M,C); PEN: 1421 (P,W).

Crataegus pruinosa (Wendl.) C. Koch. Single tree on dry hill-
side near Tarpaulin Cove. NAU: 1801 (N,P).

Crataegus spp. This genus is to be met with scattered through-
out the dry, sandy beech woods, mostly on Naushon. Several
collections have been made, but it has not been found possible to
match this material with existing specimens or to reconcile it with
available descriptions. Each individual plant appears to possess its
own characteristics which differ from those of its neighbor, and,
rather than give to these specimens names which assume a definite-
ness that they do not own, the problem is here left in abeyance.

Fragaria virginiana Duchesne. Dry open slopes and hollows.
NON: 3287 (N); NAS: Northrop (o); PEN: 1422 (W).

F. VIRGINIANA Duchesne, var. terrae-novae (Rydb.) Fernald &
Wiegand. Rhodora, xiii. 106 (1911). Seen only in a sandy meadow
near Cuttyhunk Pond. CUT: 3581 (N,P).

F. vesca L. Reported by Jordan from Penikese: no specimen.
Potentilla norvegica L., var. hirsuta (Michx.) Lehm. P. mon-
speliensis L. See Bibliot. Bot. 16, Heft Ixxi. 404 (1908). Dry open
ground. PEN: 1424 (N,P,W,M). ., ...o .r . /x

P. argentea L. Dry sandy and gravelly soil. NAS: Northrop {o);
PEN: 1423 (N,P,W,M).

P. recta L. Sandy clearing in woods near West Gutter. NAU:

3923 (N,P).

19301 Fogg, — Flora of the Elizabeth Islands, Massachusetts 255

P. PACIFICA Howell. P. Ansrriiia of Manual, in part. See Fernald,
Rhodora, xi. 1 (1909). Shingle beach along south shore of Nonames-
set. NON: 2874 (N,P,M); NAS: Northrop, as "P. Anscrina" (o).

P. PUMILA Poir. I)rv knolls and exposed sandy and grassy liill-
sides. NON: 2677 (N,P), 3288 (P,M,C); PEN: 1425 (N,P,\V).

P. CANADENSIS L., var. simplex (Michx.) T. & G. The Elizabeth
Islands material seen all has narrow, oblanceolate leaves and short,
more or less appressed, pubescence. Dry slopes and sandy clearings.
NON: 3355 (N,P,M); CUT: 2536 (N).

Geum canadense Jacq. Apparently rare; boggy woods at east end
of Nashawena. NAS: Northrop (o), 3544 (P).

RuBUS occidentalis L. Edge of woods at west end of Nonamesset.
NON: 2676 (N).

R. pergratus Blanchard. Thickets and margins of boggy hollows.
NAS: 3487 (P); PEN: 1428 (N,P,W).

R. laciniatiis Willd. Well established on peninsula of Penikese.
PEN: 1429 (N,P,W).

R. ANDREwsiANUS Blauchard. This is the commonest Blackberry
on the Islands, occurring in wet overgrown hollows and in thickets on
open slopes. NON: 2577 (P). 2626 (N,P); UNC: 3034 (P); NAU:
Pennell 3135 (W), 2496 (P), 2497 (P).

R. HISPIDUS L. Abundant; apparently as much at home on the
dry exposed hillsides as in the boggy depressions. NAS: Northrop (o),
3539 (N); CUT: 2312 (P), 2321 (N,P,M).

R. FLAGELLARis Willd. R. procumhciis of many authors. R. vil-
/o5w^ of the manual. See Bailey, Gen tes Herb. i. 234 (1925). Trailing
over dry open sandy and gravellv ground. NAS: Northrop (o); PEN:
1430 (N,P,W).

Rosa rugosa Thunb. Established and thriving, usually along beach-
es, on several of the islands. NAU: Taylor 3077 (P); CUT: 3439 (N);
PEN: 1427 (P,W).

R. rubiginosa L. Reported by Mrs. Northrop from Nashawena;
no specimen seen.

R. palustris Marsh. /?. Carolina of Manual and most American
authors. See Fernald, Rhodora, xx. 91. (1918). Wet margins of
ponds. NAS: Northrop, July. 1901 (Y), 2354 (N,W); PEN: 1426 (W).

R. CAROLINA L. R. humilis jVIarsh See Fernald, 1. c. Wet hollows
and pond margins. NON: 2631 (N,P); NAS: Northrop (o), 3477 (P);
CUT: 3461 (N).

R. NANELLA Rydb. N. Am. Fl. xxii (pt. 6), 497 (1918). Material
collected from sandy thicket on Uncatena has dwarf stature and very
narrow leaflets and corresponds in all essentials to Rydberg's descrip-
tion of this species. UNC: 3112 (N).

R. VIRGINIANA Mill. Moist hollows and swampy borders of ponds.
NON: Pennell 2843 (P), 3173 (N,P), 3346 (P,M); PAS: 3746 (N,P).

Prunus serotina Ehrh. Sheltered hollows, thickets and sandy
woods. NON: 2671 (N,P); UNC: 3108 (P); NAU: 2501 (N,P); NAS:
Northrop (o), 3519 (P); PEN: 1431 (W).

256

Rhodora

[November

P MARITIMA Wang. Apparently not very common; forms a dense
thicket at east end of Nonamesset. NON: 3206 (N), 3303 (P,M);
NAS: Northrop (o).

LEGUMINOSAE

Gleditsia triacanthos L. Sandy woods near dwellings at east end
of Naushon; originally introduced. NAU: 3925 (?)• . ^ ,^

Genista tindoria L. Well established and spreadmg m fields at east
end of Naushon and on Uncatena. UNC: 2976 (N); NAU: 796 (N,

P,W).

'Cyiisus scoparius (L.) Link. Abundant and spreading on parts of
Naushon (see p. 155); local elsewhere. NAU: ^^^**f,,f ^,\^' J^^? ^2,
1890 (N), Moore, 1903 (N); PAS: Rollick, Aug. 1898 (Y); NAS: NoHh-

rop (o). J TV/r »

Ulex europaeus L. A single large clump along road near Mary s
Lake. NAU: 797 (W).

Trifolium arvense L. Dry open ground and sandy areas. ^^^ ^
2669 (P); UNC: 3012 (N); NAS: NoHhrop (o); CUT: 2257 (N); PEN:

1434 (P,W).

T. praterise L. Grassland; occasional. NAS: Northrop (o); PEN:

1435 (P W).

T repens L. Open fields and grassy levels. NAS: Northrop (o);

CUT: 2261 (N,P); PEN: 3402 (N,P). ...^ .. .. r ^

T. hyhridum L. Open grassland; occasional. NAS: Northrop {o)\

PEN: 1436 (P,W). ^ . . .

T. agrarium L. Disturbed sandy soil near areas under cultivation.

PEN* 1433 (N P W).

Coronilla variaL. Abundant on grassy hillside toward west end

of Pasque. PAS: 795 (P,W).

Robinia Pseudo-Acacia L. At several localities along the north
shore of Naushon, where doubtless planted but now spreading. NAU :
3874 (N,P); NAS: Northrop (o).

Desmodium rigidum (Ell.) DC. Sandy woods and hills; not com-
mon. UNC: 2977 (N,P); NAU: 2600 (N,P). ^

D. OBTUSUM (Muhl.) DC. Sandy woods and clearings. NAU:
3910 (P,M,C), 3928 (N,P). i • ^

Lespedeza procumbens Michx. Not abundant; seen only in sandy
woods on " Captain's Island." NAU: 3909 (N,P,M).

L. viOLACEA (L.) Pers. Reported from Nashawena by Mrs. North-
rop; no specimens seen. , ^.t i i- ^

L. HiRTA (L.) Hornem. On Mrs. Northrop's Nashawena list; no

specimen seen. j u i

L. CAPITATA Michx. Apparently not abundant; open, sandy bank

on Uncatena, west end. UNC: 3129 (N).

Vicia tetrasperma (L.) Moench. Disturbed sandy area near landing.

PEN: 1438 (W). ,^, ^^^,

V. Cracca L. Dry, open ground. CUT: Taylor 3076 (P); PEN:

1437 (N,P,W,M,C).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 257

V. mllosa Roth. Drv sandy strip near Cuttyhunk Pond. CUT:
3579 (N,P).

Lathyrus maritimus (L.) Bigel. Sea beaches; commoner on outer
islands. NAS: Northrop (o); CUT: 2263 (N,P); PEN: 1432 (N,P,W).

L. PALUSTRis L. Known only from single collection on Nashawena.
This material approaches var. pilosus. NAS: Northrop, "July- Aug."
1901 (Y).

Apios tuberosa Moench. Occasional in wet hollows and around
ponds. UNC: 3089 (N); NAS: Northrop (o); CUT: 2328 (N,P).

Amphicarpa monoica (L.) Ell. Sandy woods near Hadley Harbor.
NAU: 3917 (N,P); NAS: Northrop (o).

LINACEAE

LiNUM STRIATUM Walt. Dry hillsides and peaty pond margins.
CUT: 2330 (P), 2538 (N,P).

L. VIRGINIANUM L. Drv sandy woods. NAU: Williams d' Collins,
July 10, 1911 (N),2381 (N,P).

OXALIDACEAE

OxALis STRICTA L. Open sandy hillsides and sandy clearings.
UNC: 3096 (P); NAU: Penncll 3142 (W), 2498 (P), 3682 (P); NAS:
Northrop (o); PEN: 1439 (N,P,W).

GERANIACEAE

Geranium maculatum L. Seen only on an exposed gravel bank
at upper margin of beach along north shore of Nonamesset. NON:
3364 (N,P).

G. CAROLINIANUM L. Bumt-over ground near hospital building on

Penikese. PEN: 1440 (W).

SIMARUBACEAE

Ailanthus glandulosa Desf. Naturalized and spreading at several
localities on Naushon and Nashawena. NAS: Northrop (o), 3481 (P).

POLTGALACEAE

PoLYGALA POLYGAMA Walt. Open sandv and grassy hillsides.
NAU: 2477 (N); NAS: Northrop, " July-Aug." 1901 (Y).

P. SANGUINEA L. Low boggy ground. Not common. NAS:
Northrop (o), 3563 (N).

P. CRUCIATA L. Peaty and boggy pond borders; apparently not
widespread. CUT: Sanford, Aug. 15, 1917 (X), 2514 (N,P).

EUPHORBIAGEAE

AcALYPHA VIRGINICA L. Kuowu oulv from a sheet of Dr. Pennell's
collecting on Naushon. NAU: Pennell 3140 (W,P).

258

Rhodora

[November

A. DiGYNEiA Raf. See Weatherby, Rhodora, xxix. 193 (1927).
Rare. Boggy depression in open hills near east end of Uncatena.
UNC: 3131 (N).

A. GRACILENS Gray. Dry hillsides and moist depressions; local.

UNC: 2974 (P), 3087 (N).

Euphorbia polygonifolia L. Sea beaches and low sandy stretches.
NON: 2870 (N); UNC: 3008 (N); PAS: 3755 (P,M); NAS: Northrop,
"July-Aug." 1901 (Y); PEN: Jordan (o).

E. maculata L. Sandv beaches and dry exposed slopes. NON:
2869 (N); NAU: 2499 (P)- PEN: Jordan (o).

CALLITRICHACEAE

Callitrichk palustris L. Apparently rare and local. NAS:
Northrop (o), 3521 (P).

C. heterophylla Pursh. Small ponds in hills; more frequent than
last. NON: 2235 (N); PEN: 456 (W), 1095 (P,W), 1758 (N,P).

(To he continued)

THE FLORA OF THE ELIZABETH ISLANDS,

^LVSSACHUSETTS

John M. Fogg, Jr.

{Continued from page 258)
ANACARDIACEAE

Rhus typhixa L. Occasional in sheltered hollows on hillsides.
NAS: Northrop (o), 3479 (N); PEN: 1441 (P,W), 3409 (N,P).

R. GLABRA (L.) Gray. No material seen ; reported by Mrs. Northrop
from Nashawena. .

R. GLABRA (L.) Gray, var. borealis Britton. See Britton s Man.
Fl. No. St. Can. 601 (1901). Forming a small grove along the North-
west Gutter at the west end of Uncatena. UNC: 2453 (P,^I).

R COPALLINA L. Open slopes and moist depressions. UNC: 3039
(N) ; PAS: 3793 (N,P) ; NAS: Northrop, Aug. 13, 1901 (Y).

R. Vernix L. Swampy woods and thickets. NAU: 2485 (N,P);
PAS: 3794 (N); NAS: Northrop (o).

R. Toxicodendron L. Thickets, copses and overgrown hollows.
Extremely variable as to habit, leaf-shape and fruit. NON: 3345
(N,P,M); UNC: 3040 (N,P); PAS: 1785 (N,P); NAS: Northrop (o),
2352 (N); CUT: 2314 (N); PEN: Jordan (o).

AQUIFOLIACEAE

Ilex opaca Ait. Occasional in sandy wooded areas. NAU: 2488
(P,M,C); NAS: Northrop (o), 3535 (N). _

I. oPACA Ait., forma subintegra Weatherby. Rhodora, xxm. 118
(1921). Sandy woods with the last; rare. NAU: 2484 (N,P,W,M,C).

264

Rhodora

[December

I. VERTiciLLATA (L.) Gray. Moist overgrown depressions and wet
borders of ponds. NON: 3174 (N,P,M) ; NAS: 3490 (P).

I. VERTiciLLATA (L.) Gray, var. fastigiata (Bickn.) Fernald.
Rhodora, xxiii. 274 (1921). In similar situations as last; occasional.
NON: 2884 (N, P, M); NAS; Northrop, "July-Aug." 1901 (Y), 3494

(N).

I. LAEVIGATA (Pursh) Gray. Reported by Mrs. Northrop for Nash-
awena; no specimen seen.

I. GLABRA (L.) Gray. The Inkberry is evidently very rare on the
Elizabeth Islands. Although sought for repeatedly, it is known only
from a single specimen collected on Nashawena by Mrs. Northrop.
NAS: Northrop, Aug. 1901 (Y).

ACERACEAE

Acer rubrum L. Sandy and boggy woods and thickets; var. tridens
was nowhere seen. NON: 2655 (N,P) ; NAS: Northrop (o), 3534 (N).

A. platanoides L. Planted in small grove near hospital building on
Penikese. PEN: 1443 (W).

A. Pscudo-Platanus L. A few trees with the preceding. PEN: 3410
(P,M).

SAPINDACEAE

Aesculus Hippocastanum L. Included in Mrs. Northrop's list with
the comment, " Single good sized tree in midst of native trees about
Choptauk Lake. Introduced." No specimen seen.

BALSAMINACEAE

Impatiens biflora Walt. Wet borders of ponds.
NAU: 2735 (N,P); NAS: Northrop (o).

NON: 2887 (N);

VITACEAE

Parthenocissus quinquefolia (L.) Planch. Psedcra quinqucfoUa
Greene. Occasional in thickets. NON: 2294 (N,P); NAS: Northrop
(o); PEN: 856 (W).

P. tricuspidata (Sieb. & Zucc.) Planch. Trailing over stone wall
near hospital building on Penikese. PEN: 3411 (N,P).

ViTis Labrusca L. Thickets and moist hollows. UNC : 3028 (N) ;
NAS: Northrop, July, 1901 (Y), 3493 (P).

V. AESTIVALIS Michx. Trailing over low trees, edge of sandy woods.
NAU: 2490 (P).

MALVACEAE

Malva rotundifolia L. Dry sandy soil, waste places. NAU: Pcnncll
3115 (W); NAS: Northrop (o); PEN: 1442 (P,W).

Althaea officinalis L. Mrs. Northrop, in listing this species for
Nashawena, says, "2 or 3 plants on diiTerent parts of shore." No
specimen seen.

IRREGULAR PAGINATION

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 265

Hibiscus Moscheutos L. borders of salt marslics. XOX: 2(173
(P); UNC: 2985 (N,P); NAU: 2378 (N).

HYPERICACEAE

Ifi/pcricuin pirforatiitn L. Abundant evervwliere on dr\- hillsides.
NON: 2()79 (P); UN'C: 29()S (X); XAS: Northrop (o), 3552 *(P); CUT:
2332 (X); PEX: 1445 (N,PAV.M,C).

H. BOREALE (Hritton) Bickn. Pond borders and wet hollows;
abundant. XOX: 2()42 (P,W), 2701 (X,P); UXC: 2983 (X); XAU:
2720 (X,P); PAS: 3738 (P); XAS: 3523 (P).

H. MUTILUM L. Same habitats as last; less fre(|ucnt. XOX: 25(i9
(X); XAU: Prnnrll 3125 (W), 23S9 (P); XAS: Northrop (o); 1*KX:
Jordan (o).

H. CANADENSE L. IVatv pond margins and boggx hollows. X'OX':
2045 (X); UXC: 3023 (X>), 3085 (X,P); XAS: Northrop (o); CUT:
Sauford, Aug. 15, 1917 (N).

H. GENTIANOIDES (L.) BSP. I hv sandv hillsides and clearings.
NON: 2694 (P); UNC: 308(; (X); XAS: Northrop (o).

II. VIKCJINUIM L. Sandx and peatv pond borders, often j)artlv
immersed. XOX: 2040 (P/, 3171 (X,P,C); UXC: 3022 (X); XAF:
2927 (X,P); XAS: Northrop (o), 2351 (P,M); CUT: Sanftml, .\ng. 15,

1917 (N).

ELATINACEAE

El.\ti.\e mim.ma (Xutt.) F. & M. See Fernald, Uhoduka, xix. 10
(1917). Sandv bottoms and borders of ponds. XOX: 2700 (X.l*);
NAU: Pcnnrlf :i2{)2 (W), 2727 (P), 2943 (Xj; PAS: KMil (P). W(\\ (F);
NAS: Northrop (o); CUT: 2533 (Xj.

CISTACEAE

IlELiANinKMLM CANADENSE (L.) Miclix. Open grassland and <lr\
sandv clearings; abundant. XOX: 2()25 (X,P), 2578 (P,M), 2903
(X);*XAU: 2474 (P,M), 2598 (X,P); XAS: Northrop (o), 355S (P).

H. PROPINQUUM Hickn. See Hull. Torrey Hot. Club, xl. 015 (1913).
Seen only on sandy bank bordering West End Pond. X\VU : 2915 (X",P).

H. nu KNELLii Fernahl. //. nut jus of Manual. See Hiiodoka, xxi.
30 (1919). Drv, open, grassy slopes. XAU: 2493 (P). 2945 (X);
P.\S: Svrnson, Sept. 8, 1920 (X); XAS: Northrop, Aug. nK)l (V).

H. Di'MOSUM (Hickn.) Fernald. See Hull. Torrey Hot. Club, xl. ()13
(1913), and Rhodora, xix. 00 (1917). (Grassland and sandy clearings.
NAU: 2494 (P,\V), 2944 (X); P.\S: Svcnson, Sept. 8, 1920 (X); XAS:
Northrop, Aug. HM)! (Y), 3557 (P,M).

HuDSONiA TOMENTOSA X^itt. Open sandv areas. XAU: U'illiant.i,
Julv 10, 1911 (X); PAS: 800 (X.P,M); XAS: Northrop, "July-Aug."
1901 (V), Williams,, My 10, 1911 (X). /V//W/291() (W), 2305 (P,M).

Lechea villosa Ell. Aj)parentlv not eonmion; sandy woods on
Xaushon. XAU: 2402 (P).

266

Rhodora

[December

L MARITIMA Leggett. Abundant on open downs and sandy stretch-
es. NON: 2584 (N), 2624 (P); NAU: 2473 (P), 2481 (P), 2o97 (N);

PAS: 3770 (P). _ ^, ,

L. Leggettii Britton & Rollick. Reported by Mrs. Northrop

from Nashawena; no specimen seen.

VIOLACEAE

Viola papilionacea Piirsh. Not common; seen only in sandy
beech woods on Naushon. NAU: 3314 (P), 3326 (N,P).

V FIMBRIATULA Sm. Abundant on open downs and dry, grassy
slopes. NON: 2581 (N), 3289 (P); NAU: Pmndl 3129 (W), 3126 (P);
CUT: Sanford, Aug. 15, 1917 (N), 2457 (P,M); PEN: 1444 (N,P,W).

[V. SAGITTATA Ait. Reported by Jordan for Penikese and by Mrs.
Northrop for Nashawena but probably refers to the preceding.]

V. lanceolata L. Open bogs and peaty borders of ponds. NON:
2568 (P,M), 2894 (N), 3307 (P); UNC: 3088 (N); NAU: Pemiell 3122
(W), 2511 (N); NAS: Northrop (o); CUT: Sanford, Aug. 15, 1917 (N).

V. PALLENS (Banks) Brainerd. Peaty and boggy hollow at west end
of Nashawena. NAS: 3499 (N,P).

[V. BLANDA Willd. Appears on Mrs. Northrop's Nashawena list.
In absence of specimens it seems most likely that this refers to the
preceding.]

LTTHRACEAE

Decodon verticillatus (L.) Ell. Abundant in swamps, borders
of ponds and low wet thickets. NON: 2563 (N,P), 3170 (N,P);
NAU: 2387 (N); PAS: 3795 (P); NAS: Northrop (o),2338 (N,P,M,C);
CUT: Saifford, Aug. 15, 1917 (N).

MELASTOMACEAE

RnEXiA viRGiNiCA L. Wet, peaty depressions and borders of small
ponds. NON: 2639 (N,P,M); NAS: Northrop, "July-Aug." HK)1
(Y), Northrop, Sept. 1901 (Y), 3548 (P); CUT: 2522 (P).

ONAGRAGEAE

LuDViGiA PALUSTRis (L.) Ell. Floating or growing on moist mud;
small pond holes. NON: 2240 (C), 2571 (P,M), 3164 (N,P); NAU:
2468 (N,P); PAS: 3784 (P); NAS: Northrop (o), 3526 (N,P); CUT:
3447 (P);PP:N:457 (P,W).

Epilobium densum Raf. Reported by Mrs. Northrop from Nash-
awena as E. lincare; no specimen seen.

E. PALUSTRE L., var. monticola Haussk. Collected in an open
sphagnum bog at the west end of Cuttyhunk on July 28, 1927. CUT:

2307 (N).

E. COLORATUM Muhl. Seen only on the shingle at French Watering
Place; perhaps more general. NAU: 2718 (N).

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 267

E. GLANDULOSUM Lehm., var. adenocaulon (Haussk.) Fernald.
E. adcnocaulmi Haussk. See Rhodora, xx. 34 (1918). Reported from
Nashawena by Mrs. Northrop; no specimen seen.

Oenothera muricata L. Dry sandy soil, east end of Pascpie;
probably more common. PAS: 3761 (P),

O. BIENNIS L. Dry sterile ground, occasionally in grassland. NAS:
Northrop (o); PEN: 1446 (N,P,W).

0. grandiflora Ait. On Penikese, where probably introduced. PEN:
1447 (W).

HALORAGIDACEAE

Myriophyllum scabratum Michx. Rather frequent; small ponds
and wet pond holes. NON: 2241 (G,N,P), 3363 (P); UNC: 2996
(N,P,M,C), 3647 (P); NAU: 801 (N); NAS: C. E. Falcon, Sept. 1875
(G), Northrop (o); PEN: 453 (N,P,W), 3398 (P).

M. humile (Raf.) Morong. Abundant and variable; forma capiU
laccum and forma nutans have both been met with but appear to be
nothing more than mere ecological variations. NON: 2705 (N,P,
M,C); NAU: 2503 (xN,P,M); PAS: 845 (N,P); NAS: Northrop (o),
3520 (N,P); CUT: 3448 (P).

M. TENELLUM Bigel. Muddv borders and bottoms of several small
ponds. NAU: 2401 (N), 2753 \N,P); PAS: 1068 (W); NAS: Northrop
(o); CUT: Hcrvey, no date (N).

Proserpinaca PALUSTRIS L. Ponds and pond borders, often sub-
merged. NON: 2570 (P), 2643 (N,P), 33S2 (P,M); UNC: 3026 (N);
PAS: 554 (P,W); NAS: Northrop, JuW, 1901 (Y); CUT: 803 (W),
Sanford, Aug. 15, 1917 (N).

UMBELLIFERAE

Sanicula canadensis L. Drv sandy woods; not common. NAU:
2459 (P,M), 3927 (N,P,C).

Hydrocotyle umbellata L. Shallow water of ponds and muddy
or grassy pond margins. UNC: 3001 (N); NAU: Williams et al., July
10, 1911 (N), 1805 (N), 2502 (N,P), 2729 (M), 2947 (N,P,C); NAS:
Northrop, "Julv-Aug.'* 1901 (Y), Pennell 2923 (W); CUT: Sanford,
Aug. 15, 1917 (N).

H. Canbyi Coult. & Rose. NAU: 1069 (G), 2948 (N,P).

In the Gray Herbarium there is a specimen of H. Canbyi labelled
H. ujnbcllata, var. amhigua which was collected at "Woods Holl,
Sept. 10, 1876, on shores of Nobska Pond." This sheet is further
labelled "Herb. W. G. Farlow," and, though no collector's name
appears, may have been collected by Farlow himself. It constituted
the only known record for this plant north of southern New Jersey
until 1925 when, on July 14, Dr. H. K. Svenson and the writer found
it growing around the shores of French Watering Place on Naushon.

268

Rhodora

[December

On Aug. 31, 1927, the writer again visited this locality and found the
plant, this time along another portion of the shore of the same pond.
H. Canhyi is thus definitely established as a member of the flora of
New England.

H. VERTiciLLATA Thuub. NON: 2895 (G,N,P); UNC: 2981 (G,
N,P,W), 3000 (P).

The reported occurrence of this species in Massachusetts has
apparently been based upon a sheet collected by Thomas Morong at
Woods Hole on Aug. 10, 1877. This sheet, which is in the Gray
Herbarium, bears the determination "H. interrupta Muhl." and is
perfectly typical H. vcrticillata as understood today. Also, there is a
sheet of this species in the herbarium of the Boston Society of Natural
History collected at New Bedford and further labelled merely " Oakes
Coll. '* Southwest of Massachusetts the plant has been known only
from Block Island before "jumping" to southern New Jersey (Cape

May County).

On August 30, 1927, while exploring the west end of Nonamesset,
the writer came upon a small pond in the open hills which was filled
with a species of Hydrocotyle. This, upon examination, proved to be
H. vcrticillata. Three days later, September 2, this same species was
found growing in profusion in two small grassy ponds along the north
shore of Uncatena toward the east end. Thus, a second member of
this genus is restored to our list for this region after a lapse of half a
century.

Ptilimnium capillaceum (Michx.) Raf. Borders of brackish ponds
and salt marshes. NON: 2595 (N), 2664 (P); UNC: 3019 (N,P);
NAU: 2370 (P,M,C), 2715 (N,P); NAS: Northrop (o), 3501 (P).

CicuTA MACULATA L. On Mrs. Northrop's list for Nashawena;
no specimen seen.

SiUM SUAVE Walt. S. cicvtacfolium Schrank. See Blake, Rhodora,
xvii. 131 (1915). Swampy hollows in woods and borders of ponds.
UNC: 2988 (P); NAU: 2386 (N,P); NAS: Northrop (o), 2350 (N,P);
CUT: 3443 (P).

LiGUSTicuM scoTHicuM L. Sandy and gravelly beaches. NON:
3210 (N); UNC: 2979 (N,P); PAS: 557 (P,W); PEN: 1449 (P,W).
► CoELOPLEURUM LUCIDUM (L.) Femald. C. actacifolivm (Michx.)
C. & R. See Fernald, Rhodora, xxi. 144 (1919). Sand and cobble
beaches; not very common. CUT: 3435 (N,P); PEN: Jordan (o).

Pastinaca sativa L. Reported by Mrs. Northrop from Nashawena;
no specimen seen.

Heracleum lanatum Michx. Seen growing only on gravelly bank
along East Gutter. NON: 3310 (N); NAS: Northrop (o).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 269

Daucus Carota L. Dry grassland; apparently not common. NAS:
Northrop (o); CUT: 2326 (N,P); PEN: 1448 (P,W).

CORNACEAE

CoRNUS FLORIDA L. Fairly abundant in boggy woods in hollows
along south shore of Nonamesset; perhaps elsewhere. Seen in full
"flower '^ on May 28, 1928. NON: 3302 (N,P,M,C) ; NAS: Northrop (o).

Nyssa sylvatica Marsh. A common tree both in the wet wooded
hollows and in the moist portions of the beech-oak forests. NON:
2636 (P), 2667 (W,M), 2886 (N,P); NAU: Simons, July 23, 1901 (W^,
2932 (N); NAS: Northrop (o), Williams, July 10, 1911 (N), 3517 (P).

ERICACEAE

Clethra alnifolia L. Thickets and pond borders; abundant.
NON: 2293 (P,C), 2627 (N,P), 2882 (N,M); UNC: 3649 (N,P);
NAS: Northrop (o), 1771 (P); CUT: 2309 (P).

Chimaphila MACULATA (L.) Pursh. Reported by Mrs. Northrop
from Nashawena; no specimen seen.

Pyrola rotundifolia L., var. americana (Sweet) Femald. P.
americana Sweet. See Rhodora, xxii. 122 (1920). Mrs. Northrop,
who reports this from Nashawena, says, "Few plants in one station."
No specimen seen.

MoNOTROPA uniflora L. Drv saudy beech and oak woods. NAU:
Williams, July 10, 1911 (N), 806\P,W); NAS: Northrop (o), 3537 (P).

M. hypopithys L. Seen only in sandy woods at east end of Nash-
awena; perhaps more common. NAS: 3538 (N).

M. hypopithys L., var. rubra (Torr.) Farwell. See Amer. Midi.
Nat., X. 39 (1926). The crimson variety was collected in dense beech
woods near Witches' Glen on Naushon; seen nowhere else. NAU:
3687 (N).

Rhododendron viscosum (L.) Torr. Thickets and pond borders.
NON: 2629 (N); NAS: Northrop (o), 1769 (N,P,M); CUT: 2317 (N).

R. VISCOSUM (L.) Torr., var. glaucum (Michx.) Gray. Known
only from a single specimen collected by Williams on Nashawena;
probably more abundant. NAS: Williams, July 10, 1911 (N).

Kalmia angustifolia L. Apparently rare; reported by Mrs.
Northrop from Nashawena, but no specimen seen.

Leucothoe racemosa (L.) Grav. Occasional around sandy mar-
gins of ponds. NAU: 2482 (P,M),^2938 (N,P).

Lyonia ligustrina (L.) DC. Dense thickets and wet borders of
ponds. NON: 2628 (N,P); UNC: 3650 (N,P,M); NAS: Northrop
(o), 1768 (N,P).

Chamaedaphne calyculata (L.) Moench. Seen growing only
around borders of dune ponds on Nashawena. NAS: Northrop, Aug.
1901 (Y), 1770 (N,P,W).

Epigaea repens L. Reported by Mrs. Northrop from Nashawena
with the remark, "On bank near Mirror Lake." No specimens seen.

270

Rhodora

[December

Gaultheria procumbens L. Apparently not common. Reported
from Nashawena and foimd there growing sparingly in boggy woods
at east end. NAS: Northrop (o), 3545 (N).

Gaylussacia dumosa (Andr.) T. & G., var. Bigeloviana Fernald.
See Rhodora, xiii. 95 (1911). Also Rhodora, xxiii. 248 (1921).
Decidedly not common; so far known only from Nashawena. NAS:
Northrop, July, 1901 (Y), Northrop, " July-Aug." 1901 (Y), Taylor,
July 6, 1920 (P).

G. FRONDOSA (L.) T. & G. Occasional in thickets and about ponds.
NON: 2236 (N,P); NAS: Northrop, Aug. 1901 (Y).

G. BACCATA (Wang.) C. Koch. This is the commonest Huckleberry
on the islands; it is everywhere in the open areas, often forming an
impenetrable growth in the hollows or under the lee of protecting
hills. NON: 2576 (N,P), 2651 (P); NAU: 2504 (P,C), 2941 (N,P,
M), 3689 (P); NAS: Northrop (o), 3559 (P).

Vaccinium corymbosum L. Abundant around ponds, in moist
depressions and in boggy woods. NON: 2637 (P); NAS: Northrop
(o), 3516 (P).

V. ATROCOCCUM (Gray) Heller. Same situations as last but appar-
ently less common; flowers and fruit earlier. NAS: 1776 (P).

V. MACROCARPON Ait. Open bogs and peaty hollows. NAS: North-
rop (o); CUT: 2313 (P), 2519 (N,P,M).

PLUMBAGINACEAE

LiMONiUM Nashii Small, var. trichogonum Blake. L. carolinianum
of the Manual. See Rhodora, xxv. 58 (1923). The following num-
bers, out of a large series being held for study, may be cited. UNC :
3122 (P); NAU: 2456 (P).

In his recent revision of the Sea Lavenders Blake characterizes the
New England material as having the calyx densely pubescent, at
least at the base. Nevertheless, specimens collected from the Eliza-
beth Islands show the calyx nearly smooth or with a few scattered
hairs and, according to Blake's treatment, would come closer to L,
carolinianum, which name he restricts to the southern material with
glabrous calyx. It is obvious that the last word has not yet been
said concerning this genus as it is represented in the eastern United
States.

PRIMULACEAE

Samolus floribundus HBK. Sandy and peaty pond margins; not
common. NON: 2873 (N) ; NAU: 1078 (W), 2722 (N).

Lysimachia quadrifolia L. Sandy woods; infrequent. NAU:
3327 (P); NAS: Northrop (o).

L. TERRESTRis (L.) BSP. Open bogs and pond borders. NAS:
Northrop (o), 2344 (N); CUT: 2316 (P).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 271

Triextalis borealis Raf. T. amcricana (Pers.) Pursh. See
Blanchard, Rhodora, xi. 236 (1909). Boggv woods. NON: 3297
(N,P); NAU: 3328 (N,P); NAS: Northrop (o). ^

Anagallis arvensis L. Cobble beaches and sandv waste places.
UNC: 2957 (N); NAS: Northrop (o); CUT: 3434 (P); PEN: 1450
(N,P,W).

OLEACEAE

Fraxinus AMERICANA L. Certainly not common; seen only on
edge of swampy hollow along north shore of Nashawena. NAS:
Northrop (o), 3480 (N).

F. PEXSYLVANICA Marsh. Reported by Mrs. Northrop from Nasha-
wena; no specimen seen.

Ligmtrum vulgare L. Escaped from cultivation on Penikese. PEN:
1451 (W).

GENTIANACEAE

Bartonia virginica (L.) BSP. Open bogs and peatv hollows or
pond borders. PAS: 3782 (P); NAS: Northrop, Aug. 1901 (Y), 3508
(N); CUT: 2515 (N,P).

B. PANICULATA (Michx.) Robinson. A single specimen of this
species was collected in the woods on Naushon by Professor Fernald
and the writer on Aug. 10, 1927, but the material was subsecpiently
mislaid. The plant is surely uncommon on the Elizabeth Islands.

Menyanthes trifoliata L., var. minor Michx. See Rhodora,
xxxi. 198 (1929). Seen growing only in a small lx)ggv area bordering
Sheep Pond. NAS: Northrop (o); CUT: Ilrrvey, no date (N), 810
(P,W).

Nymphoides lacunosum (Vent.) Fernald. Collected in a small
pond on Nonamesset. NON: 2707 (P).

APOCTNACEAE

Apocynum androsaemifolium L. Exposed sandy bank bordering
beach on Uncatena; possibly more general. UNC: 3639 (N,P).

ASCLEPIADACEAE

Asclepias tuberosa L. Sandv hillsides and clearings; rare and
local. NAU: Simom, July 16, 1901 (W), 2602 (N); PAS: 589 (\V).

A. INCARNATA L., var. PULCHRA (Ehrh.) Pers. The commonest
Milkweed on the islands. Swamps and wet pond margins everywhere.
NON: 2674 (N,P); UNC: 3059 (P); NAU: 2736 (N); NAS: Northrop
(o), 3478 (P); PEN: Jordan (o).

A. SYRIACA L. Occasional in drv sandy ground. NAS: Northrop
(o); PEN: 1452 (W).

A. PHYTOLACCOIDES PuTsh. Reported by Mrs. Northrop as A. exal-
tata with the comment "Not uncommon." No specimen seen.

A. VERTiciLLATA L. Known only from Mrs. Northrop's collection
from Nashawena. NAS: Northrop, Aug. 1901 (Y).

272

Rhodora

[December

CONVOLVULACEAE

Convolvulus sepium L. Occasional in thickets and overgrown
depressions. NON: 2890 (P); T^ASMhrop (o .

C. SEPIUM L., var. pubescens (Gray) Femald. On sea beaches,
apparently not abundant. PEN: 1454 (N,P,W). ppvr. 1453

C. arvensu L. Cobble beach of isthmus on Penikese. PEN. 1453

rimrTiTA Gronovii Willd. Swamps, boggy clearings, wet thickets,
etc nIu:p3 3127 (W), 2495 (P), 2605 (N P,M , 29 4 (P.C ,
2933 (P) 3139 (N,P,M), 3890 (P,C); NAS: Northrop (o), Wtlhams,
July 10, 1911 (N), 2353 (P).

This is apparently the only species of Dodder on the islands. A
large and variable series has been collected and has been studied in
the light of T. G. Yuncker's recent monograph of the genus (19^1J,
but it has not been found possible to recognize more than this single
polymorphic species. ^^.j^gj^^cEAE

Verbena hastata L. Collected on f ^^^^^y "PP%i'j^%r H'^'u'
tena and in swampy clearing on Naushon. UNC: 2980 (N), WAU.

24«^ (P)- LABIATAE

Teucrium CANADEN8E L., var. LiTTORALE (Bickn.) Fcrnald. Abund-
ant near beaches, borders of brackish areas wet thickets etc^^>^^^
Cnshman, July 27, 1906 (B) 2586 (N ; UNC: 2960 (P) SM^ (N,P^ ,
NAU- 1797 (P), 2500 (P,M); NAS: Northrop (o);PEN: 1460 (IN-P."}.

Trichostema dichotomum L. Frequent on open downs and sandy
knoUs NON: 2623 (N); UNC: 2973 (P), 3120 (N); NAS: Northrop

^° Scutellaria lateriflora L. Listed as " common " on Nashawena

by Mrs. Northrop; no specimen seen. lUnmial See

S EPILOBIIFOLIA Hamilton. S. gdericulata of the Manual, bee
Femald Rhodora. xxiii. 85 (1921) Abundant - -* ^oll^w^^
around ponds. NON: C,«Aman, July 27, 1906 (B 2298 (N,P.C)

SSAu?iMS\N),8\5kp);PE^

Marrvbium vulgare L. Sandy soil ^^ste places. NAU Moore 1904
(N), 814 (W); NAS: Northrop (o), Wtlhams, July 10, 1911 (!S),

Xi'Sa^L. Dry, sterile soil. UNC: 3054 (N); NAS: iVor^A-

rop(o); PEN: 1458 (N,P,W). ^ „.

Prurlella vulgaris L. Apparently not common; dry ground. NAS.

'^R v7Li:LTv2:x.ANCEOLATA (Bartou) FemaR RhoooR'. xv
179 (1913). The indigenous variety was met with on dry, sanay
slopes near center of Uncatena. UNC : 3032 (P).

1930] Fogg, — Flora of the Elizabeth Islands, Massachusetts 273

Leonurus Cardiaca L. Fairly abundant on Penikese; not collected
elsewhere. NAS: Northrop (o); PEN: 1455 (W).

Pycnanthemum flexuosum (Walt.) BSP. Known only from spec-
imen in Woods Hole herbarium collected on Naushon. NAU: Stjuons,
July 23, 1901 (WO.

P. MUTICUM (Michx.) Pers. Reported and collected by Mrs.
Northrop from Nashawena. NAS: Northrop, Aug. 1901 (Y).

Lycopus virginicus L. Rare and local on the islands; seen growing
only on the sandy beach bordering West End Pond. NAU: Cushman
iSc Morse, Aug. 25, 1906 (B), 2920 (N).

L. UNIFLORUS Michx. The most abimdant species of the genus
here; pond margins, wet hollows, etc. NON: 2565 (N,P,W), 2663
(P); UNC: 2969 (N); NAU: Williams, July 10, 1911 (N), 2487 (P);
NAS: 2346 (N); CUT: 2525 (N); PEN: 1457 (W).^

L. AMERICANUS Muhl. Frequent; moist situations. UNC: 2970
(N); NAS: Northrop (o), 3749 (N,P); CUT: 2524 (N,P), PEN: 1456
(N,P,M,W,C).

Mentha spicata L. Occasional in wet hollows. NAS: }iorthrop (o),
3491 (P,M).

M. Cardiaca Gerarde. Reported by Mrs. Northrop from Nasha-
wena; no specimen seen.

M. crispa L. Collected in a swampy hollow at the east end of Cutty-
hunk. CUT: 2284 (N).

SOLANACEAE

Solanum Dulcamara L. Seen only trailing over a dense thicket
bordering pond on Uncatena. UNC: 3042 (N).

S. NIGRUM L. Very abundant as a littoral plant, often fringing the
sandy or cobbly beaches with a solid growth. NON: Cmhman, July
27, 1906 (B), 2872 (N,P); NAS: Northrop (o); CUT: 3446 (P); PEN:
1462 (N,P,W).

S. rostratum Dunal. Sandy waste areas; infrequent. NAU: Taylor,
July 24, 1921 (P); PEN: Hollick (o).

S. triflorum Nutt. This western species was foimd growing on
beach near Tarpaulin Cove; possibly introduced in the days when
the Cove served as a refuge for sailing vessels. NAU: 1806 (N).

Datura Stramonium L. Sandv and cobbly beaches; of casual occur-
rence. UNC: 2958 (N); NAU: Cushman & Morse, Aug. 25, 1906 (B);
PEN: 1461 (W).

D, Tatula L. On Mrs. Northrop's Nashawena list; may refer to
the preceding; no specimen seen.

SCROPHULARIACEAE

Verhascum Thapsus L. Open, sandy ground and waste places.
NAU: 2720 (N,P); NAS: Northrop (o); PEN: 1465 (N,P,W).

Linaria vulgaris Hill. Decidedlv not common on the islands; seen
only on Penikese. PEN: 857 (W').

274

Rhodora

[December

L. CANADENSIS (L.) Dumont. Grassy slopes and open sandy
stretches. NON: 3373 (P); NAS: Northrop (o), 3483 (N); CUT:
2331 (N); PEN: 1464 (P,W), 3403 (P).

LiMOSELLA SUBULATA Ives. L. aquatica, var. tcnuifolia of Manual.
See Rhodora, xx. 160 (1918). Sandy and muddy margins of ponds
on Pasque and Nashawena. PAS: 3745 (N,P); NAS: Northrop,
"July-Aug." (Y), 2356 (N).

Ilysanthes dubia (L.) Barnhart. Reported by Mrs. Northrop
from Nashawena as /. gratioloides; no specimen seen.

I. inaequalis (Walt.) Pennell. /. anagallidca (Michx.) Robins.
See Pennell, Torreya, xix. 149 (1919). Sandy and peaty pond borders.
NON: 2702 (N); NAS: Williams, July 10, 1911 (N); PEN: 454 (N,

P,W).

Gratiola aurea Muhl. Peaty margins of small ponds. NON:
2641 (P), 3194 (N,P), 3383 (N,P); NAU: Taylor 2347 (P), 1079 (P,W).

Digitalis purpurea L. A few plants persistent around leper cottages
on Penikese. PEN: 1463 (W).

Veronica serpyllifolia L. Reported from Nashawena by Mrs.
Northrop; no specimen seen.

V. peregrina L. Occasional in sandy soil. The Naushon material
is very sparingly pubescent, approaching var. xalapensis (HBK.) Pen-
nell. NAU: 3330 (P) ; NAS: Northrop (o) ; CUT: Hcrvey, no date (N).

V. arvcnsis L. Seen only in a sandy field near center of Nonamesset.
NON: 3306 (N,P).

Agalinis purpurea (L.) Pennell. Gcrardia purpurea L. See
Bull. Torrey Bot. Club, xl. 126 (1913). Listed by Mrs. Northrop for
Nashawena; no specimen seen.

A. MARITIMA Raf. Gerardia viaritima Raf. Seen only on border of
brackish area on Uncatena ; probably more general. UNC : 3005 (N,P).

Melampyrum lineare Lam. Mrs. Northrop reports this for Nash-
awena; no specimen seen.

Pedicularis canadensis L. Dry, sandy beech woods near Hadley
Harbor. NAU: 3313 (N,P).

LENTIBULARIACEAE

Utricularia geminiscapa Benj. U. clandestina Nutt. In a small
pond on Nashawena; probably commoner. NAS: 3504 (P).

U. GiBBA L. Known only from sheet in Woods Hole herbarium
collected on Naushon. NAU: Simom, July 23, 1901 (W).

U. BiFLORA Lam. Reported from Nashawena by Mrs. Northrop;
no specimen seen.

U. CORNUTA Michx. Reported and collected by Mrs. Northrop.
NAS: Northrop, Aug. 1901 (Y).

OROBANCHACEAE

Epifagus virginiana (L.) Bart. Fairly abundant in beech woods
on Naushon; should be found on other islands as well. NAU: Wit-
Hams, July 10, 1911 (N). 2937 (P), 3138 (N).

19301 Fogg,— Flora of the Elizabeth Islands, Massachusetts 275

BIGNONIACEAE

Catalpa hignonioides Walt. Well naturalized in certain areas on
Nashawena. NAS: 3546 (P).

PLANTAGINACEAE

Plantago major L. Cobbly beaches, grassy slopes, etc. NON:
2697 (P), 2900 (N); NAS: Northrop (o); PEN: 1467 (N,P.W), 3407

(N,P).

P. MAJOR L., var. intermedia (Gilibert) Dene. Material so labelled
was collected by A. H. Moore on Penikese in 1904. Here also may be
referred tentatively specimens collected by Hollick on Naushon and
by Mrs. Northrop (labelled P. halophila Bickn.) on Nashawena. This
is done with the realization that the maritime phase of P. major is in
need of very careful studv in order that its status may be satisfactorily
determined. NAU: Hollick, Aug. 11, 1898 (Y); NAS: Northrop,
Aug. 1901 (Y); PEN: Moore 1916 (G, N).

P. JUNCOIDES Lam., var. decipiens (Barneoud) Fernald. P. viari-
tima of American authors in part, not L. See Fernald, Rhodora,
xxvii. 100 (1925). Collected on exposed gravelly bank along north
shore of Naushon west of Kettle Cove. NAU: 3876 (N,P).

P. OLIGANTHOS R. & S. P. maritima of American authors in part.
See Fernald, 1. c. 93-104. Salt marshes and brackish muds. PAS:
3740 (N,P,M,C); CUT: 3568 (P). x... xt ,

P. lanceolata L. Grassland; not common. NAS: Northrop (o);

PEN: 1466 (P,W), 3394 (N,P). . .^ . ^. .. , i

P. lanceolata L., var. sphaerostachya Mert. & Koch. See I'ernald,
Rhodora, xxiv. 203 (1922). Sandy and cobbly beach on Uncatena.

UNC: 3055 (N,P). . ^

P. lanceolata L., var. sphaerostachya Mert. & Koch., forma eriophora

(Hoffmans. & Link) Beck von Man. See Fernald, 1. c. Sandy soil at

east end of Cuttyhunk. CUT: 2269 (N). _

P. aristata Michx. Occasional in sandy fields. UNC: 36o2 (N,P);

NAU: Simons, July 23, 1901 (W); CUT: Northrop (o).

P. VIRGINICA L. Mrs. Northrop reports this from Nashawena,

saying, " Common on downs, about 2 in. high." No specimen seen.

RUBIACEAE

Galium Aparine L. Listed by Mrs. Northrop for Nashawena

no specimen seen. , xiz-ixr o^no /t>\

G PiLOSUM Ait. Dry sandy woods and clearings. NON : 2ob2 (F;

NAU: Pennell 3150 (W), 2930 (N), 3686 (P) ; NAS: Northrop (o).
G. ciRCAEZANS Michx. Collected by Pennell on Naushon. NAU

Pennell 3110 (W).

G. TRiFiDUM L. Borders of brackish areas; not common. NAU
2513 (N), 3684 (P,M); NAS: Northrop (o), 3541 (P).

Erroneously reported from Penikese; see Rhodora, xxvi. 228
(1924). The Penikese material proves to be G. Claytoni,

276

Rhodora

[December

G. Claytoni Michx. Wet thickets, pond margins and grassy
slopes; very abundant. NON: 2572 (C), 2646 (N), 2666 (P), 2880 (P),
2906 (P); UNC: 3027 (N); NAU: 2374 (P), 2716 (N,P,M); PAS:
3748 (P); CUT: 2516 (N,P); PEN: 1468 (N,P,W).

G. TRIFLORUM Michx. Collected only at edge of beach near east
end of Naushon; possibly more general. NAU: 2457 (N,P).

MiTCHELLA REPENS L. Reported by Mrs. Northrop from Nasha-
wena; no specimen seen.

Cephalanthus OCCIDENT alis L. Not common; collected in boggy
pasture on Nashawena. NAS: Northrop (o), 3564 (P).

CAPRIFOLIAGEAE

Lonicera japonica Thunb. Occasional as an escape. UNC : 3037
(N); PEN: 1469 (N,P,W).

Triosteum perfoliatum L. Seen only on dry open hillside on
Pasque. PAS: 567 (N,P).

Viburnum acerifolium L. Reported by Mrs. Northrop for Nash-
awena; no specimen seen.

V. PUBESCENS (Ait.) Pursh. V. venosum Britton. See Blake,
Rhodora, xx. 11 (1918). This is the only Viburnum seen growing
upon the islands. It is of frequent occurrence in moist hollows and
in thickets bordering ponds. UNC: 3010 (N,P); NAU: 2458 (P,M).
3320 (P,C); NAS: 3489 (N), 3518 (P).

Viburnum dentatum L. is reported by Mrs. Northrop from Nash-
awena. It is possible that this refers to the preceding, but in the ab-
sence of specimens this record is permitted to stand.

Sambucus CANADENSIS L. Occasional; thickets and moist depres-
sions. NAS: Northrop (o), 3488 (N); PEN: 1470 (N,P,W).

CUCURBITACEAE

Cucurbita maxima Duchesne. Sandy soil near cultivated area on
Penikese. PEN: 1471 (W).

CAMPANULACEAE

Specularia perfoliata (L.) a. DC. Reported from Nashawena
by Mrs. Northrop; no specimen seen.

LOBELIACEAE

Lobelia cardinalis L. Found only in a swampy clearing at the
east end of Naushon. NAU: 2466 (P,M).

COMPOSITAE

Eupatorium verticillatum Lam. Mrs. Northrop reports E. mac-
ulatum from Nashawena, but it is probable that this refers to the plant
treated by Wiegand as E. verticillatum (See Rhodora, xxii. 57 (1920)),

1930] Fogg,— Flora of the Elizabeth Islands, Massachusetts 277

a coastal plain species the general absence of which from the Eliza-
beths is indeed surprising. NAS: Northrop (o).

E. HYSSOPIFOLIUM L. Rather frequent on open grassy and sandy
slopes NON: 2590 (P); UNC: 2953 (N); NAU: Oakcsr (Herb.
Lowell) (B); PAS: 1790 (P,W).

E. verbenaefolium Michx. Reported by Mrs. Northrop from

Nashawena; no specimen seen

E. perfoliatum L. Wet hollows; uncommon. UNC. 29bb (IN;,

NAS: Northrop (o). ^ , i i •

Chrysopsis falcata (Pursh) Ell. Open downs and sandy clearings.
NON: 2583 (P); NAU: 2596 (N,P); PAS: 3788 (N); NAS: i\orthrop

Solidago bicolor L. Dry, sandy woods; not abundant. NAU:

3161 (N), 3905 (P). ^ , , 1.T X' u

fS ULIGINOSA Nutt. is reported from Nashawena by Mrs. Northrop;
this doubtless refers to some other species, but since, m the absence o

material, it is impossible to say which f,'^^'/^^ V''"" K^ON '^2 H fpi •

S. sempervirens L. Sandy and cobbly beaches ^^N. 3211 (P ,

UNC: 3110 (N); PAS: 3744 (P); NAS: Northrop (o); CUT: 22/0 (N);

PEN' 1492 (N P W)

S. JUNCEA Ait. ' Known only from bare gravelly slopes on Penikese.

PFN- 1489 (P W)

S. suAVEOLENS Schoepf. S. odora Ait. See Standley, Rhodora,
xxi. 69 (1919). Collected only in dryish peaty l-^'^^f ''"^/"^titty-
hunk; perhaps more abundant. NAS: Northrop (o); CUl : 2o20 (N.f,

S. Eluottii T. & G. Dry thickets on Uncatena. UNC: 3105 (P),

3111 (N,P,M). , , „ ., ppxT.

S. RUGOSA Mill. Dry, boulder-covered slopes on Penikese. PtJN .

^''s^ iuGO^MiU.. var. aspera (Ait.) Fernald. S. aspera.\it. See
Rhodora, xvii. 7 (1915). Open grassy downs and pond borders.
NON:3186 (N.P.M); UNC:3093 (P); CUT: 2521 (N?)

S. NEMORALIS Ait. Dry hillsides. UNC: 2971 (N,P). PAb. 3797
(P); NAS: Northrop (o); PEN: 1490 (W)

S. CANADENSIS L. Collected only on dry southwest slopes of Pen-

'T a.™;T Sected only in thickets on dry hillsides of Un-

"f t" nZ"!:!!' ?uS!^' Everywhere on open downs and borders of
Gaylvssacia thickets. NON: 2561 (P) : 3 68 (N P); UNC : 3084 (N.
PM); NAU: Cuskman, Aug 25, 1906 (B), 2o05 (M,C), 2719 (N,P),
2929 (N.P); NAS: Northrop (o); CUT: Sanford, .Aug. 15, 1917 (N);

TciiSFoK.) Salisb Same ^tuatiT as the pre^di^
somewhat less abundant. NON : 3179 (N.P.M.C) ; UNC : 2952 (N, P) ,
PAS: 3750 (P).

278

Rhodora

[December

S. MINOR (Michx.) Fernald. This southern species, ranging from
Mississippi and F orida to Virginia and reported by Bicknell from
Nantucket, was collected by Professor Fernald and the writer on open
grassy hillsides near the central part of Naushon. NAU- 2506 fN P)

(N) rNAu73?60''(N P)' ^'''*''°""' '" ^'^ "P*" ^°°ds. ' NON : 3193

XT^;,''^™'^®.^'*- Grassland and sandy thickets. NON- 3190 ("?■) •
NAU: 3142 (N); NAS: A^ortArop (o). v^in. ^lau (r;,

A. UNDULATUS L. Dry woods; common. NON- 3IS3 I'M P"! -^is?
(P,C); UNC: 3080 (N,P,M); NAU: 3158 (N,P;M); PEN 14^^^^

A. ERicoiDES L Reported by Hollick from Pasque;' no specimen
seen nor has this Aster, so abundant on the adjacent mainland, been
found growmg on the Elizabeth Islands, despite diligent search for it
010; ^^^LTiFLORus Ait. Fairly abundant on grassy slopes NON-

?J98 S w)'"" ^""'^^'^ ^^^' '''' ^^'^^^ ^^^-^ '''' ™^ p^n;

A. puMosus L. With the last. NON: 3178 (P) 3189 CN P W^

A DUMosus L., var. coridifolius (Michx.) T. & G. Clearing in
dry beech woods near Mary's Lake. NAU: 3145 (N,P)

A. DUMOSUS X ? This plant, probably representing A. dumosus
Xmmmeus, was collected m sandy woods on Naushon. NAU: 3143

xt;^;/'^^^^^® ^^"^- Abundant in sandy woods and on open downs

A LATERiFLORUs (L.) Brittou. Apparently not common; dry,
shady beech woods on Naushon. NAU: 3146 (N).

A NOVi-BELGii L. Abundant in dry, open ground, especially along
borders ot thickets; extremely variable NON- 3181 (W r\ "^is?

ffifATJKfiisl&V'''^'' '^'"'^ 309r(S;pf 3\o?^P?;' pTs^

A. UMBELLATUS Mill. Reported by Mrs. Northrop from Nasha-
wena; no specimen seen.

o«no .xn"^^"/^"^^ ^- ^^y' open hillsides; not frequent. UNC-
3092 (N); PAS: 3790 (P);NAS:A^or<Arop(o).

tm^f^''V!l!°^''^ 1^'^'*-) ^^P- Open sandy slopes. UNC: 3643
(f); JNAb: Northrop (o).

E. CANADENSIS L. Dbturbed, sandy soil; rare. PEN; 1485 (W)
h PusiLLUs Nutt. See Robinson, Rhodora, xv. 205 (1913) Onen
sandy slopes. NON: 2902 (N,P) ; PAS: 3768 (P) ; PEN- 1486 (P W) '

(M;; PAS: 3789 (P); N.\S: Northrop (o).

S. LiNiFOLius (L.) BSP. Reported from Pasque by Hollick- no
specimen seen. '

Pluchea camphorata (L.) DC. Margins of brackish ponds and

19301 Fogg— Flora of the Elizabeth Islands, Massachusetts 279

borders of salt marshes. NON : 2289 (N). 2662 (P) ; UNC : 3036 (N) ;
NAU: Cushman & Morse, Aug. 25, 1906 (B) 1799 (N ,P,^^ ).

Antennaria plantaginifolia (L.) Richards ^<f™^%,'\"°"^.,'V^{'.

exposed hillsides. NON: 2670 (N), 3280 (N,P); UNC: 3W0 (N.P);

NAU: 3312 (P); N.\S: A'orf/.rop(o). vnv- -^-JSl fPV

A neglecta Greene. Same situations as last. .NU.N . JZ»i l.r;,

NAS: 3506 (P), 3554 (N); CUT: 3458 (N,P). ^, , , ,

Anaphaus MABGARIT..CEA (L.) B. & H. Grasslaml and open
ground everywhere. NON: 2560 (P); UNC: 3035 (N); PAb: 3/96
(P); NAS: A^or<Arop(o); CUT: 3454 (N,P).

Gnaphalium obtus.folium L. G. poyceph»'i,^ ^I;f ''• «P^":
sandv and grassy slopes and clearmgs. LNC: 29o4 (N ; 30 ' (P),
NAU- 2730 (N); PAS^: 3700 (P); NAS: Northrop (o); CUT: Sanford,
Aug 15, 1917 (N); PEN: 1482 (N,P,W,M,C).

G. ULIGINOSUM L. Upper beaches and low san.ly ground, occasion--
ally bordering salt marshes. UNX: 3006 (N); NAU: Cu^hmanA
mLTmS-25, 1906 (B), 2388 (N,P); NAS: Northrop (o); PEN:

G PURPUREUM L. Dry, sandy soil ; rare and local. NAU : 834 (W) ;
N\S- 3507 (N); CUT: /^crwy, no date (N).

IVA ORABIA Bartlett. Reported by Jordan from Pen.kese; no spec-

'Tmbrosia ARTEM.S.IFOL.A L. Not comiiion exccpt on P;";k;««
where abundant on grassy slopes of peninsula. NAS: .\orlhrop (o),

^TAN-Sf ^ANADENSE Mill. Reported by Mrs. Northrop from

Nashawena; no specimen seen. vnv. -iiT? (^Ml

X ECHINATUM Murr. Occasional on beaches. M)?s. .nn (is),

3914 (P); PAS: 3759 (P); PEN: 458 W

Rudbcckia hirta h. Open grassland. UNC: 3641 (P), NAS. Aort/1
rop (o); CUT: 2329 (N); PEN: 1487 (P,^^). „•„*;„„

HeUanthus annuus L. Sandy ground near area under cultivation

on Penikese. PEN: 1483 (P.W). , ^ „„u

Coreopsis rosea Nutt. Apparently rare : seen only on sandy beach
bordering West End Pond. N.\U: 2916 (N,P,M,C).

C lanceolata L. Persistent around leper cottages on Penikese,
where planted. PEN: 1480 (P,VV).

BiDENS CONNATA Muhl. Border of small pond in hills at north end
of Penikese. PEN: 1476 (N,P,\V,M,C). r„o„or^

B. CONNATA Muhl., var. petiolata (Nutt.) .^•■"•Y*^";,^*.^"?.^'',ps'
X 197 (1908) Wet borders of ponds; occasional. UiNU. JUio (r),
3115 (N,P); NAU: 3133 (N,P,W,M,C).

Achillea Millefolium L. Common everj"«-here on open g^as^y

hillsides and in sandy fields and ^le^^nKf^V^^i?^ ^P\T^^ 232ifN^-
2959 (N); NAS: Northrop (o), 3555 (P); CUT: 2260 (P,M), 2322 (i\),

PEN: 1472 (N,P,W). . , , , rrxjr. 90=^=; fNV

Anthemis Cotula L. Disturbed sandy ^^ff- UNC 295o (N),
NAS: Northrop (o); CUT: 3580 (P); PEN: 8o8 (N,P,\\).

280

Rhodora

[December

Chrysanthemum Leucanthemum L., var. pinnatifidum Lecoq. & La-
mo tte. Certainly not common; occasional on open grasslands. NAS:
Northrop (o) ; PEN : 1 477 (N,P, W) .

Tanacctum vulgare L. Sandy soil near habitations. PAS : 3767 (N) :
PEN: 1097 (W).

Artemisia Stelleriana Bess. Occasional on sandy and cobbly beaches.
NON: Cushnan, July 27, 1906 (B); CUT: 2275 (N,P,M).

Erechtites hieracifolia (L.) Raf. Sandy clearings; moist pond
margins and depressions. NON: 2696 (N); NAU: 2928 (N); NAS:
Northrop (o); PEN: 1481 (W).

E. HIERACIFOLIA (L.) Raf., var. prealta (Raf.) Fernald. See
Rhodora, xix. 27 (1917). Moist hollow. NON: 2661 (N,P).

E. MEGALOCARPA Femald. See Rhodora, xix. 24 (1917). Moist
depressions; local. UNC : 2965 (P,C) ; 2984 (N,P,W,M).

Arctium viinm Bernh. Reported by Mrs. Northrop for Nashawena;
no specimen seen.

Cirsium lanceolatum (L.) Hill. Sandy and gravelly banks and hill-
sides. NON: 2693 (N,P), Cushman, July 28, 1906 (B); NAU: 2717
(N,P); NAS: Northrop (o); CUT: 827 (P,W); PEN: 1479 (P,W).

C. HORRiDULUM Michx. C. spiuosissiinum of the Manual. See
Robinson, Rhodora, xiii. 239 (1911). Occasional in open exposed
situations. NON: 2299 (N); NAU: Miss Weir, June 22, 1890 (N);
NAS: Northrop (o).

C. DISCOLOR (Muhl.) Spreng. Occasional in sandy clearings and
borders of thickets. UNC: 3104 (N); PAS: 3752 (P).

C. PUMiLUM (Nutt.) Spreng. Not common; met with only in
grassland on Naushon. NAU: 828 (P,W) ; NAS: Northrop (o).

C. arvense (L.) Scop. Abundant on Penikese, where it forms a
fringe on the upper beach to the south; local elsewhere. NAS: North-
rop (o); PEN: 1478 (N,P,W).

Centaurea arcnaria Bieb. Reported from Nashawena by Hollick
as the first record for North America;^ no specimen seen.

Cichorium Intybu^ L. Listed by Mrs. Northrop from Nashawena;
no specimen seen.

Krigia virginica (L.) Willd. Sandy fields. NON: 3305 (N,P);
NAU: 3879 (N,P).

K. BiFLORA (Walt.) Blake. K. amplexicaulis Nutt. See Rhodora,
xvii. 135 (1915). On Mrs. Northrop's Nashawena list; no specimen
seen.

Leontodon autumnalis L. Occasional in grassland or on dry banks.
UNC: 3060 (N); NAU: 2392 (N); NAS: Northrop (o).

L. autumnalis L., var. pratensis Koch. In tall grass on slopes at
west end of Penikese. PEN: 1484 (W).

Taraxacum officinale Weber. Not common; collected only near
landing on Penikese. NAS: Northrop (o) ; PEN: 1496 (N,P,W).

Sonchus arvensis L. Occasional on beaches and sandy areas. NAS:
Northrop (o); PEN: 1493 (N,P,W,M,C).

> Hollick, A. Cont. Geol. Dept. Columbia Univ xi no 72. (1901 r

1930] Fogg —Flora of the Elizabeth Islands, Massachusetts 281

S. okraceus L. Seen mostly ^o^}^i\^^th^_^ti^rol upper sea

^^..(LTmi^i^tSi"^^^^^^^

^^S?; Z\^L L. Reported by Mrs. Northrop from Nash-
awena; no specimen ^J^"- ,^ (53; j ) Gray, forma angus-

L. CANADENSIS L., Var. I^TEGRIFOUA VOlgtM p^',. „.-j :„ A^y

TATA Wiegand. See Rhodora, xx.k 10 a920). Collected m dry,
QonHv soil at east end of Pasque. rAb: eSu^ l,r;.

L HiRSUTVMuhl. Seen growing only in sandy woods on Naushon;
nrLblv more common. NAU: 2464 (N); NAS: Norlhrop {o\
Tre!an™ Serpentaria Pursli. Mrs. Northrop reports th« fron.

^?r;:uorTTtSrss.)Xnald. Sandy woods; apparently rare.

^mERAacf PXN,ct;L..TUM L. Dry, sandy woo^is; not common.

^RmS?an™ Willd. Reported from Nashawena by Mrs. Northrop;

""H^'slTurMichx. On Mrs. Northrop's list for Nashawena; no

'•'h "grc^ovh L. This is the commonest ."ember of the genus on
these islands; dry woods, sandy clearings etc. ^pN: 2634 (P), 318o
^m MiiT- wq VNV P\S- 572 (W), 3775 (N); NAS: 3o3b (P).
^%''cF.>^.^^l^^ -^V^^'^ not a.."ndant collected o^^^^^
in dry, sandy soil at east end of Pasque. P.\S. 3//4 (N), >.V&.
Norlhrop (o).
University of Pennsylvania.

MEMBRANES IN THE PLANT CELL

I. MORPHOLOGICAL MEMBRANES AT PROTOPLASMIC

SURFACES

Janet (^. Plowe

With 11 Text-figures
Keceived for publication, September 8, 1930

INTRODUCTION

The qiirstioii of t\\v existJMirc and function of plasma membranes
is of concern to the pliysiologist both fi'om a pi'actical and from a theo-
retical pomt of view. It is (d practical interest becanse, in attempting-
to analyze the action of chendcals on the livin<»: orjz'anism, it is essential
to know wh(d.her substances in th(^ extei'iial medinni iiecessarily comr
into direct contact with all internal (M)nstituents (d" the c(dl which they
affect, or whether action on an external reji'ulatcu'y layer alone may b«'
responsible for chaii^'es in other parts of the c(dl which the chendcals
never reach. It is of theoretical interest because, if it can be shown that
the existence and function of plasma mend)ranes is essential to the life
of the protoplast, it follows that it is as incorrect to think of pridoplasm
as a liviniT substance a])art from its oi'^'a ideation into protoplasts as it
would be to think of pndeins, or water, or carbohydi'ates as li\ in«i- unless
organized into protoplasm.

In that case, the pr(>toplast, rather than protoplasm, heconu's
"the physical basis (d life".

The cimcept of specialized, diffei"(Mitially permeable lay<M's in thr
cell is so convenient in the explanation of cell ])henomena that it has
widespread acceptance. The statement is made in a curnMd text <>n plant
physiology, with no implication of uncertainty, that "The nucleus, plastids.
etc., as well as the cytoplasm, have surface membranes on the innrr and
outer surfaces which perndt certain substances to pass through whilr

Membranes in the plant cell. 1

197

1 • ..r ^.. Thp cell is able to carrv on nwiny diff^T^'Ut
keeping others from doing .o. ^^ ^ /^^^^^ ^,^^^.,,^, mend>ran.. which
processes at the same time, owing ^^ >t'^^' ^.^

urround the different structures in it R. m. ^^'J

The view that membranes surrouml '^" /^ ^ •;"^.;: ^,,^,^, f,,,,

the behavior of a ^-^^^^'^^;^^^;^^ ^ on which

hut it is granted by its adherents that ho ^^^^ ..p.nuuntal

it rests is inadequate. Ibd.KU, wh.; ha> -^^^^^ ^' ^ ^.,^,, ,^,.,

work < ■•■«" ;■»*;:':;:,;:;,:;; V L,!.!- •■«- - • ■'■

"hislicr iiborlmupt uu'lit - (102H). „;„„:„„ uoi -ulv .m its ^wii

11. fi^iv fiirtlu'i' invest iir«inon. n'" •":

othov lines.

I-HKVIOIS INVKSTir,ATl«»NS

„.o plant vacuole by rapid plasmolysU. "-""i.^t' ">;,:,,. ,., ,.«.. lo-U.... up..n
..ro^rties of this layer, which he calkn < > " ' ' hu, p ■ bablv ,hc .livis,,,,. he .a«
,.e isolated t.moplaBt a. living. ..r.ce .t '"•^' '";".;.,,,,„,' .,„„,„. „ „.,. fact that the
resulted from surface tension orces alone. ^^^"^^^^ „j, „f ...e enti.e livn.«

.Ufferential pern.eability "^ ^->^^^ t , ^t .!r,„na,es as an artifact.
,,,ot.plast that it seems >'"'''^^'> ''^^t ,, ,.„ ,, ,„,f„..e of a plan, protoplast rctauud

Pr..KK (';2^J-— ;^ ;,:, „i,„ acid, altl .h. «, he surface layer

.lifferential ,«-ru,eabdit\ for<l>e. ,.vt„plasu..

.as ruptured, the dyes pc-netrated ^"^ J'^' Z l^,,,.;.,, .,„„„.., i„ physical pro.K.r.ies
In view of our present ''"';;; 7'*-; . .,.,„ ..„.„Hbu...>ns ,0 ,he physiological

:^.^^fr:eir::K c^i J .- -- -^ '■ ^ --'-

Has been directed toward the "I'-f '"V' / '^7 ! ; . .,p,„s„,.wan
„bservati,,no„linarilyfailston-vealsu. .al > ra^^^h^ ^^^^^_^_ ^„„^^ „^,,^ ,, ,

interface, in the plasn.olyzed "V""'" ''>^ , . ,' , ,.„ ,„„..,. layer distin,., fro„, the
fU-ld iUumination. was able <"/'.^'"' ''; ' ' 'j^ „, r,„«W„7- protoplast. In the first,
.....uainder of the plasn.oly.ed ^>""^';^:f^„,„,, „...„. no movement. su....estin,
,Uolayerc.mtaiusfincpart..lesnwap.dm tj ^ ,,omo,ene.ms. lu S,.rooyru a

H,at it is in the ,el comlition: .n '^J^ "J^.^^J:,,, Uu.... amis which .K-rs^

--"-^-7"— t X': in e «i are HUe the material posi,,. th- -r

!;;r£ .nt,"n.C;i^ . Ceany discernible iu cases.

198

Plowe

KiJSTER (1910) describes the appearance of a fine membrane in cells of Allium
treated with acid on deatli from rapid deplasmolysis; this later collapses against the plasma
mass. He reports (1926) that when a protoplast divides on plasmolysis, a thin strand
of tonoplast may persist, running through the strand of cytoplasm connecting the two
portions of the protoplast. The strand of tonoplast becomes evident on "foamy degener-
ation" of the cytoplasm. In another paper (1927) he describes the occurence of naked
vacuolar membranes in the sap of ripe solanaceous berries, where they could hardly be
considered to result from the use of reagents.

GiCKLHORN and Weber (1927) report that when mesophyll cells are placed in
conductivity water or in isotonic solutions, the vacuole, surrounded by a thin layer of
protoplasm, may contract, although the remainder of the protoplasm remains next to
the cell wall. When these cells are placed in stronger solutions, the protoplast undergoes
normal plasmolysis and the vacuole contracts further. The thin layer about the vacuf)le
may be a distinct osmotic membrane.

On the basis of measurable potential differences between vacuolar sap and the
external artificial sap in which cells of Valmiia are immersed, Osterhout, Damon, and
Jacques (1927) conclude that the inner and outer layers of the protoplast differ.

THE MICROMANIPULATION METHOD

Some time ago Pfeffer (1897) remarked that the question of the presence of
differentially permeable membranes * 'Would be definitely answered ... if it were found
that substances commonly present or artificially introduced, diffused through the central
mass of the plasma, but did not appear in the vacuole or in the water outside." Pfeffer's
idea can now be carried out, using fine glass pipettes and needles in connection with a
micromanipulation device.

The original term, "microdissection", should be abandoned, for dissection implies
dismemberment of a dead organism, and micromanipulative methods are now associated
with the study of the internal physiology of the living cell. It is bv such means that the
internal conductivity, the internal hydrogen ion concentration, and the internal oxidation-
reduction potential of active, living cells are now being investigated.

Less work with micromanipulation apparatus has been done on })lant cells than
on animal cells, because the layer of protoplasm in the plant cell is relatively thin and the
cell wall offers an obstacle to the insertion and free movement of needles or pipettes.

Kite (1913 a, b; 1915) who was the first to publish extensive reports of micro-
manipulation experiments, attempted to investigate the consistency of cytoplasm and
nucleus and the relative permeability of the internal and external cytoplasmic layers in
plant as well as in animal cells. It is unfortunate that in many instances he made no
distinction between dead and living protoplasm. This lack of discrimination causes
uncertainty in interpreting his accounts.

Seifriz (1921) employed such plant material as Vaucheria, pollen tubes, and
bread mould in investigations of the physical properties of protoplasm, working chiefly
on expressed cytoplasm.

Chambers and Sands (1923) used the pollen mother cells of TraJescantia, in some
cases piercing the cell wall and in older cells removing the wall before dissection. They
directed their attention towards mitotic structures. Here again the distinction between
dead and living structure was not always carefully drawn.

Membranes in the plant cell. I

199

scKTu (1927) contributed ^ i^^^^^i;:::^:^!^^ ^

plant cell indisputably alive and >n "'''-'^'j^«"^27„,,d,es with horizontal tips, p.ercmg
Flla Symphoricarpos, and ,Sp.roff;/m. "^ "^'^ ";.^„,„t were made ,K,s8ible by the

;, ..,.1,.* ".«" '7,;;r t x»"i^-v'' "^ -" «>•""""" » ""' '

pipette free to move m anv direction,
with the cell wall.

?,Itoit of matorinl from a nucrop„K-.to.

„HA,. ,U-:VKHSUU.K '^•^'•'^\.'^,^;:^,.,, ,,„,,. Temporary

Protoplasts, like organisms, may '^J^^ ,,_^^., ,„;,„ n^ade familiar by the

...an^et-n l^He^ ~n;-;;:S; .'S.re arises: Is i^ -J.^n^ t

suggested b\ worKim

200

1* 1 () w e

•guarantee, for them, that the behavior of the protoplant ha^5 been normal tlir(>u«iii(>ut
the investigation. Sueh signs as cessation or increase of Brownian movement, aggregation
of granules, sweUing, must constantly be watched for. The appearantte of acidity ma\-
be used in detecting local injury. (Chambers and Pollack ( 1927) and ( 'hambers, Pofj.ACK.
and HiLLER (1927) have shown that the reaction of the proto])lasm of certain animal
cells changes from its normal level (pH 6.7 — 7.5) to as low as pH 5.3 on fatal injury.
Rezntkoff and Pollack (1929) state that, although a churning inoticm with a needle
or pipette causes a distinct temporary local acidity in the cytoplasm of Amoeba duhi/u
the ordinary quiet iusertitm of a needle, or even the injection of Xa(M or K(1 solution,
causes no change in reaction.

PROTOPLA^^MIC STHEAMIXG IN PLANT '^KLLS AS A VUKVK ON

NORMAIi CONDITION

Changes in protoplasmic streaming offer, in many plant cells, a s.Misjtive indicator
of changes in the condition of the protoplasm. Wliile protoplasm in cells which normallv
show streaming may cease streaming without having undergone appreciable injury, it
is also true that known injury is regularly accompanied by changes in the manner f
streaming or by (u)mplete cessation. If we disregard all results in which streaming is
affected, we may feel reasonably sure that the behavior of the ])rotoplasm in other respects
will not be abnormal.

SCARTH has reported that a strand of cytoplasm traversing the vacuole may
continue streaming while stretched by a microneedle. Taylor's experience with Nitflht
and Seifriz' with mycomycetes were cited above. To these may be added the follow ing
facts from the present work. When a needle is pushed intt) the protoplast of an onion
cell, carrying a layer of cytoplasm with it, streaming will take place in the layer about
the needle in a wholly normal manner. Strands may form ruiuiing from the needle to
the wall. The nucleus may move out along the needle to its tip. Further, if the needle
is thrust through the protoplast until it touches t!»e protoplast-vacuolar surface on the
opposite side, then partially withdiawn, a strand of protoplasm will be ])ulled out from
that surface into the vacuole, and in this strand, too, normal streaming will occur. Or.
if the needle is withdrawn from the plasmolyzed protoplast, pulling a strand of [trotoplasm
out after it, streaming will occur in this strand both towards and away from the [irotopiast .
A protoplast may be pinched in two with the needle, and yet streaming in the two portions
continue undisturbed. An opening may be made through the j)r()toplast which brings
the vacuole into comnnmication with the external soluti(m, and yet streaming continue
uninterrupt^^d.

These results encourage us to believe that, with careful manipulation, it is p(»ssible
to carry on intraceUular investigations of plant protoplasts as well as of animal protoplasts,
in wliich the ])rotoplasm under observation is normal insofar as any organism, organ.
or tissue can be ctmsidered normal while under ex])erimental conditions.

MKilOMAMPULATIVK STUDIKS OV TIIK IMIKSl^^NCK

OK M KM HH A NKS

Evidence from microdissection is generally in support of the presence of differ-
entiated layers at protoplasmic surfaces. Chambers (1917) states that Paramoecia and
marine over can recover from dissection only if a new film, or series of films, is formed,
cutting off the injured regitm.

Membranes in the plant cell. 1

2^1

r «i iirf'wi' is fatal to somati*'

....Us ,r... vari..u« a„in.a. tissues ^^^:j::Zne.i m....... u.av .ht-.s, an.. .-1

SK-KH.z (192.) fim.s that --"'''- ">''7;,.,, „,,„,„„.,...... ,.«—'"-" ^ ^""l' "'-

„ hen the eytoplasn. disintegrates. V\ he n a --^ >> ^ ; ,,,„„„,. ,„a,v .«■ ..ra« n
|aBtpartt.,tearisalways,.>eou.erb<.r<U. . AnAn . ^^^^ ^^^^.^ ^^ ^^^^^^^ „ „„.,

,..«etl.er a«ain .>.v t..e e„ntrae -on '"V" ' ; ~ , :u,,..en,ia,i..n nU... out.-v .aye^
,,.;rder, Thes.- resu.ts in.lieate ..»■ ytese" - 1^ -^^^^ surrounding .•y...|.l-n. fron. ...e
., the protoplast. He des.ri.,es ^^^'^ ''^^^J p.„ „,, ....eleate e,v..n-.Ky.e of
vaeuolar n.enduan.. of ,lu- p.asn.o.yv.e.l o, on • ^^^^^^^ ^^^ . , ,„,.„„.,,.

rryplol,r,„„'ln,.^. Skik.u/. ( H.2.i) ...■s.v.>,e. an out.,

whieh is „,od..rate.y elastie .,,,.„.„, „,,. „, i„„.,est u. v,e«

Memhranes surroun..n.« th.^ e, nt.a.t.le v. („„ „, ,.,„„ra,tde

,., ,,,,v,.s report tha, vaeuo.es in the «;";;;;;'; '2" outraeti.e vaeuo.e ,„ .Vu.o...,a
vaeuo.es (I92S). K.tMI^I:^.) aeser,..es u». .^ ^^^^^ ^^_^,,^ ,,,,,,,,, ,^^

„, .,f hi... eonsisteney and adhesiveness, a,. .» n a ^_^^ ^^_^^^^ ^^^_ ^^^^^^^^^ ^^__. ^^^
as of i-reater density. ..antheendoplasm. """'•'„ ,.;„,,, ..„„„^d. to indent, rall.r

,han pieree. wi... a l.lun, ne..dle. . AV .. R ( «" ^ , „„„ .,f hi... .ou.s.s.enev .

..„.uo.e» i.. K«,>'..'-.^ ...ay '"■";;'"■:.;;: ..' "..iarv vaeno.es n.ay son.et.n.es ..rs.s,

an<l t..at s.,re.ldy ren.an.s of the v.a..s of

after t..e va.i.o.es hav.- fuse.. .,,,,,, „f „ „„..l..,u. „,e,....rane

'<•'.«• """•-■ f"';.' ""^'.T;;'., T Ir I definite n..elear n,e,n..ra,.es for n.ar.ne

credits, by eyt.,...uists. '-'"='""•''•''';[,"„, X.Hun,^. a.t..ou.h, he di.saurees «.th

tlie idea t..at ,uos, p,.otop.asn.,e surfa,. s p. ss.« ^^^^ „„,h.arf..nd ,. «..>.-

^definite nu.-.ear .nen.bra,.e. w..,e.. <"' « - • " ^ , , sk.kk... , .92. , re,.,..s disseet....

..rawn with a pipette, i.. t..e ".-"»■ ;;''! J "^Xt ,, A '.'•■ S.v.m. ^ V-^^O bel.eyes

„„ the degenerate n,en,..ra,.e ,on. '- <J ' ^ ,^ „, „„,,,. „., „. worked is fhnd ,n ,..e
that t..e me,nl.rane of t..e nnel,-, n> the pl..nt
living condition

... ., ,|,.finite diffeie..tial.oli of th.
,,ide„eO .... ......V Oiffor..... sourees ^^^- ^^^^^^ ^ „ ^^,,„ ,..,„ ,,.,, ,„., ith

Uversof protop ■•-. i" the livi"^ '"'''L"''",'' , ',., t f •'- -rf-e .ayer of ,..e ...neus

th;M,..cleL. .. i.. the last ease. poss,b.y d,f..a^^^ _^^^^ ,,_,,.,,,^ „„., „,.,, .„f,er-

ttf. Certain points render 'h.-- -■<''■■.' ■-", .^ ,,„, ,.„..,„ain to what ..x.ent

Elation re....ar.y oeeurs in ,..e ---;, ' , \.„..,„.,a,e p. -.. para..e.s th.- -^

t,.e or«ani.atio,. of the .-'"l''-^ /; ,;.,,, .,,,uf,e,d studies, evidenee for pi....-

1 II With the exception oi •
'!Z^l ."....a.ed on .ivin. protoj^st. ^ ^^^^^,,,,,„„ ;„„ , , ,<,„.vvle.l.-'.-

„(.linV,T>..i.itum..fs,..-l.r..l;i>.'.''>. .1.

202

Plow e

TEHMINOLOOY

In discussing- the litcraturo, th(* torrninol()<ry of t\w papers iindor
consideration was lar<i:ely followed, but consi(lerabl(.' confusion exists.
De ViUKS called tin^ layer surrounding the vacuole the tonoplast and that
at the outer surfaces of the protoplast, the ectoplast. Pfeffer referr(^d
to tln^ two as plasma membranes. From the first, some difficulty has
arisen from the occasional nsc of the word ''membrane" as synonymous
witli cell wall. Now the conjmon use oi the word "membrane" alone
to desi<i-nate an osmolic membrane makes it a physioloo-ical rather than
a pundy structural term.

]:)K Vhies^ "tonoplast" presents fewer difficulties. It is this term
which the present writer has us(^d to denote a distinct layer of cytoplasm

about the vacuole.

"Ectoplast", however, is wididy used in a wlndly different sense,
[n a myxomycete Plasmodium or in a protozoan, the ectoplast (U* ecto-
])lasm is a thick lay(M' of cytoplasm whi(di is certainly not the outer "plasma
membrane" in eith(M' a structural or a physiolojrical sense. Mast (1924)
has given us the term "i)lasmalemma" for the thm (^\ternal layer whi(di
is the "membrane" if an osmotic function exists. This term seems free
fnnn the difficulties ])resent(Ml by both "ectoplast" and "plasma UK^n-
brane". It has seemed permissible, therefore, to extend its us(^ to the
botanical world, and to employ it to denote a distinct, differentiated
layer on the {>ut(M' surface of the plant protoplast.

Since plasma lemma and tonoplast are themselves cytoplasmic
structuiMN, we cannot refer to the lay<'r of protoplasm betwiM'ii them
simply as cytx^plasm. It obviously is not ectoplasm. Endoplasm, on the
other hand, would include the tonoplast. No term in use at present
adequat(dy distinguishes this portion of the cytoplasm. A new one seems
necessary to avoid confusion. Mcsoplasm is proposed as indicative of its
position in relation to tlu' other layers.

Plasma lemma and tonoplast, then, will be used hereafter to denote
differentiatiHl layers at the interfaces betwecMi protoplasm and wall and
protoplasm and sap, respc^ctively, and uKsoplasm for that portion of the
cytoplasm which lies b(*twe(Mi tlu^e layers.

METHODS AND MATERIAL
OiiAMBEKS (1924) describes micromanipulation methods in general,
and Sc^vRTH (1927) and Seifriz (1927) (lescrib(> the application of these
methods to plant cells.

MembrancH in the plant cell.

203

c

. ,. .viri of this W(»rk was the inner
The material used for the greatei P;^ /^^ ^^^ ^,.,„,,na <.uons,

,esulls ohtained with other cl. ben MM ^^^^^ ^^^^^ occasionally a

oil gives a layer only oiu. ^^^'^\ ^^ /' J ' ^.^.,^.., ,, it. This provides ex-
;^!;ch ot large, -lorless nn^soph^^^^^^^ Sections of the stripped

'optionally favorable ma on. foi ^^^ ^, ,,p ,ater in the mu^o-

., -dermis may he mounted m ; J^^ ;;" ;^^^ ,^,,,,a by horizontal m«^.ll|'>
iuupnlation moist f -^ -•' ; J^ ' t luetliod Scunu has us.l .. h
^^,^ pi.vee the ^'lU walK Ihl • \ , ,,, „,.y .n.pl- tin

..her plant cells in his -ovk Mo ^,,i,, ,, .pidermis are p asmy-

H^thod described by SKinuz and -H.^ 1^^^^^^^^ ^^^^^ ,, ,,.

ivzed in IS percent (O.ob M ) ^'^^ \;;^^^.^^^,,.,sts are well rounded awa>
^.out one half its original ->!-- ^^ ^,, ,, the plasm.dvzing .>lu-
,rom the end ^valls after ^^-;;;^^ ^ , ^ ,,,, transversely to th. h.ng
,,,, A strip is then cut xsth ^^-' ^^,^^,, ,,,,.... the end wal

,,is of the leaf, on a glass slid. ^^J^;^^^ u.^ untnuehed and

^--^ ^^- ^^''''''''''' '''■ ""7 " t-mi ^nioumea in a hangu.g drop o,
,^,^.^^j,,,a. This section of ^T^^ » ^,, ,,..,• .f th. nucn-

...iudyzing solutKm on a -- ^ ; ,, the cells pointing towards

manipulation moist (dwunbei, th ^^^''\yl ,^^, ,,,terial may 1- held

: i'u end of the moi^ ^'^-y;^-'';., ; 1: A dissection needl.- with
,. pi. by a i>htnt needle wi h ^ ^^ J^^,^^, ,,,.,,u the open end>

, 1 oit/ontal tip now has access to th P I ^^^^ y,,,,,u^uon on the

■'"• :::r:^:::zzi::::'i' •■ -""'■- "■*

movement oi 111' 'i" :^|j

..„ plant u.ar,.n.l is .1ms 'l-;;^' ,.,,,,,,, „, ,„i> w...k. Uu, ..u,.las-

,„„,,,,.,i ....„s W...V ^''-\^-''' ;;■,..„ .... ;.s...i u. ...u.k.- ..>-•

;,flrr plasmolysis.

f niter hver distinet trom

;;;,, ....ovlas.,. and 1"-";; ;;.""! ,'";,.,, ,i,„.., .naterial fro.u ....■

• 1 A fri^in (Hie aliotnel. I oi

distinguished now on

204

P 1 o w e

Membiaiu's in the plant ct'll. I

2()r»

protoplast ^vill contiiuic to flow into thr strand as the needle is drawn
back, then this flow will cease, and the material coniposinji- the strand
cloiij2:at(^s as the needle moves. The lenp^th to which the mesoplasm can
be eh)n^at<'d is not very <?r(vit. When this limit is reached the mesoplasm
ronnds into (lro])lets, while the plasmalemma persists as a slender thread
conniM'tinji; the droplets and forminjj: a lay«M' over each droplet (Kiii'. 1 ).
Tf th(^ n(»edle is drawn still fnrther back, the thread stretches and th<'
balls are carried farther apart. If the ne<4le is moved towards the proto-
plast d^'din, the strand contracts and the balls reapproach one anothei-.
They maintain their ji^lobnlar form mitil actnally in contact. If the «;iobules
and the conn(H*tinji: thread were composed of the same material, we should
(*xpect to see the same shapes shown as the <i:lobides reapi)roach (nie
anotlKM' which were shown as they sepai'ated and ronnd<Ml \\i>. just as

-<2> <S>-

Fi^r. I

an elon^'ated soap bnbble passes thronjxh the same foi'ms when allowed
to shorten as it did when eloiij2:ate(l. Hnt no si^'n of fusion of the droplets
appears, until thox are actually in contact with on<' another. Similarly,
a globule i'eapproachin<2: the protoplast itself must actually c(une intcv
ccmtact with the mass from which it arose l)ef(U'e it loses its spherical
shape and starts to flow back into the plasma mass, even though the
two are ('(umected at all stages by a thread <>f plasmalemma.

It has been known for some time (Chodat 1911) that when meso-
plasm is included in strands persisting between protoplast and wall on
plasmolysis, the cytoplasm rounds into balls on a thin threa<l which
appears to arise from the outer layer of the protoplast. The behavi(»r
of the protoplasm on micromanipulation is evidently am)ther mani-
festation of the same tendency. The concdusion that the outer layer
and the inner cytoplasm are distinct seems inescapable. When a drop
of homogeneous fluid is elongated until surface forces cause it to si^parate
into droplets, these droplets are wholly free from one another; no con-
necting tlii'ead will iv-main. A highly viscous fluid such as tar or molasses

If »

*•>

^

FiL'. '2

,nay be pulled out into a long, slench'r thread, but th.Mv will nm be
droplets on this thread. The layer forming the thn-ad and the layer
i'ormnig the balls must l)e distinct.

PLASMALEMMA IN TllK DKAD I'Ut HOPLAST
The smooth surface of a dead, swollen i)rotoplast can actually be
torn awav bv the microneedh- (Fig. 2) allowing the mesoplasm to fh^w
out into the' surnmnding m.'dium as a shreddy, granular mass which
does n(d mix with the water. |

This brittle external layer is the — ,...— «t-

structure which Pfkffkh rup-
tured in acid-killed cells by
de])lasmolysis. allowing dyes to
penetrate through the crack>
so formed into the mesoplasm.

It seems to I'etain some of the
tlifferential ])ermeal)ility of its

living condition, but otherwise

is very different from the fluid,

extensile living layer from which

it arose.

Swelling of the pr(d«>-
plasm toHows the death (d'
unplasmolyze(l as well as plas-
Uiolyzed cells. Since the ])roto-
plast of an iinplasm«»ly/e<l cell

is in ccmtact with the wall, it is impossible to determine wh.'ther the dead
plasmalemma can be t(^rn awav from the mesi^plasm. but it a needle i>
pushed throng hthe wall until it pierces the outer layer Imt not th. toim-
plast the swollen protoplast willc(dlapse until the tonoplast lies almost flat
,,o,nnst the wall. Appaivntlv the dead plasmalemma held back the thnd
which dist('nded tluMmsoplasm, amlits pumture allowed t he tluid to escape.

It is m» easv matter to puncture or tear the plaMualemma of tlir
hving pn»toplast. ^Kven a very shar]> needle carries a layer of protoplasm
Nvith it as it enters the cell: tin' needh- invaginates. rather than pierco.
the pndoplast (Fig. 3); the plasmalemma lie< next the needle, and the
mesoplasm and tonoplast are also indi'Uted and carried m. When the
needle is pushed through the ])r(»t(adast until it traverses the vacuolr
,,nd passes thromrh the opposite' si(h'. the second protoplast wall stn-tclie.

FiL'. :i

Membranes in the i)lanl cell. I

207

206

PI owe

nnd covc^rs the advancing needle like a tent. It is evident from tlie move-
ment of the granular mesoplasm that material foi' this cone and for the
layer covering the needle is supplied by flow from other regions, but if
the needle is pushed through far enough, particularly if it is moved
I'apidly, this flow will be inadequate and a strain will result which may find
relief in one of several diffe'rent ways. The plasmalemma may be ruptured,
resulting in death of the cell. The protoplast may be completely punctui'iMl
at the point at which tlui needle enters and at the opposite side wluM'e
the needlfi tip stretches it into the form of a cone. In this case the severed
edges of the layer about the needle come into contact with the edges

of the puncture in the cone and
fuse. The protoplast resumes it
normal contours, but a tube of
protoplasm now surrounds the
needle which joins the proto-
plast at both ends (see Fig. 4).
Periiaps the situation may be
more clearly visualize(l by
analogy with invertebrate em-
bryology. The n(;edlr first
invaginated th<* i)rotoplast,
resulting in a gastrula stage,
the end of th(* " archenteron " was then brought into contact with the
opposite wall of the gastrula, and an opening, the "stomadaeum'\ was
form(Hl. The vacuole corresponds in position to the coelom, and is at
all stages completely shut off from the outside solution. This intr^rpretation
of the state of affairs can be corroborated when the needle is uiov^mI
backwards and forwards. Material from the needle flows into the wall
of the protoplast, and materi;U at the other (Mid of the needle flows from
the w^dl of the protoplast onto the in^edle. In addition to these two possi-
bilities, the strain produced by the needle may have a third result.

PUNCTURE OF THE VACUOLE AND CONTRACTION OF THE

PROTOPLAST

Sometimes the layers of protoplasm about the needle remain intact
while those on the opposite side of the protoplast, stretched by the neeiUe,
are ruptured. The opening enlarges rapidly, forming a gaping aperture.
The cell sap is in direct contact with the external solution, and in tlie
case of cells whose sap contains colored pigment, its outward diffusion

Fi^^ 4

>

,.

can be follow(Ml. The hole grows slowly after its initial rapid lormatmn,
and as it enlarges the pn.tnplasm fh.ws t.^gether until finally only a ball
of protoplasm remains, c(U.taining a nucleus l)ut uo cmtral vannde
(Fig o) What is now the outer layer of this mass ci)nsists m part of the
layer originally in contact with the cell sap. The cytoplasm is normal,
the streaming of the graindes continues, and no swelling or appearamv
of Brownian movement results.

An enlarging puncture id' this sort mM inlnM|Uently arises at the
point at which the needle first enters the protoplast, .u- more rarely.

a

e

TTv

(/

Fig. o

at, a point .euu.t. from th. n-.U. as a .vsult cf ,1... fus.-m p.-oduccl.
The last case is more eo.nmon amon^' .ells whiel. have stood for son,-
time in plasmolvzinfr solution. This sug,..sts that the internal stnutu.v
of the protoplast undergoes ,ra.l«al readjnstn.ent to Us new form a .1
loses in extensibility. Knlar-infr pnnetures, not resultn.jr m d-ath of tl.
cell, also occur at times in unplasmolyzed n'lls.

FLASTICITY AS A CAUSE OF l'R..TOl-LAS.MU: ft.NTHA.rri. .X
The fact that the hole formed enlarsres so slowly after its fu-st ap-
pearance suggests that its enlargement and the contraction of t '« P-jOt^'
Em may be due to elastic forces rather than surface forces 1 is hard
o why the rate of enlargentent should change so abruptly .f .t w.re.

208

Plow e

Membranes in the plant cell. I

209

tli(^ result of surface tension. Elastic substances, however, often show
la*>* in recovery, retnrninji* nipidly partway to theii' orijrin.il form, then
conii)letin{^ the retnrn xory slowly.

There are other points in the behavior of the protoplast wiiic-h are
more readily explained on the assumption of elasticity than of surface
tension alone. When a strand is x)ulled out from tin; surface of the proto-
plast and stretched farther and farthei', it will finally break, and contract
very rapidly, almost snapping back, and usually crumpling as it recoils
(ScARTU, 1927, has described sinular behavior of internal cytoplasmic
strands wluui broken.) These strands lag in the final stages of theii' con-
trac-ti(Ui. The retui'u of the final lagging portion to the plasma mass can
be accelerated by pi'essing on the adjacent piution of the protophist.

ELASTICITY IN THE PLASM ALEMMA

If a strand is pulled out from the protoplast but uol elongated to
the point at which the nu'soplasm s(;parates int(> balls, the heavy granuhir
nuiss will at first flow back into the protoplast when the needle reap-
proax^hes it almost as fast as the iummUc itself moves, but at the last will
flow more slowly, so that a portion persists foi' a time as a iijund pro-
trusion (ui the surface of the protoplast. This will also flow back into
the main mass in t'wuv. and can be ma(h' to flow back instantly by pressing
with the needle on an a(lj(uniug ])orti(Ui of the protoplast. Now, such
pi'essnre inchuits the surface of tlu^ protoplast, makiug its curvature
uKU'e sharp. When (Uie lifjuid droplet flows into amdher as a result of
surface tension forcis, the speed of flow will incn^ase as the difference
betw^een the curvatui'e (d' the two droplets increases. Hence, the effect
of such pressure should be to retard the flow, were it caused by surface
tension. Again there is a strong suggestion that it is elastic forces, pro-
bably in the plasmalemma, that cause the flow. If the elasticity were
rt^sident in the mesoplasm, strands of varying thickness should flow with
equal rapididity, whereas thicker strands flow mon^ slowly than those
in wiiich the proportion of mesoplasm to plasmalemma is low^er, and thin
strands arising from the phismalemma ahuie and carrying balls of meso-
plasm move most rapidly of all.

ELASTICITY IN THE CYTOPLASM

The behavioi* of the mesoplasm suggests that it, too, possesses
elasticity in a low degre(\ It will be noted that when a strand is pulled
out fi"(un the protoplast, some lines of granules appear to snap and flow

(

♦>

back towards the protoplast before others on either si(h' of them do.
This and similar phenomena when the needlo is movrd through the proto-
l)Iast are explicable if we think of thr mesoplasm as based on a conti-
nuous, labihs elastic framework. Some ii])rils in sucdi a framework might
be under greater tension than other, and break and contract s(M>ner.
Such a structure has been suggested by Skiffuz ( 1924, 1929) and supported
by SCAHTU (1927).

TEAHINd THE LIVING PLASMALEMMA
The layer of protoplasm in the plasmoly/rd nr unplasmolyzed cell
is so thin that it i^ difficult to demonstrate' that t(>aring the out^r surface,
has wh(»lly different effects from
piercing the entire thickness (d

the protoplast. Hut if our of .:^;S%^

the contracted protoplasmic ___
i)alls resulting fnmi a widening
puncture of the prot(»plast is
used, it can be seen that thr
n(MHlIe may ])ierce the ♦•ntire
thickness (d" such a ball without *'''^'- '^

iniurv. while if ihroutrr surface

is town with a rapid m«dion. the mesoplasui swells aud ceases streamiug,
the swelling commenciug at the ragged edge of the tear (Fig. G).

WHAT IS THE SKrN IFICANCE oF STILVNDS PKHSISTlNlr
HETWEEN PHOTOPLAST AND WALL AFTEH PLASMoLVSlSy
KiiSTKii (1910) has noted that portions of a prntnjdast separated
by plasmolysis frecpiently do not fuse on deplasm.dysis. This fact is
occasionally cited as evi(h'nc(; for th«' existence (d" a distinct external
layer on the prot(»pIast. This is questionable, for two soap bubbles may
likewise fail to fuse when brought int(» contact, and the two porti(uis of the
protoplast, filled with and surrounded by liiiuid, are more analagoiis
to soap bubbles than to li(,uid droplets. The Ixdiavior of isolated portions
of prot(»pIasts brought into contact with (Uie aiudher could be further
followed here. If the soluti<ui in which plasmolyzed, sectioned material
is mounted is diluted, the protoplasts will niove toward the open eud of
the outer cells as they swell, until they partially protrude or are even set
fre(^ in the surrounding nu'dium. A partially protruding protoplast can

Protoplasma. XU '^

210

Plowe

be pinched in two with the needle, much as a soap bubble can be dividrd,
but in this case a thin strand of plasnialemnin connects the two portions
(Fig. 7). One portion is anucleate, yet streaniinj>* in it will continue in
exactly the same manner as in protoplasts containing' the nuchnis, even
after the connectmg thread is broken. A ball of protoplasm tiuis s(^paratc()
will not fuse when brought again into contact with the other portioti
of the same protoplast or with another protoplast, which confirms KiisTKii's
observation. However, when the two balls of protoplasm <irc scparntcd
again, a newly formed plasmalemma thread will connect tluun. TIk'sc
threads suggest plasmodesmae. It is already known that plasmodesmae
must arise secondarily, since they occur between cells of differeet genetic
origin in ^raft hybrids (Hume 1913), but whether or not there is true

Fi^. 7

cytoplasmic continuity in plasmodesmae is still an open (luestiou. The
threads seen here originate from a secondary contact of piotoplasm, and
apparently involve no true fusion of mesoplasm.

Strands running from a plasmolyzed protoplast to the cell \\i\\\
are sometimes thought to arise from plasmodesmae. Others suggest
that their presence indicates the continuity of substance between wall
and membrane which Cranner (1919) has supported. The greater number
of the strands disappear after prolonged plasmolysis. The few that remain
can be broken by careful manipulation. Bnt whenever a free protoplast
is brought into contact with the wall, a new strand will form at th(^ new
point ot contact. In or(h»r to remove the protoplast this new sti'and mnst
be broken, and in breaking the first a new contact is apt to be made
and another strand formed, and so on. These newly formed stran«ls
fasten the protoplast to the wall as securely as those persisting frcun plas-
molysis. In one case a plasmolyzed protoplast, lying in a, cell which had
both end walls removed, was rolled over and over by two needles from
one end of the cell to the other, all previously existing strands thus being
broken. The plasmolyzing solution was then made more concentrate' 1.

Memhranes iit the plant crM. I

211

and on further contraction of the protoplast it was evident that numeions
new strands now <'onnected it with this new region of the wall. When
we also c«msi(hM' the fact that the strands betW(H'n needle and protoplast
are fullv as persistent as those between protophist and wall, it seems
that the strands an' more likely the result oi the glutinosity of the [)|;is-
malemma than of continuity of substance between wall and pr(»to])l;ist,
or of the presence of plasmodesmae.

TllK roXOPLAST

The tonoi)last is the most striking of the differentiate(l protoplasmic
layers in that it can be completely isolated and still retain many -d' the
properties shown in the livinii* cell.

DISTINCTNESS OF TllK TONOPLAST IN 'I'lIK LIVIN(r (KLL

The hr'st indication of a distinction between tom.plast and meso-
])lasm in the normal cell is obtained when a needle draws out a strand
from the wall (d" the vacnol(>. Frecpiently the LTanular material in such
a strand will round up into balls (Ui a hyaline, toiioplast thread, much
as the mesoplasm rounds into balls on an external plasmalemma tliP-ad
when a strand is pulled out from the outer surface. The fact that iliis
does n(»t invariably occur is probably <lu.' to the fact that an internal
strand cannot be elongat^'d to any great extent, whereas a thread pulled
out from the external surface can be ehmgated to several times the leno-th
of the protoplast, if necessary, in order to reach the limit ..f extensibility

of the mesoplasm.

On the death <»f the cell, tlu^ material composinii- tin- irans\acu«»lar
strands, as well as the remainihM- of the cyt<»plasm. swells. The meso-
plasm in the strauils flows together t»» form globules (Ui a thin, hyaline
thread. Threads which originally contained n«» Ln-anular material d. i.ot
swell visibly, and show no rounding into droplets. Apparently the matrrial
composing the tonoplast swells little, if anv. on .h'ath.

ISOLATION i)V TllK TONOPLAST
The tonoplast will (d'ten remain intact when a suibh-n, tearing
motion of the uvmUv ruptures the plasmalemma. The mesoplasm shells
and contracts, slipping back over the surface of the tonoplast (Fiir. 8).
Plasmalemma and mesoplasm can be completely removed by the needle,
leaving th«^ tonoplast free, as a transparent bag filled with cell sap. 1 he

14-

212

P 1 () w e

Mombrant's in the plant cell. I

:^i:^

toiioplast is clear, colorless, hyaline, apparently qnite fluid and only
slightly sticky. The isolated vacuole swells in a diluted nirdiuni and
shrinks in a, concentrated one. If it contains colored sap (as in cells of
the beet, occasional cells of Elodea and onion) it is (evident that the pigment
is retained. If placed in a colored solution, the dye does not penetrate.
Since this layer retains its differential permeability, it seems not unlikely
that its physical condition may also closely approach that of tlu' same
layer in the normal cell. When the tonoplast is elongat<'d by the needle,
it breaks or separates, like an elongated soap bubbl(% into small globules
containing cell sap. Only short strands can be pulled frinn its surface.
Torn, it flows together mto a transparent mass. In the isolate(« tonoplast
we have a. striknig instance of a fluid membraiu^ possessing differential
permeability approaching that of the living protoplast.

Separation of the tonoplast (Hi death also
occurs in cells mounted in tap water, so its
isolation cannot be attributed to the plas-
molyzing agent. In iKui-plasmolyzed cells cut
by the razor in sectioning, the ton(>plast f(>rms
a layer distinct from the swollen mesoplasm,
and can easily be separated with the needle.
The separation of tonoplast fi'om mesoplasm
can b(^ follow(id as an unplasmolyzed proto-
plast swells as a I'esult of injury with the
needle. Starting at the point of injury, the tonoplast appears to blister
(d'f in a manner recalling the separation ot the fertilization membrane
from an echinoderin egg. At times, the tonoplast is not. at first, distinct
from the swollen mesoplasm, but becomes distinguishable lat(^r as the
mesoplasm shrinks away and collapses against the plasmalemma and
wall. The tonoplast isohited from unplasmolyzed cells behaves on mani-
pulation just as (h>es the same layer from plasmolyzcMl cells.

The number of different ways in which separation of the tonoplast
can be brought about is strikini'-. If i)lasm(dysis in* deplasmolysis is very
lapid the tonoplast m;iy separate from ths^ rest of the cytoplMsm. 11 the
protoplast is cut by the i-azor, the tonoplast may sei)arate while the rest
of the cytoplasm swells and flows tog(;ther. If an injection oi' the inser-
tion of a needle produces too great a strain in the cytoplasm, th:j outer
layers may be ruptured or swell, and the tonoplast persist. If the nucleus
is injured, the tonoplast will remain distinct whih^ the rest of the proto-
plasm undergoes marked changes. Certainly a layer which appears in
both plasmolyzed and unplasmolyzed cells, whether death occurs sh>wly

Fig. 8

>

or smhlenly, whether (h'ath is caused by mechanical injury «»r l)y toxic
l)roducts of nuclear injury, which is invariably distinct fr«»m tli. nmainder
(d" the pr(>toplasm and which always poss«^sses the same pi'(.[Mrt ir<, must
r<'present a layer distinct from the remainder of the protoplast in the
living condition. Taking into consideration the difference in behavior
of the granular and hyaline material when a strand is pulled .Mit int.»
the vacuole from the vacuole walL it seems certain that tlir livinir i)rot«.-
plast possesses a ditferentiated layer about the vacuole.

riiK Nrn.Krs

The nucleus in the oinon cell is iri'egular. usuilly flalh'iird and
roughly disk shaped. This flattened f(»rm jM'isists cvm when thr uucIcun
is suspend(^d m t he middle (d" the vacuole rat her
than lying in the peripheral layer of cytoplasm.
Many nuclei show (h'ep, transparent gr(M»ves
which occasi(»nally pass completely through
the nucdeus (Fig. 9). The mudear substance
presents a finely mottled, apparently alveolar
structure. Two nucleoli occur.

ELxVSTKUTV OF TlIK XrCLFFS

The suggestion id" elasticity and rigidity
which is given by the irregular shape of the
nucleus is confirmed by its behavior when
handled with microneedles. The nucleus may

be pushed slowly from end to end (d" the cell without chauLdng shap.-,
and if a disk— shaped nucleus which is in smh a position that it appears
circular is caught on one rd^yi^ by the neciUe tip. it will be tipp.'d u}) and
appear as a narrow .dlipsc. When the needle is withdrawn it falls into
its original pi^sition and again appears circular.

If i^reater force is applied, the niudeus may be disKMted. but resumes
its shape with striking regularity. When pushed rapidly throni/h the
cytoplasm, the pressure of tlu' cytc^plasm against its five ciLn-s will bend
the nucleus until, in profile, it is sausage shaped. The nucleus siMimrs back
to its original shape when the needle is bn»ught to rest. This experiment
may be Repeated many times, and the niudeus invariably straiLditens out.

The return of the nucleus to its original f<»rm cannot be the result
(d' surface tension forces, since the form whiidi it resumes is flattened
and highly irregular, in no way approa(diing the spherical form towards
which thes(^ forces would work.

Fiu. 9

214

P 1 () \V V

Membranes in the })larit cell. 1

216

Fig. 10 shows the n'turn of a liviiij>' nucleus to its original lorin when
distorted by the needle. The nucleus was so irregulai' in contour that
the needle tip could be hooked under one of the proti'usions and i)ulled
laterally, producing marked distortion, yet after each of several successive
distortions the form resumed was very close to the original.

One protoplast afforded a demonstration of elasticity in the nucleus
without a needle touching the latter. Occasionally, when a needle pierces
a protoplast and passes on through th(» end wall of a cell, the pr(»toplast
will flow through the hole left after the needh; is withdrawn. This is
pi'obably a result of suction, the wall of the second cell being somewhat
stretched as the needle is withdrawn. Flow can take place through a
hole so small as to be perceptible with great difficulty. The protoplasm
which passes through the hol«^ remains alive, for active stn\Mming takes

Fig. 10

place. In the case under consideration, the protoplast continued to flow
through the hole into the adjoining cell until the nucleus was carried
against the hole, passed part way through, and stopped. The hanging
drop was then diluted, and the expansion of the protoplast in the second
cell now forced out the intruding protoplast. Th(^ nuchms sprang back,
and resumed its original form. When the concentration of the drop was
increased, the first phenomenon was repeated, and once more the flow
stopped when the nucleus plugged the hole. Again dilution of the drop
forced the protoplast back, and again the nucleus resumed its original
form. Tts rigidity was evidently too great to allow it to pass through the
oi)ening through which the protoplasm flowed so easily.

CHANliES IN THE DYING NUCLEUS: THE NUCLEAR MEMBRANE

The nuchnis undergoes striking and instant changes when the
cytoplasm is torn or when tin; nucleus itself is subjected to sudden pressure.
The first sign of death is the disappearance of the mottled structure;
this is followed by swelling. As the nucleus thus changes, a death wave
passes over the cell, oiiginating at the nucleus and becoming evident

ill cessati(ni of streaming and swelling «d" tin- cytoplasm. Thr nucleus
continues {o swell until almost spherical. The ciuitents of the two nucleoli
form granular masses. In a (M'11 which has hern plasniolyzed Ww nucleus
may remain in the rounded, hyalim- form f(»r som«' time. In an uni)las-
molyzed cell, tin- m(»ttl<'d ap])(\Mrancr reap])ears almost immediately
after the nucleus beccunes spheriral. The marking may be even more
distinct than in the living nucleus, but is c(»arser. The smooth, si)herical
form and the gjanular nucleoli ;ii'e ch'arly uidike those of the living nu(leu>.

In the plasmolyz<'d cell these changes may be deferred f(.r several minuMs
iifter swelling. After the mottled pattern reappears, shrinking sets in,
slowly in the ])lasmolyzed cell, but very rajudly in the unplasmolyzed
cell. In the latter th«' collapse is very irregular,
the nucleus shrinks in some sp(>ts until con-
cave, and from these ;i membrane lifts off

(Mg. 1 1). It is imi)ossible to say whether this

membrane originates from the cytoplasm or

from the nucleus. It is something more (h-finite

than the result of shrinkage. sei)a rating nucleus

and cytoplasm, for when the nucleus is pulled

(Hit of the cytoplasm by the microneedle, the

blistered membrane ^i)^'< with it.

If the nucleus is roughly handled with the

microneedle while swelling, the reai)pearance

nf mottled structure and the c(»mmencement of shrinkage are brought

nil ;it once- bef(»re swelling is complete in unplasmolyzed cells, or without

the lapse (.f time, after the first stages of swelling, which intervenes in

]>lasmolyzed cells.

TIIK NUCLKUS AS AN OSMOTIC SYS'IKM
The following exi>lanation is offered for the behavior nf the dying
nucleus. The nucleus is surrounded by a differentially permenblr mem-
brane. Swelling is a result of the .iction of .icid (from injury) on the
nuclear ])roteins. At a certain point, swelling stretches the membrane
nntil its differential permeability is losl. (Similar suggestions have been
ma(h' for the loss of haemoglobin at a certain stage in the swelling of
<'rvthrocytes.) At this point electrolytes fr(un the cytoplasm and sm-
rounding medium petietrate the imcleus. and the proteins are 'salted
out", bringing alxuit the reapi>earance of the mottled structure. Shrin-
ka-'e (svneresis) follows. The reas(»n that a longer interval elapses before

Fiu. II

216

P 1 () w e

•MemhiaiH's in tlu- plant ct'll. 1

1^1

the mottled structure reapp(^ars in plasuiolyzed cells is that the hij^iicr
external osmotic concentration retards the last stages of s\v(»lling. Rouj^^h
manipulation with the needle ruptun^s the membrane and allows the
electrolytes to enter immediately. In the case of the unplasmolyzed ('ell.
puncturing the membrane allows t\u) electrolytes to enter before swelling
has gone far enough to destroy the semi-permeability of the membrane.
A combination of colloidal and osmotic th(M)ry se(Mns necessary to account
for the behavior of the nucleus.

ELASTICITY OP THE DEAD XriT.Ens

The dead nucleus, removed from the cell ,ind stretched l)etw(^en
microneedles, shows marked elasticity. It may be stretched to five times
its original length and contract to a little over twice its original length.
It if is stretched untd its elastic limit is reached (10-15 times its length)
it tears and snaps back as it contracts, but is now merely a fibrous mass
clinging to the needle tips.

CONFIRMlNd RESULTS FROM OTHER (^ELLS

Leaf cells of Elodea, cortical paj'enchyma cells of beet and earn a
roots, mesophyll cells from the cabbage leaf, cortical parenchyma cells
from stems of young seedlings of Lu/pinns aJbus. and I'oot h.ni's of Triancn
W(5re used as material to suppleTuent the work on onion cells.

Separation of tlu^ tonoplat- 1 as a result of mechanical injury w<is
seen in the first four n.imed, .uul in the List, but did not occur in the
(•ells of Tjupimis.

The form.ition of balls of mesoplasm on a thread of plasmnlemma
was seen very clearly in Elodea, both hi strands pei'sisting between proto-
plast and w^all after plasmolysis nnd in strands pulled from the outer
surface of the protoplast with the needle. In Tnanam ro(>t hairs, such
balls could Ix* seen on strands persisting after plasmolysis and .ilso when
strands wen^ pulled out with the microneedle from the surface of living,
(ixtruded cytoplasm. Ralls appeared regularly on strands pulled out
from the protoplast of the carrot cells, but the smnlhn^ss of the protoplast
and the thinness of the layer of cytoplasm mnke it impossible to say with
certainty, as can be said in the other cases, that a part of the vacuole
w^as not carried out into the strand pulled out from tin* protoplast, and
that its separation into smaller vacuoles was not the cause of the formation
of balls.

,

;

The failnr*' of such balls to form in cahbagr. brct. ami lii)>iih' crlls
is connected with the fact that thr stfands j)idl«'(| frnm iIm' smf.icr n\'
these jU'otoplasts are short, orcliiiarily bi'caking and stalling tn cniitiMcl
before they have cxcrcdrd tin* length of tin* prntopl.ist.

In all the ct'lls, coiiti'action of tlw strand aftr!- ln'«'akinii' i«'Sfmbl«'s
contraction <d" an elastic body rather than flt»\\ <•[ .1 tiair fluid, hrin^r
miH'h morr i*api<l in the first than in thr last ^tagrs. ( '•►nti-action is always
accelerate(l by pi'essurr <»n tin- i)rotoplast.

Cells (d* sevcfal types showed that jninctnfc i^i the entire iliiekne^<
of the protoplast, followed by cont I'act inn of the protoplasm into ,1 h.dl.
was not fatal. This conld be seen most (deafly in HltHlm.

In in> case did liberated cytoplasm mi.x with water. Its l>r|ia\ior
can be bdlowed i)rst in the liaifs of Tr/mmrn. Mere (•yt(tplasni whi(di
is forced out violently swells at fiist. and some partitdes enter into Rrow-
nian moveuH'iit. The mass may l«»ok as if it were comiMisecl (»f discrete
partiides. but when a n«'edle is thrust into it and I'etraeted. the entile
mass is j)ulled out after the needle, finally breakiiiir and (•ontl•aetinL^
Evidently a continuous stnictiire persists, even thouii'h not \isih|r. The
swollen pi'otoplasm shrinks and darkens soon after libei'ation. It then
Ixdiaves like a fairly rigid body when pushed about with tin- iierdle.

(•ONCIJ'SIONS

I he inh'UHil OH(l c.rh'rndl stnfucc hif/rrs of Ihr pidlifpla^in, nj ounni
cells itfdlslinfinishaftlc nnCritscftitirad f/ or fillrnnucrosroitfrdll n. (irr nrrrrfhrlrs.s'
disdncl front ihc rcsl of Ihr ct/htphistn. The coiroboi-ative evidence from
other cells suggests that a distinct plasmalemma and toiioplast. ordinarily
too thin to Ix' detected optically, are regular features of the onraiii/atioii
(d' the plant protoplast.

Ihc })l(isnH(lcnnn(( Is prolccfirc n\ fnnctfon. This is shown by the
fact that it is fatal to tear this surface layer, although the cell is not or<li-
iiarily injured by pr<donged [)rtd)ing. The fact that the proto|)last may
\)c punctured as a whole. (»pening the \aciio|«' to the outside s(dontin.
without fatal effects, suggests that the tonoplast affords ju'otection to
the mesoplasm similar to that given by the plasmalemma.

The jdnsnudcnnno is hiuhhf cinsllc. Hlosticitif is less tnorktd. Uul
dclcchfhie. in the nicsopldsnt. Ihc n miens is dtc nutsl hifihh/ elashr of ihc
annponcnls of ihe oniitn prolojthisl. The protoidasm thus behaves as thoULdi
base(l npoii the continoiis structural frainewoi'k of .1 n-cl. I'ather than

218

PIr

) w V

Mt'inbrant's in tlio plant cell. I

219

111)011 i\ coiitimKnis a(jii(M)ns plinsc like that of an (Mimlsion or suspension.
This is in aii'ircnK'nt with the familiar fact that rapid (',hanj;<' in confi-
ii'iiration of tfic prot(»plast (plasmolysis, (h.'plasniolysis, j)ressnnO i^ ordi-
narily fatal even thon<>-h tho sam(' result, lu'odnced hy jiTadnal steps,
can be f)ron^'ht about without injury.

Plasrnaleiiiina and tonoplast are elastic fluids. Xotinj*' the exten-
sibility of the plasnialenirna. the ease with which it accommodates itself
to chan^^'s in confi^niration, the fact that the ^Tenter part of a pr(»toplast
may flow through a minute opening- without injui'y, a new explanaticm
su'»'gests itself for the phenomena first noted by Nagki.i (1855) and repe-
atedly ('(uifirnied and cit(Ml since. It has p'nerally heen considered, in
agreement with his account, that the fact that exuded or expressed proto-
plasm from root hairs, pollen tubes, V(tn(hrri(u etc.. Ixdiaves as if sur-
rounded by a m(Mnl)ran<' wholly similar to that of the intact protoplast
is ail indicati(m that the pndoplasmic membrane is "autiUHunous",
i)V that the formati(ui of surface memf)i'anes by protoplasm is a result
(d" surface forces at the protoplasm-water interface. Is it not likely that
in these cases we are not dealing* with the reformation (d" a membrane
l)y naked mesoplasui, but that plasmalemma as well as mesoplasm is
forced (Mit, and that the mesoplasm is covered, at all stajres, by the fluid
plasmalemma? The droplets wiiich fail t(» •re-form" a membrane are
then those which consist o\ mesoplasm with no coverini>- plasmalemma.

SIIMMAHV

1. Existing' evidence foi' the formation id' morpholoji^ical mijmbranes
at protoplasmic surfaces in the living* plant protoplast is reviewed
and judjj:e(l ina(bH|uate.

2. Cases fnun the literature and from the author's work are cited to
show that micromanipulati(ui may be employed in the investij»-ation
of this problem without peiceptible disturbance (d" the normal condi-
tion (d' the proto{)last.

\\. Investigation of plasmcdyzed and unplasmcdy/evl pr(d(>i)lasts with the
microneedle indicates:

a) That there is an exteinal layei', oi' i)lasmalemma, surrcuindin^-
the ])rotoplast, whiidi. while fluid is more (dastic and more exten-
sile than the remainder (d" the cytoplasm.

b) That a similai', but less (dastic layer, the ton(4)last, surrounds
the vacu(de.

I

c) That the meso])lasm is mu(li less extensile than the ton<»plast
and plasmalemma Ix-twcen whiidi it lies.

<l) That the nucleus is suri'ounded hy at least one nicnduane id either
cytoi)lasmic or nu(dear oi'i«rin. The nucdear material, while hijrhly
elastic and showing' the ])roperties (da hydi'o])hihc c(dloid. never-
tlndess is part (d' the osnndic system whi(h the entire muleus
(•(Mistitutes.

Depai-tment (d" IJolany, Tniveisity (d" Pennsylvania. Thiladelpliia. Ta.

LlTKUATl'HK CITKI)

CnA.MBKKs, H.. MM". .Micnxlisscction stu(li<'s. I. 'i'hc visible structure of cell prutoplasni
aiul death changes. Amer. .lourn. Physiol. 48, I 12.

— 11)24. 'J'he physical structure of protoplasm as <letermine<l hy mien »<lisseet ion an«i

injection. Cowdry, (hneral ('t^toloyt/. University of Chicago IVess.

— iV- H. PoLi.AC'K. 1927. Micrur^ical studies in cell physioloj^y. I\'. Colorinietric det<T-

niination of the nuclear and cytoplasfnic pH in the starfish r^^. .lourn. (ieii.
Phvsiol. 10, 739 755.

— — ^ S. HiF.i.ER. 1927. The protoplasnii*- |>H nf liviiiL' cells. Proc. So( . Kxp. Kiol.

& Med. 24. 7H(I 7HI.

— il- (J. S. Rknvi. 1925. The structure of cells in tissue's as revealed hy nntrodissection.

I. The physical relationships of the cells in c|)ithelia. .\nicr. .Inurn. Anat. .H5.
385 W2.

— iS: H. ('. Sam»s, 1923. .\ dissection <>f the chnmiosnnies in the pollen mother cells

of Trnfhsrantia rinfinim L. .lourn. (ien. Physiol. 5. H15 820.
('noi)AT, P., 191 I. Princi|)es de hotani«jU<'. .1. B. Haillierc et fils. Paris.
(HANNKK, j^. Hanstkkn. 1919. Peitra^c /ur Physiolo^ne der ZelKvand un<l der plasma-

tischen ( Irenzschicht^'n. F^er. Deutsch. Hot. <ies. 87, 380 391.
<;i<'KLM<)KN, .1.. iS: F. VVkrkk. 1927. I'lu'r \'akwolenkontraktion nnd Plasn)olysef(»rm.

Protoplasma 1, 427 432.
HoBKR, P., I92(). Physikalische ('h«'mie der ZelN- und der ( Jewehe. \\ . Kn»relmann,

Leipzig.'.
HoWLANi), P. P>., 1923. Studies <>n the contractile vacuole (»f Anto*b(i rtrrurnsn and

ParatiHHciuui t/iutlntitm. Proc. Soc. K.xp. Pi«tl. and Med. 20. 47<> 471.
HcMK. M.. I9I3. On the presence of connectin^Mhrcads in irraft hyhri«ls. New Phytol. 12,

2U> 221.
KrrK, (J. L.. 1913 a. The relative permeahility of the surface and intcri(»r p<»rtioiis of
the cytoplasm of animal and plant c.'lls. Biol. Bidl. Marine Biol. I.^il). 25, I 7.
1913 b. Studies on the physical properties of protoplasni. I. The physical properties
of the protoplasm of certain animal and plant cells. Amer. .lourn. Physiol. 32
14() 1(>4.

— 1915. Studies on the jH'rmeahility of tin' internal lytoplasm of aninuil and ]»lant < dls.

Amer. .lourn. Phvsiol. 37, 282 299.

220

Plow t', Membranes in the plant cell. I

KusTER, K., 1910. Cber Vcranderung der Plasmaoberflache bei Plasniolyse. Zeitsclii .
Bot. 2, 689-717.

— 1926. Beitiage zur Keimtnis der IMasmoIyse. Frotoplasma 1, 73 104.

— 1927. Cber die (Jewinnung nackter Protoplasten. Frotoplasma 8, 223 234.
Lloyu, F. K., 1928. The (;ontractile vacuole. Biol. Rev. 8, 329 358.

Mast, S. O., 1924. Structure and locomotion in Amoeba proteus. Anat. Rec. 2U, 88.
Nageli C, 1855. Pflanzenphysiologische l^ntersuchungen.

OsTERHOUT, W. .1. v., 1922. Injury, recovery, and death in relation to conductivity
and permeability. ,]. B. Lijipincott, Philadelphia.

— K. B. Damon & A. (1. .jacqttks, 1927. Dissimilarity of inner and outer surfaces in

Vahiiia. .lourn. (Jen. Physiol. 11, 193-205.
Pfeffer, W., 1897. The })hvsiol()gy of platits. (Tr. A. J. Kwart) Clarendon Press.

Oxfoid.
Pollack, H., 1928. Micrurgical studies in cell physiology. VI. Calcium ions in living

protoplasm. .lourn. (Jen. Physiol. 11, 539 — 545.
Price, S. R., 1914. Some studies on the structure of the plant cell by the method of

dark ground illumination. Ann. Bot. 2H, 601 — 632.
Rarer, ()., 1928. Principles of plant physiology. Macmillan Co., New York.
Reznfkoff, p., & H. Pollack, 1929. Intracellular hydrion concentration studies. II.

The effect of injection of acids and salts on the cytoplasmic ])H of Amoohn (Juhia,

Biol. Bull. Marine Biol. Lab. 5H, 377—382.
ScARTH, (J. W., 1927. The structural organization of plant protoplasm in the light of

micrurgy. Frotoplasma 2, 189 — 205.
Seifriz, W., 1921. Observations on some f)hysical properties of protoplasm by aitl of

microdissection. Ann. Bot. 85, 269 — 296.
- 1924. The structure of protoplasm and of inorganic gels: an analogy. Brit. Journ.

Exp. Biol. 1. 431 443.

— 1926. The f)hvsical pr<)perties of erythrocytes. Frotoplasma 1, 345 365.

— 1927. Xew material for microdissection. Protojilasma 8, 191 — 196.

— 1929. The structure of protoplasm. Biol. Rev. 4, 76-102.

— 1929 a. Protoplasmic structure. Froc. Int. Cong. Plant Sci. 1, 251 — 258.
Sharv, L. W., 1926. An introduction to cytology. Mc(Jraw Hill, Xew York.
Taylor. C. V., 1923. The contractile vacuole in Euplotes: an example of the sol-gel

reversibility of cytoplasm. Journ. Kxp. Zool. 87, 259-290.

— & W. P. Farber, 1924. Fatal effects of the removal of the micronucleus in EupJofts.

Univ. (^alif. Pub. Zool. 2«, 131 — 144.

— & D. M. WiriTAKER, 1927. Potentiometric determinations in the ])rotoplasn\ and cell

sa|) of Nitella. Frotoplasma 8, 1 — 6.
Vries, H. i>e, 1885. Plasmolytische Studien iiber die Wand der Vakuolen. Jahrb. Wiss.

Bot. l«, 465 598.
Wilson, F. B., 1925. The cell in development and heredity. Macmillan, Xew York.

• ♦

•(

T

V/1

MEMBRANES IN THE PLANT CELL

11. LOCALIZATION OF DIFFERENTIAL PERMEAHILITV

IN THE PLANT PROTOPLAST

Janet (^. Plowk

Received for i)ublication. Septeniber S. |!>;jo

111 ])art oiu' (tf this jkijkt (Plowk. \\y.\\) rviib'ncr w.is gi\rii indi-
v'.itiiijr thnt the lixiiiu' pl.int pi'ntojd.ist pnssrsscs diflt'i't'Hti.itrd l;iy<'r>.
rln' pLisiii.ih'imii.i .iiid tlu' tonophist . .it llu' cytnjjl.isin-w.ill .iiid cytn-
plasm-sap intrrl.iccs. Tln' fidlnwiiiii- study was m.idc with tlir pmpoM-
of d^'teriiiiiiinii" whrllKT or not iln- diflt'miti.d jui'iiUMlMlity pnssrssrd
hy tho y)laiir protnpl.ist diirinii- WW is Idcali/t-d in llu'sc pliysic.iMy distinct
layers.

It dot's not, of coiirsr. in'c«'ssai'ily fnllnw that th<' «iitif»' tliicknrs^
of these layers I'nnctinns as an osmotic mrtnhraiic. rvm if it can Im- shown
tJbit the diflcrcntial pcmicahility nf tlic protHpl.ist is ,i pioprriy o| tnno-
plast and plastnalcmina and imt (d" the mcsoplasiiL This distinction will.
howev<M'. be \vai\cd for the j>rcscnt. and discnssrd .iftcr tlic inajoi- pi<ddcni
lias hccn taken np.

From the rcsnlts of connllcss cxpciinnnts on rrcoj-d. .i f.-w concin-
sions may l)c drawn which arc apparently applicaldc to any lixini! cell.
Water can move icidily hotli into the c«'ll and ont The movement <d
water is influenced by the composition of the externnl milieu. Sohites
as a ride move less freely than water, and Ijetwecn different solutes. >iic|i
as urea and sucrose, there may be wide differenc«'s in the ease with move-
ment takes ])lace. Certain substances, sn<di as basic dy»'s. may mnvf
into tile cell readily but move out of it with infinite slowness, so that
an appearance of "one-way i)ei'meability " results.

In the «rreater part of th«' mass of work on cell pei-me.ilnlity. the
permeability (d the cell to n certain substance is coiisideied svnonvmon>
with p«Mn'trability by that snbsiance. It has been reco<rnized in a smallei-
number of cases that ability of substances to p.iss out (d the cell must
also be considered.

222

Pl«) we

Mnnhraiics in the plant ct'II. II

22:5

Two basic theories liav(^ b(MMi advanced to (explain tlie relation of
the cell to water and solutes. The first, and classical, theory is that the
cell is surrounded by a differentially permeable membrane, and thus
constitutes an osmotic system. The second theory is that th(^. intake and
output of water and solutes by cells are the results of hydration and
adsorption in the colloidal protoplasm. Those who advocate the second
explanation do not claim that the osmotic conc(;pt is invahd for vacuolate •
plant cells, where the colloidal protoplasm itself would function as a
differentially permeable membrane around the vacuole; but tfiey maintain
that the relations of this protoplasmic layer with external mediiim and
vacuolar sap are colloidal, not osmotic.

Our present concern is not the specific permeability of the protoplast
for different solutes, or the mechanism by which membranes function,
or the details' of hydration and adsorption. The question umhM' consi-
deration is: Is the permeability of the surfa(!e layers of the protoplast
the limiting!: factor in the permeability of the cell?

Three pictures of the orjjjanization of the vej>:etativ(^ plant cell can

be suj];gested:

1. A layer of differentially ])ermeabl<' protoplasm surnMindin^* a

vacuoh\

2. A differentially permeable membraui^ at tin; outer surface of
the (comparatively) freely permeable mesoplasm. with the vacuolar
sap in direct contact with th(^ nn^soplasm.

;j. I)iff(MTntially permeabh^ membranes at both outer and vacuolar
surfaces of the (comparatively) freely permeabh; mesoplasm.

Kite's microinjection studies on the permeability of plant cells (1913a, 1915)
led him to the conclusion that there is no essential difference between the permeability
of internal and external cytoplasmic layers. Pfeffer (1897) as already menti(.n(Hl.
demonstrated localization of differential permeability in the coagulated plasmalemma
of the acid-killed Hydrocharis root hair. With these exceptions we have no direct evidence
either for or against differentially permeable membranes in the plant cell. Considerable
indirect evidence is to be had on either side of the question, but it must be rememlx ie<l
that inevitable osmotic effects in the vacuolate plant cell add to the difficidty of correctly
interpreting indirect evidence.

TFTE liKLATlON OF THE CELL TO WATER AND SOfAITES ON A

COLLOIDAL BASIS

Lloyd (1915, 1917) finds that acids and alkalies, in concentrations too low to be
osmotically effective, first increase, and in higher concentrations decrease the rate of
growth of pollen tubes in sugar solution. He believes this to be an indication that the
swelling of growth is a hydration phenomenon rather than an osmotic one. The fact

♦^>

f A

iV.'

^' V

ff

that it is possible to explain these results as Lmjvd does without hypothe( atin;,' a diffen-n-
tially permeable membrane should not obscure the fact that it is ('(jually possible to
explain them on a similar basis in the i)resence of a membrane. Changes in the swelling
capacity and osmotic pressure of gelatin run closely parallel, according to LoK.n (1922).
It might easily be ccmceived that the acid used by i.Lovu altered the «»smotic pressure
of the cell proteins within a membrane, and that the volume change was osmoti* . Cse
of acids of different penetrating power at the same pH miuht throw some light (.n the
subject, since, if the osmotic explanation is the correct one, it is the effect on internal,
rather than external. pH w liiih is of importance. ^

I'urely colloidal permeability theories have Ix'en advocated by workers on animal
tissues. .VIaktin Fischkk (1907, I9()S) and 1). . I. Lloyd (19H)) suggest explanations
of volume changes in animal tissues in terms of protein hydration, the first pointing out
the parallel effect shown by different acids and neutral salts on the swelling of fibrin and
of animal tissues, and the second, the failure of the v«)lume changes of muscle in acid and
basic soluticms to follow osmotic laws. Moore and Koaf (I90S) and Moore, Koaf and
Webster (1911) propose to discard the employment of membranes in explanation of the
distribution of solutes between plasma and bloo<l corpuscles, substituting the idea of the
formation of loose chemical compounds and adsorption products. Hut Siebeck ( 11*12)
and Heutner (1913) find that volume changes in animal tissue due to the effect of arid
are distinct from demonstrable osmotic effects present at the same time.

Bakhuyzen (1928) advances a combination of adsorpti<m and hydration theorii s
to explain water intake, penetration «»f solutes, growth, excretion, and accumulation
of substances inside the cell, pointinii out, on data from the literature, that the Hof-
MELSTER serit's holds for th<' penetration of cells by ions.

WATEH AM) SOLLTE DELATIONS AS OSMOTIC IMlENoMKNA

Evidence supporting the membrane theory for plant cells is wholly imiirect. Alt-
hough the behavior of plant cells in plasmolyzing s(.luti.>ns was the starting point of
osmotic theory, our lack of knowledge of non-vacuolate cells (and Dancjeard, W)'!*,
says •11 n*y a pas de cellule sans vacuole") prohibits us from rigidly termini; tlu' plant
cell an osmcjtic system unless we limit the term t<. the relations Ix'twej-n vacuolar sap and

ext^'rnal milieu.

The high electrical resistance of plant tissues (Ostkrhoit, 1922) has been attri-
buUnl to impermeability of the surface layers to electrolytes. Such an interpretation
for resi.stance in animal tissues is corroborat<'d by the demonstrati<m of Hobkr (I'.HO)
and McClendon (1926) that free electrolytes are actually present within the cell. Homkk
shows that erythrocytes placed in the dielectric between the two plates of a comlenser
affect the capacity in the same manner as a conducting substance in the .same |K)sition.
McC/LENDON finds that the resistance to i)assage of current decrea.ses as the fre«|U< iicy
of an alternating current employed increases and the path which ions travel in carrying
the current becomes shorter. Neither metlnMl would serve to distinguish decisively
iH'tween electrolytes in the vacuole of plant cells, unable to pass thr()U«h the prot<»plast,
and electrolytes in the protoplast unable to pass thnuigh a nu'mbrane. HEMiN(iT«»N
(1928), using McClendon's method on beet root tissue, assumes, in order to interpret
his results, that each cell possesses a sin»ile plasma membrane.

224

Plo W V

Mcinlnaiies in the plant ct'll. II

225

THE A^slMAL VK\Aj AS AN OSMOTIC SYSTEM ON THE BASIS

OF MI(U^OINJE(TION EXPERIMENTS

There is diieet evidenee in support of the exi.stenee of differentially permeable
membranes in animal cells. Kite (1913) shows that injected dyes may diffuse readily
through the eytoplasma of Amoeba. Chambers (1924) shows that many dyes which
do not penetrate Amoebae and sea urchin egjrs on immersion can diffuse readily through
the cytoplasm on injection. In the case of the ova he first reported that cytolysis accom-
I)anied the diffusion of the dye, and some (|uestion was raised as to whether the per-
meability of the cytoplasm was not an effect of injury. Subsequent im])rovements in
technique have done away with the difficulty. Today, microinjection is a standard method
for the introduction of pH and rH indicators to which the cell is normally imi)ermeable.
(Xeedham 1926a, b; (^hambers and Pollack 1927; Chambers, Pch.lack and Hiller
1927; (Chambers 192S; Rapkine and Wurmser 192()a, b.) The viability of injected
material is attested by normal movement of amoebae and lack of cytolysis in ova.

I)emonstrati<m by microinjection of localization of differential permeability at
surfaces has not been confined to dyes, (^hambers (1922) finds that, while immersion
in ammonium chloride or in sodium bicarbonate changes the color of starfish eggs vitally
stained with neutial red to the basic and to the acid color res]>ectively, injection of thesi'
solutions will cause change of color in the other direction. Csing the explanation offered
by Jacobs (1920, 1922) for tlie effect of immersion in solutions of COg in sodium bicar-
bonate and of ammonium chloride on intracellular reaction, Chamberss results are to
be interpreted as showing that the hydrogen and hydroxyl ions which fail to pass through
the outer surface of the cells from the external solution diffuse freely through the cyto-
plasm when the solution is injected.

There seems to be little question that the internal cytoplastn of the animal cells
investigated is normally more j)ermeable than the surface layers. Are we justified in
concluding that since this is true for echinoderm eggs and amoebae and a few other animal
ceUs, it is also true for plant cells ? The gap is rather too wide to bridge without further
evidence, and, even assuming that the two cases are analagous, leaves us still iminformed
as to the nature of the relation between cyto])lasm and vacuole.

THE VACroLAR MEMBRANE

Very little evidence is forthcoming on the permeability of the vacuolar membrane.
In general, if there is an external membrane and the cytoplasm is freely permeable, it is
necessary to assume that the vacuolar membrane is differentially permeable, in order
to account for the failure of colored sap to color the cytoplasm, the close analogies between
the properties of de Vries' tono])lasts and entire protoplasts (1885), and the anomalous
contraction of va(uioles noted by Osterhoct ( 1913 a, b), Ki ster (1926), and (4icklhorn
and Weber ( 1927). This is especially true of Osterhouts observation that phycoerythrin
set free from the plastids in the swollen cytoplasm of Grifjithsia does not penetrate the
vacuole. But, unless it is true that an external membrane exists, no one of these cases
really tells us anything about the permeability of the vacuolar membrane, for in that
case the first becomes but the logical result of the differentially permeable nature of
the cytoplasm, and the last two represent retention of differential ])ermeability by one
int^rfacial layer longer than by the remainder of the injured cytoplasm.

1

<^

1

Kite's uork (I913a, 1915) leads him to believe that nn <liffcrentially |K'rmeable
membrane surrounds tlie vacuole.

HowLAND (1928) finds that the acid dyes of the Clark-Lubs pH indicator series
do not diffuse into the cytoplasm of Artivosphaniuw when injected into the food vacuole,
but we cannot safely assume that a food vacuole in a rhizopod and a saj) vacuole in a
pia!it cell are analagous.

K.TKs EVIDENCE AGAJNST JM^ESENCE oF DIFFERENTIALLY

J'ERMEABLE MEMBRANES

Since Kites exp<Miments on plant cells give us our «mly direct evidence on the
exist^^nce or non-existence of differentially ]H^rmeable membranes in the plant cell, and
since no careful considerati<m has hithert(» been given to the direction as well as the
extent t<. which injury may have influenced his results, his ))apers require further dis-
c«ission before proceeding with the investigation nf the j)roblem.

Kite (1913a, 1915) reports investigations using a considerable number of plant
< ells. He concludes that the cellulose wall, and not the protoplasm, of plant cells prevents
the entrance of dyes; from these results and his work on animal cells he concludes that
•partial permeability t<. water, dyes, and crystalloids, is a property of all |>orti<m8 of
proto})lasmic gels"*. His conclusion, however, must be considered ojH'n to serious tiuestion
for the following reasons. His own accounts indicate that in many cases tlu' cells wen-
injured by the injection, and in othei ca.ses there is no menti<m of any criterion used U>
decide whether or not the cells were injured. His intracellular injecti<»ns in plant cells
were alwavs vacuolar injections, not cytoplasmic injecti«ms. No dyes were u.^ed to which
the protoplast is imi>ermeable. Consequently, although to Kite belongs the credit for
pointing the way to direct experiinental investigation of the question of differentially
permeable membranes in the living protoplast, in order to find a satisfactory answer
to the question it is necessary to elaborat^^ up<.n his methods and above all to carefully
<listinguish between permeability as a result of injury and normal permeability.

METHODS

Dyes which <h» ind pciictnitc the cell iiniiiciscd in thciii wciv iiijcctd
into tlic v;icn(»lc. to (h'tcnninc whether the .ippareiu inipenneahility on
iiniTKTsion w.is in reality the result of an nne(|nal equilibrium distribut-
ion between exK'rnal so|uti(Ui and vacucde. and into the cytoplasm.
to ascertain penetrating^ pc^wer when passa^'e throuirh surface layers was

md involved.

A Cuamhkh's microinjectinn apparatus was used in cunnrctinn
with a Peterfi micromanij)ulator. JMpettes having: the same form as the
n(M^dIes already descril)ed (Skifhiz 1927) wvre used, with hnrizontal
instead (d' vertical tit^s. The sealed ends wen- broken «»pen ajrainst the
end of the coverslip rather than the lower surface. When the Inut/ontal
idpette is used, there is no (|uestion as to the exact position of the tif)

Protoplasma. XU 16

226

P 1 o w e

oi the pipette at tlie time of injection, as there often is if one attempts t(>
decide whether a vertical tip lies in the vacuole or in the cytoplasm. In
work on animal cells ther(^ is not this difficulty to be met, and the simplir
typ(^ is satisfactory. The material to be injected was held in place with

a blunt, vertical needle.

Two questions arise with regard t(» the soundness of this method
of investij^niting permeability. First, is it possible to distinguish accu-
rately b(^.tween a cytoplasmic and vacuolar injection? Second, can th«^
behavior of a substance injected into the cytoplasm give informatiim about
the permeability of protopkism in the normal condition? A convmcing
affirmativ(^ answer to the first question is obtained when the indicator
dye brom cresol purph^ is used. Its color changes from the intermediate
(half neutralized) color in the pipette to the acid yellow wIkmi injected
into the vacuole, and to the basic blue when injected into the cytoplasm.
In order to leave no doubt as to the answer of the second (luestion, \vr
may consider as normal only those cases in which streaming of cyto-
plasm is uninterrupted (in Allium and Trianea) or in which the (uisily
disturbed normal configuration of the plastids remains unchanged and
no Brownian movement appears (in Griffithsia). Further evidence of the
normal condition of injected cytoplasm will be given later.

The cells used for this study were root hairs of Trianea hogolcnsis,
epidermal cells of Allium cepa, and the coenocytic cells of Griffithsia
horneiiam. The results for each will be taken up separately.

HOOT HATRS OF TRIANEA
The large root hairs of Trianea boyotensis offer good material loi^
microinjection studies. The cytoplasm forms a. fairly thick layer, and
frequently masses temporarily at the tips of the hairs. Streaming con-
tinues within these masses. The wall is easily pierced. The absence ol
plastids, the transparency of the cytoplasm, the thinness of the walls,
make internal details clearly visible. Injections into cytoplasm or vacuoh'
can be made with no ambiguity as to location. The active streaming
enables the observer to constantly check the condition of the protoplasm
under observation.

FAILURE OP DYES TO PENETRATE ON IMMERSION
Aniline blue, acid fuchsin, and the indicator brom cresol purple
were used. The first two never penetrate the living cells. Cells immersed
in one percent aniline blu(^ for twenty four hours showed no color in cyto-

Mrnibraiies in the plant cfil. 11

227

plasm or vacuole. Acid fuchsin is rather toxic, hence th«' longest immersion
used was two hours forty five minute's in a 0.5 prrernt solution, hi hoth
cases the cells were plasmolyzcd aft<'r removal fmni the dye, to avoid
mistaking coloratiiui of the wall UiV prnrtration of tlu' protoplast hy th«'
dye, and to intensify any color which might br prrsmt in thr vacuoh'
if dye had penetrated. Shortage of the Trianea, aupply pn'vented immer-
sion tests with brom cresol purple, but in no case was dye in the surroun-
ding solution taken up by living cells.

FAILURE OF DYES TO PENETRATE THE rUoToPLAST KKo.M

THE VACUOLAH SAP

Aniliiu' blue or acid fuchsin injected into the vacmde diffuses freely
untd the sap throughout the length of the vacuole is colored. Thr spreading
(d' t\n' dye is accelerated by the motiim given U\ the sap l)y the streaming
cytoplasm It is interesting to note that the streaming of the cytoplasm
is much mor«^ rapid than diffusion of the dye. Cuhtis (1!)2H) maintains
that streaming in phloem cells accelerates transfer of foiul substances,
and the picture here seems to support the idea that i)rotoplasmic streaming
is a more rapid means of transportation than frer diffusion.

When the pip^'tte is withdrawn, a small amount <d" prott»pIasm
flows out after it (very little if the c»dls are in isotonic, (o.OHS M) KNO;,.
solution) and the hole in the protoplast is this closed. If plasmalemma
as well as mesoplasm is included, a globule is formed which is pinched
off. If not thus protected, the mesoplasm coagulates.

Cells with dye in the vacindar sap were ])reserved in hanging drops
and observed from time to time to note whether the dyr passed out of the
cell or diffused into the cytoplasm. In no case did color appear in the
cytoplasm. If any dye penetrated, the resulting concentration, compared
with that in the vacuole, must hav(; been very h)W. Aniline hlue injected
into the sap was n'tained in every case until after streaming had ceased
and death changes in the cytoplasm set in. In oiw case, streaming con-
tinued and the color was retained for twenty hours after injection. The
same statenuMits describe all but three cases in which cells were injected
with acid fuchsin. These (after thre(» hours twenty-five minutes) had lost
color from the vacuole, though streaming continued in most of the cyto-
plasm. The cytoplasm at the tips of the cells was abnormal in appearance,
suggesting that local changes might have been responsible for the loss
of color.

15*

228

P 1 o w e

Membranes in the plant cell. II

229

We may conclude that the protoplast of the Trinaea root liaii-
is ordinarily impermeable in the active condition to aniline bhu^ and acid
I'uchsin. This is a true case of impermeability of the protoplast, not ol
unequal distribution of a diffusing substance ^vhen equilibrium is reached,
for the dye is unable to pass through the protoplast when in contact
with either internal or with external surface.

DIFFUSION OF DYE INJECTED INTO THE CYTOIM.ASM

When a small injection of either of the dyes named (less than ojk-
half the volume of the nucleus) is made in the mesoplasm, it is clearh
visible for a moment as i\ colored region, in contrast to the immediate
diffusion of a vacuolar injection. The color promptly fades. All the
cytoplasm involved continues streaming without interruption, and no
subsequent signs of injury appear. Actually, one cannot here follow tlir
diffusion of the dve into tlie cytoplasm, but sees instead the disappearance
(d- the coh)r and^he absorption of the w^ater. It might be questiontMl
whether this disappearance is due to diffusi(»n from the injection into
the cytoplasm or to reduction of the dye. To answer the question, larger
injections were made with a tw^o percent solution. From these, tlu^ color
may actually be seen to spread into the cytoplasm. Larger injections
are usually followed after an interval by cc^ssation of streaming, but th.'
cytoplasm into which the dye diffuses is, at the time, in the living, noimnl
condition. Later effects may be du(; to the action of the dye on the cyto-
plasm after diffusi(»n.

It would be difficult to reconcile these results with a (uuiceptioii
of the entire protoplnst as the osmotic ni«Mnl)r;me (d" the cell.

1N.IK(^T10N i)V HHOM CRESOL PKHPLE
(0.01 M solution, Na s,ilt)

Hrom cresol purple is yellow in the acid and blue in the MkuWur
range. Injected into the vacuol(\ it takes n\\ the yellow, acid c.»1(M-, regard-
less of the form in which injected. It shows some toxicity in the vacuole,
and causes th(^ protoplasm t(» give (df balls which are carried along by
the motion of the saj). When streaming ceases, the indicator in the vacu(de
slowly changes toward the alkaline color, presumably as a result of inward
diffusion of the external medium. The dying prot(q)lasm takes up the
dye in th(^ yellow, acid form.

?

When the dye is injected into the mesoplasm it instantly takes on
the blue, basic color, regardless of the form in which injected, and it is
in this form that it diffuses into the streaming cytoplasm. A violent,
injurious injection takes on the acid color. If a pipette is suddenly with-
drawn froni a region showing the blue coh.r, the cytoplasm changes to
v«dlow as it escapes explosively.

Since only cytoplasm which has undergruie marked injury give>;
tln^ yellow color to the dye when it is injected into the cytoplasm, and
since the yellow form is the only form in which the cytoi)lasni takes up
th.^ indicator which is injected int(» the vacuole, we have further con-
firmation (d" the inipernn-ability of the vacmdar niend)rane while in the
normal state.

PRESENCE OF KUEE ELECTROLYTES IN THE CYTnPLASM
The yell(»w c«dor of the dye in the vacurde indicates the presence
u( free hydrogen ions, and the blue cid(»r in the cytoplasm must in.licate
either that tin' cvtoplasm contributes hydroxyl ions or takes up or neu-
tralizes hvdrogen" ions. Simv the color of even a large cytoplasmie in-
jection changes immediately to the alkaline, and the coh.r (diange takes
place before the dve spreads into the cytoplasm, the sul^tance causin-
the (diange in reaction canmd be cidloidally bound, but must be free to
diffuse from the cvt«»plasm into the injected siduti.ui. llow <an we recon-
cile the presence of freely diffusible hydroxyl ions, or other ions .'apable
of cimibining with hydrogen ions, in the cytoplasm with the pnsence
of free hydrogen ions in the vacu.dar sap. unless a differentially pei-
,,^,..,1)1,. nu'inbrane intervenes? Let us assume for a moment that no
membrane is present, but that instead the non-diffnsible protein< of the
cytoplasm establish a IhnuHin niuilfhrium with elrctrolvtes; m the sur-
rounding solutituis. Then the excess of hydroiivn ions in the cell sap
over that in the cvtoplasm imiicates that the prot.-in> >npplv . olloidal
cations The cvtoplasm shoidd take up hvdrogen ion< onlv from s.dutions
more acid than the vacuolar sap. On the c.mtrarv. the indicator solution
is less acid than the sap. remh-ring such an explanation ot the .liange
in reaction inade(,uate. Similarly, if we assume that ih- ehan-e in re-
nction of the indicator is due to adsorption .d' the hydrogen ions rather
than to the diffusion into it of some substance ivactin- with hydrogen
ions, it is difficult to see why the cytoplasm, alreadv in mntart with the
acid vacmdar sap. should take up hydrogen ions from the <liglitly a.id
indicator solution.

280

P 1 o \v e

Membranes in the plant cell. J I

231

THE KXISTKXCP] OF DIFFERENTIALLY PERMEABLE PLASMA

MEMBRANES

]Jiff(T«'iitial permeability of the vacu()lar nienibrane seems the
most reasonable (explanation available of the difference in the behavior
of the cytoplasm towards the vacuolar sap and towards solutions injected
. directly into it. The membrane appears to be differentially permeable
both to the injected dye and to the electrolytes normally pi'escnt in the
cytoplasm.

ANOMALOUS BEHAVIOR OF CERTAIN INJECTIONS

Possible causes of anomalous bcehavior of cytoplasmic injections
occasionally seen (Kite, 1913a, and work on Griffithsia, this paper)
can be determined, thanks to the clearness with which details of injections
may be followed in the root hair.

It is worthy of note that it is almost invariably larae injections
which fail to diffuse into the cytoplasm. This is just the reverse of what
one would expect if the permeai)ility of the cytoplasm to injected dyrs
were a result of injury. A larjj;e injection will occasionally be sharply
delimited from the cytoplasm, the injected drop will tend t(» beccnnc
spherical, then suddenly the color will jxrow more intense, without an
accompanying* decrease in volume, which would explain the chan^<' iu
intensitv hv concentration of the solution. Careful focusiu«r shows that
the dye has been adsorbed by the (presumably) coaii'ulated surface hiyei*
at the dye-cytoplasm interface. A second inj(H'tion will not pass throufrh
such a coaji-ulated film until it is broken, indicating- that a coa^rulated
region may be an effective barrier to diffusion. ('uamhF'.hs { 1917)sugirested
formation of a "coagulation film"" in explanation of Krrifs (1913a)
account of "vacuohN" from which the dye does not diffuse as a result
of injection in animal cells. The film just described seems to tally with
Cjiamheus' suggesti(»n and also with KrrK's statement that in some
cases the dye is taken up by the. wall (d' the vacuole.

Occasionally one small mass of injured cytoplasm will take up all
the dye from the siu'r(Uinding region, giving a small, int(Misely colored
spot. Such a dens(dy stained coagidum may be what Kite saw (1913a)
W'hen he stated that the water may diffuse cmt from the vacuole formed
by injection (starfish eggs) leaving the "precipitated" dye behind.

EPIDERMAL CELLS OF ONION

The (Uiion material described in the earlier pari (d' the work was
used to supplement the work on Trianea. It is more difficult to secui'e a
nu'soplasmic injection in these cells, for the layer of protoplasm lining the
walls is thin, and the details of an injection cannot be followed so easily,
but the active streaming makes it possible to check, at all times, on the
<'(»ndition of the injected material.

Tin" metlKKl used was varied according to the h)cation of injection
<lesired. Vacm^lar injections can easily be secured with the cells mounted
in tap water or in an isotonic sugar solution. In plasnudyzing scdution
the pipette invaguiates rather than pierces the ])rotoplast, and the ejected
fluid enters neither cytoplasm nor vacuole. In isotonic solutions, or
sidutions slightly moiv or less conc<^ntrated, it is possible to pierce th.'
wall and plasmalemma without puncturing the lonoplast, ana inject
into the mesoplasm, with a fair degree of regularity. The greater part
<d the work was done with the c -lis in an «dght perc(>nt siduti-ui of sucn»se.

KMLriO^: OF DYES TO PENETRATE THE PROTOPLAST ON
IMMERSION AND YxUTJOLAR INJECTION

In additi(Ui to the d,yes used in the work on Triancd, phenol n^d,
i\ hydrogen ion indicat(»r whiidi covers a more alkaline range than that
<d" brom cresol purple, was ♦Mn])l(»y(ML In onhT that any effect of sucrose
H.n permeability might affect all methods of application of the dye ahke,
the s(duti«»ns for immersion were maih- up in eight percent sucrose, con-
taining approximately one percent of aniline blue and of acid findisin,
and 0.005 molar concentration of the indicators. Neither aniline blu.;
nor acid fuchsni visibly penetrated living cells after 28 hours immersion.
Neither of the dyes appeared to be toxic. Twenty-four hours in brom cresol
purple failed to bring about penetration td" living cells. Pheiud red occa-
sionally penetrates alxuit one cell out of one hundred, but pndonged
immersion produces no increase in the numbr-r pem^trated, and twenty-
eight hours of immersion may produce no penetration whatever.

Acid fuchsin, aniline blue, and phenol red in no cast- escape fnui.
the vacuole when injected before vi-xible abnormalities appear in the
cytoplasm, and generallv are retained for a while after the death (d the
<'eil. Seven hourb was the longest observation for acid fuchsin, four hours
.ind fortv-five minutes for aniline blue, and ninety-six hours {or phenol

232

V I o w ('

red. Tlie jJTontcr luinibci' of iiij(M't<'(l crlls wei'c only wntclicd for .ui lioiir,
Hiiic(3 it would srL'iii tliat retention of the dye in the v.iciiole for .111 hour
spells relative imperirKiability of the toiioplast wlicii contrasted with iin-
niediate diffusion on inj('eti(ni into the rncsoplasin.

The toxicity <>f hroni cicsol i)nrpl<' prcv<'nts th<' use of this method
of testing' the pci'nicahihty of tiic vacn(dar inrnd)ran('. No visihh' pene-
tration (d' the cytoplasm can be seen, however, until the dyin<z- prntnpl.ism
takes up the yellow, acid form (d' the dye.

The results oi immersion and vacuolai* injection indicate that the
surface (d' the pi'otoplast is impei'meable to the foui* dyes whethei' they
are bi'ou^iit into contact with the external or with the internal surface.

DIFFUSION OF MESon.ASMIC 1NJF(TI()NS

Acid fu(disin, injecte(l into the mesoi)lasm, diffuses almost imme-
diately, without interruption (d' streannnji-. The c(dor can Ix' seen to
spread in the peripheral cytoplasnnc layer. The coloj* leaves the injection
more rapidly than the water is taken up by the cytoplasm. The injection
usually forms a ''blist<'r " in the thin cytoplasmic layei* whi(h i)i'ojects
hito the vacuole, and this protrusion may persist foi* a lidnute or two
after the coloi* has left it. The outline id' the injection fivuitiii"!' on thr
mesc>plasm is irreji'ular, and becomes mure so,, is the water is tak<'ii up.
but the appearance presented is (hd'init<'ly indicative that the water is
taken up by the cytophism; the cytoplasm does not mix with the wat<'r.
Sliji'ht, tem])(U'ary sw(dlinjr of the cytoplasm near the injection may
follow the absorption (d' the water.

Did the entire thickness (d* the cytoplasm constitute the osmotic
membrane (d' the c(dl, it would be wholly out (d order for diffusion (d'
dye to thus precede diffusiim of s(dvent. wdien scdvent ahme [M'lietrates
from the external milieu.

While the thinness (d" the cytoplasmic layer increases the difficulty
of injection, it also serves to (h'lnonstrate more forcibly than in the c;is«'
of Trianea that the mesoplasm is m(»re readily permeable than the
tonoplast. In the case of Trianea, most injections wei-e ma(h' into a mass
of cytoplasm throu^'h which the dye could spread in all directions. Here.
even when the tip of the pipette projects into the vacuole, covered only
by a thin film, the injected dye spreads back throuj^h this layer into the
layer of cytoplasm ajrainst tin; wall. Were the permeability of the tono-
plast as ji;"reat as that of the mesoplasm. this would mean that the dy«'
w%'is diffusiuii" alon^ the path (d jrreatest resistance.

.Membranes in the plant tell. 11

'>X\

:

When th(> medium is sliii'htly hypertonic, so that the pipette fails
to pierce the flaccid prot«)plast. the fluid ejected flows back al«m;r the
pipette into the external solution, no matter how violently it is «'jected.
If injury to the cytoplasm were the cause c.f the permeability shown
bv diffusion (d" mest>plasmic injections, one would expect sindi a violmt
injection to diffuse into the cytoplasm rather than the numH mesoplasmie
injection in whicdi no suidi force is emi)loyed.

Aniline blue diffuses in the same manner as arid furhsin. I»iii is
markedly toxic. ('oa«i-ulatioii of the injected reLdon results, and fliis
rej^ion will not resume stre.indim". thou;.ih noj-mal niotinn may contiiiiie
in the rest td" the protoplast for some time.

rilK DIFFKUFNTIAL PKH.MKAHILITY oF IM.ASMA .MKMUli.WKS
INDKWTFD \\\ INdKl^TlONS ()F INDK'ATnU DYKS

(().()] M solution. Na salt )

IMiemd red in (i.oi M s(dution has lowei- cohuinii- |M.wer than ilif
other dyes used, hence the injection fades more lapidly as the dye diffii'-es
than does one (d" acid fucdisin. and the spread .»r the dy.- in the e\toj»la-m
(*annot be ffdlowed well in small injections. Streamin^r may be Ineallv
interrupted, but is soon resume<l. The dye takes tui the yellow, acid (mImi-
in the vacuole, and the rose colored alkaline form when inject. d iiiio
the m»'S(>plasm. and, as in the case of brom civsol purple and Trinnni,
the color (dianire is immediate, and takes i)lace befoiv the dye spread^ in
the cyt(»plasm. In n<» case in whi(di local streamiiiii- was resumed di<l
phenol red diffuse into the cytoplasm in the yellow form: (dianiivto y.Hnw
was an imlication <d' injury prece<liiio- cessation <»f streamin^^ In nne ca^.-
the dye spread in the orange, intermediat*' fofiii into cytoplasm which
showed irre«rular streaminji: but swellin^i' followed and ^tieauMiiii' ^'hhi
ceased. Since phemd red covers a more alkaline raiiiie than br..m civ-..|
[)urple. it will indicate sliiihter ( han^cs in reaction on injury.

Krom cresol purple is lemon yell(>w when injecte.l into tin- \aciiM|e.
while a mesoplasmic injection, even when blisteriim- mit into thr vacii.»lr.
immediately takes (Ui the blue color. When injected into tie- mesoplasm
near the mndeus. that body takes up the dyr in the alkaliie' form. When
the dye is injected into the vacuole, on the other hand, the nucleus remains
uncolored until the dyinir cytoplasm ami nindeus both take up the yellow
color. The nucleus may appaivntly take up the dye from a mesoplasmic
injectiiUi without injury, for <me cell in whicdi the niiclens was en|n|vd
remaine(l mn'inal for an liour after the injection.

234

Plo wo

Attempts to conti'ast tin.' effects of vacuolar and mesoplasmic in-
jection further ai*e not (juite safe, for the conc(»ntration of the diffnsin<>:
dye will, of course, vary in the two cases, but it is of interest that brom
cresol purple unfailinirly ])roduces different cytoplasmic pictures when
injected into the vacuole and into the mesoplasm. When it is injected
into the vacuole, the mesoplasm of the transvacuolai" strands rounds
into balls on the tonoplastic portion. Usually Brownian movenn^nt of
the cytoplasmic jj^ranules also results. Vacuolar injections are fatal unless
th(^ puncture of the i)rotoplast at the entry p(nnt of th(^ pipette enlarges,
as described above foi' puncture by a needle, and the dye is allowed to
(escape from the vacuole, when the protoplast will survive. When the
dye is inj<'cted int() the cytoplasm, an appearance su^jrestinfr alveolation
of the mesoplasm, a lace-like pattern, is produced in a larjre part of the
cell, and. over this area, all movement of jrraiiules is temporaiily sus-
pended. Movement is I'esnmed, the 'alveoli'* elongate and become
irregular in form, and the granules enter into streaming motion. This
re(X)very b(;gins at the region farthest frcun tlu' center of the injection
and pi'oceeds centripetally until all jxM'tions of the injected region show
normal streaming. There is no balling of strands and no Brownian move-
dment. 80 consistently do these two i)ictures result that it is difficult to
avoid the suspicion that the first represents an acticm of bnmi cresol
purple on the tonoplast alone and the second on the mesoplasm.

Th(3 coloring power of brom cresol purple in the basic form is si)
great that the spread of the dye through the cytoplasm can be traced
for half the length of the protoplast or more. The alveolation described
usually proceeds slightly farther than the visible diffusion of dye. The
question arises, is the alveolation a specific effect of the diffusing dye,
01' is it indicative of injui'v which renders th(^ cytoplasm permeable?
That the first is the case is indicated by tin* fact that phenol red, acid
fuchsin, aniline blue, eosin, indigo carmine, and isotonic sugar solution
can all be injected into and taken up by the mesoplasm without producing
this picture. Injection of brom phenol blue gives a similar picture.

The immediate change in reaction of phenol red and brom cres(d
purple when injected into the mesoplasm, the fact that brom cresol purple
can be seen to diffuse half the length of the cell from a small injection,
through a thin peripheral layer of cytoplasm, are alike inexplicable unless
we pictun* the mesoplasm as separated from the vacuole by a structure
which pn^vents the passage of substanci^s to which the mesoplasm is
freely permeable.

Membranes in the plant cell. II

235

(IKIFFITIISIA

(iriffUhsia hornetiana is composed of branching chains (d cocnucytic
cells which reach a maximum length of 3.2 mm. and breadth of 0.fi5 mm.
The wall, although thick, is soft and is easily ])ierced with the micr(>-
pij)ette. The thin lining layer of pr(>loplasm forms thick pads against
th(^ cross walls. Hy inserting a pipette parallel to the cmss wall, it is
fairlv easv to make a injection into this mass of cvtoplasm. Th«' ryto-
])lasm c<>ntains numerous nuclei, plastids. some starch, and small protein
granules. In the young cell the plastids are cl(>sely packr'd, and polygonal
in outline, giving a mosaic pattern. In the older cells the plastids are
4'longated, oval oi* ii'regular in outline, and occur in rows, forming a net-
work. When the cytoplasm is injured, the plastids round up so rai)idly
that one can follow the change in form, the nuclei swell and become
dearly visible, and the protein granules enter into Brownian m(»vement.
The change in the plastids and the movement of the granules are the
most immediate and striking effects of injury, and may precede loss (d
<'olor by the plastids by an hour or more, or may occur as reversible injury
4'ffects. These changes were used as an index of injury to the cytoplasm.

FAlLrHK OF DYKS TO PENKTKATK ON IMMKK'SloN

Ib'anches of GriffftJtsia were immersed in a saturated solution of
e(>sin in sea water, in 1 gm. of aniline blue to KM) cc. (d sea water, and
in a oile perc(*nt solution of methylene blue in sea water, and examined
foi' penetration of the dye after removal to uncolored sea water. Th»'
<'ells were torn oj)en with a microneedle to detect color in the vacuoKir
.sap. Methylene blue gave a faint color to the sa]) after four hours immer-
sion. Eight hours in the other dyes gav«' no color to sap oi* cyt()plasm
of livinjr cells. Methvlene blue was used, as well as eosin and anilin*'
blue, to see if results (d' cytoplasmic injection showed differences corres-
ponding to results (d immersion.

DIFFFSION OF DYES INJE(TEl) INTO THE CYToPLAS.M

The soluti(»ns (d" aniline and methylene l)lue in se;i water were used
foi' cyt(>plasnuc injection. The sea water solution of eosin is too faintly
colored t(» be useful. One percent solutions of all Wwrr dyes in distilled
water were also used.

286

P 1 <i w o

Matci'i.il w.Ms kept in h.niji'inji' drops of sea \vat<'i', al'tri' injection
witli cacli of tli(' fiv(^ solntiinis, to (l(!t(*i*nii!i(' Avlicthcr injnry wliicli was
not (let(M'tahl(^ at the time wonld make itself known later. One cell, into
the cytoplasm of which aniline bhio in distilled water had Ixmmi in.)<'Cte(l.
was in ^'ood condition over forty-live honi's after the inject ioiL The
lonj2;(?st (d)servations for cells injected with other dyes were: eosin. o\er
forty-one honi's; aniline bine in sea water, three honi's tw(^nty-five minutes;
methylene blue in distilled watei*, foni' houis and for'ty-five nnnutes;
methyhMie bine in sea watei', seventeen hours thirty nnnutes. The
injecti'd cells fiMMpieidly outlived their uninjcH'ted neij^'libors. Injuiy
eff(H*ts, unless I'eadily I'ecoo-ni/able within an hour, faileil to appeal*
alto«r<'thei'.

Small injections, from one to four times the v(»lume <d' a nucleus.
ai"e usually wholly absorbed within two minutes. The time required foi'
absorption does not chan«re perceptibly when the dye is dissolve(l in sea
water. The visible color usually j)asses out of the injected droplet l)efor«'
all the water is taken up. The injected fluid usually shows a shai'p mai'iin
only where it fronts on the va('U(de. hijections diffuse without swellini*'
oi cytoplasm, rounding' of ])lastids, Hrownian movement of pi'otcin "rra-
nules, or staininjj; of (hither nuclei or plastids. Slightly lar«rer injections
may caus(^ staininji; of one or two nuclei (aniline blue) or plastids (methylene
blue, eosin). Methylene blue shows no perceptible difference from t hr
two acid dyes in behavioi- on cytoplasnuc inject i<Mi.

In th(^ case of injections so lar^e as to be injurious, the visible injury
chan^-es in the cytoplasm usually follow, rather than pnu'ede, the diffusion
of tlu; dye. Occasionally a '"vacuole" with a. coaji;ulated, stained, rela-
tively impiu'meable wall is produced, as described for Trinvon. It is
to be note<l that it is only from the larjre inj<M'ti(ms that tiu^ (lye fads
to (liffuse into the cytoplasm; injury inhibits, rather than facilitates,
<iiffusion.

The plastids make it diffi(*ult to deterndm^ with finality whether
dyes inject(Ml into the vjicuole ndfiiit be taken up by the cytoplasm.
Kosiu is occasionally taken u^), but always as the cytoplasm be^nns to
show si^'us (d' injury.

The details of injection are less easily followed than in Itianocn
and onion c«dls, but it is of interest that all results obtained confirm,
for this n^presentative of a wliolly different <j:roup of plants, thos(i des-
cribed for the cells above.

fS

N>

M cm bra lies in the |»lai»t ct'll. II

I'ONCLI'SIONS

•2'X

I. IIh' ((iffnrHlidl iHrnHdhiltl n nf ihr pntinplasl huranls nnin umi
ihfr.s is (I pntftntfi o/ ihr stirfdrc hnfcrs nilhrr iIhih ihr wrsophism.

In order t«» intei'pret results obtained in any other mamn'r than this.
it w(udd be necessary to assume, first, that mechaidcal injury produces
abnormal jM-rmeability in the mesoplasm Imt does m.t affect the per-
meability of the two membranes, and srcuiid. that the meso|»lasm can
underp. injury which radically altera its permeabditv without showini!
any siiiii (»f injury in its appearance or behavior.

'J. Ihr xin-jdcc Unjcis (tic also (liffcrrnliull H pnntnihh liHrnnls fm'li,
(liffusthlr rlrclnti flics nnrmnlln ftrcscnl in cell sap, ciflnphism. mul cilcnuil
snhllKm.

T\m is shown by instantaneous c«»lor chaniivs ..f injrcfd hydro^n-n
i..n indicator^ in mesoplasm and vaeiioh'. which canmU be explained a-
residts of .idsorption nr .1 hontKui c(iiuhhtnini.

3. 77/r presence 0/ (liffcrcnlidlhi pcrntcahlc nicinhnmcs al the surfaces
nf Ihe rcndthi permeable mesoplasm makes 0/ ihc racuiflalc plant cell a annplex
sffstem. Ihe hehariin nf ahlch rests nn nsmata as ncll as rnlli,alal phmnmenn.

The mesoplasm evich'iitlv retards diffusi(»n. rvm nf water, tn soiic
v\\n\\. The permeability .»f the jMotoplast i^ iidlueiicrd by the per-
meability of all layers, thou-h in m(»st casesthr p**>'»ii«'.ihilitvnf the.xternal
;ind internal membranes i> the limitinii' factor.

The marked effects of osmotically inactivr cniiceiitrations ol acid
and alkali <.n cell volume result from action mi the hydrnphilie prntein^
within the membranes. The interventinn of the plasma m.-mbrane mak«-
the ])enetratinu- power of the acid a cniitributinii- fact(»r in its ♦■fb-etiveness.

4. 117^^// is the relaliiai hehceen ihe nnaphitUnncal memhranes. ihc Innn-
jdasl and plasntalennna. and Ihe nsm(dic memhrancsf

\{v^\\s^vv<^\ (1!>2H) measuriu'i- the capacity of tin- plasma membran.'
in the beet foot. calculate> its thickness as |(M> A (0.(H //). If. however,
we alter his calctdation by assumiufj;, as the work abnvr would indicate,
that each cell contributes tan mend)ranes. the vacunlMr as well as tlh-
external plasma membrane, the calculated thickness will be o.(M>r,//.
<.r if we assume that the dielectric constant ..f the mend>rane is <rreater
than teti. the calcidated thickness increases in pr(»portion. Cnnse.piently
it is imiM.ssible to (h'ci(b' fnmi this whether the entire thiekie-ss of totrnplast
.,nd i)lasmalemma. or .Mdy a surface layer, funetions as an nsmotic mem-

238

P 1 o w e

Menibranra in tlu' j)lant cell. II

2:^J»

bmne. SiiKT such <i layc^r, if present, is inseparable from the iiiorphoh^-
jj;ical membrane, which j,nves rise to it, is ruptured when that membrane
is ruptured, destroyed when it is destroyed, isolated when it is isolated,
it may be permissible in any case to speak of the "differentially permeable
tonoplast and plasmalemma" just as we speak of a "waterproof epi-
dermis^' wiien only the cutinized outer wall is really waterproof.

The plasmalemma and tonoplast are physioloffical, as trell as mor-
phological, entities.

AOKNOWLEDOMENTS

The work upon which this paper is based was carried out with th<'
cooperation of Dr. William Seifriz of the University of Pennsylvania.
1 wish to make jj^i'ateful acknowledgment of the help which he has given.
I also wish to thank Dr. M. H. Jacobs for the use of a table at Woods
Hole while w^orking on the part of the paper dealing with peruieability.
The 0.01 M. solutions of the indicator dyes were supplied through the
kindness of Dr. Mary Kilpatrk^k.

SUMMARY

1. Direct evidence foi* the existence of distinct osmotic membranes in
the living plant pi'otoplast has not hitherto been presented. The
presence of a vacuole interferes with efforts to answer th<' ((uestion
by indirect methods.

2. Epidermal cells of onion, root hairs of Trianea, and the coenocvtic
cells of the alga Griffithsia were used as material for microinjection.

3. It was found that acid dyes which do not penetrate the protoplast
immersed in them during its life likewise fail to penetrate the proto-
plast when injected into the vacuole. On the other hand, when in-
jected into the mesoplasm itself, they W(U^e able to spread in the
protoplast but unable to pass through the tonoplast into th<' vacuole
or through the plasmalemma into the external solution.

4. This evidence for the existence of osmotic membranes at cytoplasm-
wall and cytoplasm-sap interfaces is given further support by color
changes of injected indicator dyes, explanation of which is possible
only if a membrane differentially permeable to electrolytes occurs
at each of these interfaces.

5. It seems reasonably certain that local injury to the cytoplasm did
not enter into the results obtained.

Department of Botany, University of Pennsylvania, Philadelphia, Pa.

•*.

n^

\ATK\<AT\nK CVVVA)

Hakucvzen, H. L. v., 1928. The fuiu-tion of plasma coUoid.s in m'uwth and pt'rnu'ability.

Paper presented at New York meetings of A. A. A. A., Decenilx r, 1928.
Beutner, R., 1913. Kinige weitere Wrsuehe betreffend osniotisclie luid knlloidalf

QuelluniJj des Miiskels. Biocheni. Zeitsehr. 4S. 217 224.
('iiAMBKRS, R., 1917. MierodisseH'tion studies. 1. The visible structure of cell j)rnt<»plasin

and death changes, .\nier. .lourn. Physiol. 4.*J, I 12.

— 1922. A microinjection study on the j)ermeability of the starfish egg. .lourn. (len.

Physiol. r>, 189-194.

— 1924. The physical structure of protoplasm as determined by microdissecticm and

injection. Cowdry, Geufnt/ Cytology, I'niv. of Chicago Press.

— 1928. Intracellular hydricm concentration studies. I. The relation of the environm<'nt

to the pH of protoplasm and of its inclusion bodies, liiol. Bull. .Marine Biol. Lab.

oo, 369 — 370.

— & H. Pollack, 1927. .Micrurgical studies in cell physiology. 1\'. Colorinu'trii- deter-

minati(m of the nuclear and cytoplasmic pH in the starfish egg. .lourn. <;en.

Physiol. 10, 739—755.
& S. HiLLER, 1927. The protoplasmic pH of living cells. Proc. Soc. Kxp. Biol. an<l

Med. 24, 700 761.
CuRTLS, O. F., 1928. Studies indicating that translocation is dejH'ntlent on the activity

of living cells and the possible bearing of this on solute distribution. .Vmcr. .lourn.

Bot. 15 (supp.), 630—631.
Dangeard, p., 1927. Sur Torigine des vacuoles. I^' Botaniste IS, 245 2(>4.
Fischer, M. H., 1908. Cber die .Analogic zwischen der \Vas.serabsorption durch Fibrin

und durch Muskel. Pfliigers Arch. ( Jes. Physiol. 1*24, 69 79.

— & (i. Moore, 1907. On the swelling of fibrin. Amec .lourn. Physiol. 2i>. .330 342.
(iiCKLHORN, .r., & F. Weber, 1927. Cber \'akuolenk(mtraktion und Plasmo|ysef».rm.

Protoplasma 1, 427 — 432.
HoBER, R., 1910. Fine Methode, die elektrische Ltntfahigkeit im Innern von Zellen zu

messen. Pfliigers Arch. (ies. Physiol. 133, 237 — 253.
Rowland, R. B., 1928. The pH of gastric vacuoles. Protoplasma 5, 127 1.34.
Jacobs, M. H., 1920. The pmduction of intracellular acidity by neutral and alkaline

solutions containing COg. Amer. .Journ. Physiol. r>3. 457 463.

— 1922. The influence of ammonium salts on cell reaction, .lourn. (ien. Physiol. 5,

181—188.
Kite, G. L., 1913a. The relative permeability of the surface and interior portions (»f the
cytoplasm of animal and plant cells. Biol. Bull. .Marine Lab. 25, 1-7.

— 1913b. Studies on the physical properties of protoplasm. I. The physical properties

of the protoplasm of certain animal and plant cells. Amer. .Journ. Physiol. 32,
146—164.

— 1915. Studies on the permeability of the internal cytoplasm of animal and plant

cells. Amer. .Journ. Physiol. 37, 282—299.
KiJSTER, E., 1926. Beitrage zur Kenntnis der Plasmolysc Protoplasma 1. 73—104.
Lloyd, D. J., 1916. The relation of excised muscle to acids, salts and bases. Proc. Roy.

Soc. (London) B S», 277—290.

c^^Q Plowe, Membranes in the ])lant cell. H

Llcyd, F.K., 1915. Behavior of protoplanni as a colloidal complex. Yrbk. Carnegie

Inst. Wash. J4, 66—69. . „ ^ v

- 1917. The effects of aeids and alkalies on the growth of protoplasm m pollen tubes.

Mem. Torrev Bot. Club 17, 84— 89.
LoFB J 1922. Pioteins and the theory of colloidal behavior. Mc(.raw Hill., New York.
Moore, B., & H. E. Roaf, 1908. On the equilibrium between the cell and its environment

in regard to soluble constituents. Biochem. .lourn. 8, 55 -81.
_ & A Webster, 1911. Direct measurements of the osmotic pressure of casein in
alkaline solution, experimental proof that apparent impermeability of a ";;';"l>ran-
to ions is not due to the properties of the membrane. Biochem Journ (>, 1 1(>--1 2 1
Mc(5LE.noN, J. F., 1926. Colloidal properties of the living cell "• ^^-^^if -;*^^ ^^^

capacitv of blood to alternating currents. Journ. Biol. (hem. ♦>.», 73,3-754
Xeeoham, ,).; & 1). M. Neeoham, 1926a. The H-i<m concentration and oxida^on-
reduction potential of the cell interior before and after cleavage. Proc. Uoy .
Soc. (London) B. 99, 173— 199.
- 1926b. Further microinjection studies on the (.xidation-reduct.on potential of the

cell interior. Proc. Roy. Soc. (London, B. 99, 383—397.
OSTERHOITT, W. .1. V., 1913a. Protoplasmic contractions resembling plasmolysis which

are c'aused bv distilled water. Bot. (Jaz. 55, 446—451.
_ 1913 b. The organization of the cell with respect to permeability. Science n. s. rfS,

408 — 409. . . , , .,.,

_- 1922. Injury, recovery, and death in relation to conductivity and permeabiht> .

,1. B. Lippincott, Philadelphia. r. * «

Peeefer, W., 1897. The. physiology of plants (Tr. A. J. Kwart). Clarendon Press Oxfonl
Ploxve, .1. Q., 1931. Membranes in the plant cell. 1. Morphological membranes at

nrotoplasmic surfaces. Protoplasma.
llAPKivE L and R. Wirmskr, 1926a. Sur le potentiel de reduction du noyau et les

oxVdations cellulaires. Compt. Rend. Soc. Biol. 94, ^^^^^^^^^
- iq26b Sur le potentiel de reducticm des cellules. Compt. Rend. Soc. Bio . 9o, 604-60.
REMiNOTON, R. K., 1928. The high frequency Wheatstone bridge as a too in cyU,logieal

studies; with some observations on the resistance and capacity of the cells of the

beet root. Protoplasma 5, 338 399.
SFIFR17 W 1927. New material for microdissection. Protoplasma *J, 191-196.
SrEBECK. rI 1912. Uber die „osmotische Eigenschaft^n- der Nieren. PfUigers Arch.

Ues. Physiol. 14H, 443—521.
HE Vries, H.,* 1885. Plasmolytische Studien iiber die Wand der \ akuolen. .lahrb.

Wiss. Bot. 16, 465 598.

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■« F

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\

THE SPIERP:R lens AND WHAT IT REVEALS
IN CELLULOSE AND PROTOPLASM

BY WILLIAM SEIFRIZ

Fig. I

The ellipse of scattered light from a
colloidal particle.

The Spierer Lens

Charles Spierer, Swiss physicist, has devised a lens for ultramieroscopic
observation^ which involves the principle that light scattered by colloidal
matter is greatest in the direction of the illuminating ray (Fig. i). It would
naturally be of advantage to view matter against a dark field yet toward the
source of illumination. It was Spierer's task to accompHsh this. He did it
by placing a small mirror in the lens system of an oil-immersion objective
(Fig. 2). The illuminating rays come directly from below as in an ordinary

microscope; consequently, that part of
the scattered hght which is of greatest in-
tensity is toward the observer. The small
mirror in the objective gives the dark field.
The principle involves two primarj-
prerequisites; first, the mirror must re-
flect all of the direct light, second; the
mirror must be smaller than the lens aperture, in order to leave room for the
scattered rays from the object to enter. The first prerequisite is accomplished
by having the aperture of the substage diaphragm at least as small as the
mirror; in practice this is 1.5 mm.

The mirror may be of gold, silver, platinum, or aluminum. In the ex-
perimental stage, and later when Spierer lenses were made by Nachet of Paris,
the mirror was a piece of gold or aluminum foil placed between the lenses of
the objective. The method now employed, by Zeiss, is that of electric deposi-
tion. The lens is placed behind the opening of a screen situated near the
anode of a cathode tube at the opposite end of which is a cathode of platinum;
the opening in the screen is of the size the mirror is to Yk\ Platinum cathode
rays are given off and deposited on the exposed surface of the lens.

Aluminum is fully opaque and gives a perfectly black field. (Jold or
platinum have some transparency and permit a little of the direct light to
pass which is an advantage at times. The gold mirror gives a dark-green
field and the platinum mirror a gray field.

The Spierer lens is an oil-immersion objective of 1.25 aperture (112 inch =
2.12 mm). It is provided with an adjustable iris which is useful in that it
allows the observer to keep oflf part of the diffracted light when the diffraction
phenomena are too intense and give a blurred image. (This intensity depends
upon the nature of the object and of the embedding medium.) It is necessary
to close this iris partially when using the latteral illumination of a cardioid
condenser, otherwise the field is not sufficientlv dark.

» Arch. sci. phys. nat., 8, 121 (1926).

iig

THE SPIERER LENS

WILLIAM SEIFRIZ

I20

It is sometimes of advantage to use a cardioid (dark-field) condenser in
conjunction with the Spierer lens; this is not, however, necessary since the
Spierer lens is itself a ''dark-field," and gives very satisfactory results with
an Abbe condenser, or none at all. The advantage of a cardioid condenser,
when used in addition to the Spierer lens, lies in the fact that it increases the

v///////////////: d

Fig. 2

The Spierer lens and special (Zeiss- Spierer) cardioid condenser: o = microscope ob-
jective; I = lower lens of the oil-immersion system; m = platinum (Spierer) mirror;
r.r. = reflected rays from the (Spierer) mirror, e = cover-slip, / = colloidal material;
g = slide; c = cardioid condenser; d = special fixed diagraphm; a = 1.5 mm aperture for
direct light, d.r., to Spierer lens; 6 = slit for cardioid rays, c.r.

illumination of the colloidal structure by adding its share of scattered rays,
which help though they do not come from the region of maximum dispersion
as do those produced by the Spierer lens.

Spierer has combined in a special condenser the parts necessarj' for using
the cardioid with his lens (Fig. 2). The condenser c contains the usual car-

■'f

^ %

i

V ' ♦

<-t

dioid elements and in addition a fixed diaphragm d with a permanent aperture
a of the desired size. This arrangement also gives perfect alignment of the
illuminating ray, an important condition for the Spierer lens. The condenser
possesses a movable disk which obstructs or leaves open the aperture. W ith
the disk in the center, i.e., the aperture closed, no direct rays pass through the
condenser to the objective; the cardioid alone is then in use and the Spierer
lens functions like any oil-immersion objective. With the disk aside, direct
rays pass to both the cardioid system and the Spierer lens.

To use the Spierer lens alone, the iris diaphragm, which is part of the
microscope substage and is situated below the condenser, must be closed
enough to allow no light to strike the reflecting surface of the cardiod system;
the small aperture in the fixed diaphragm of the condenser remains open to
permit the vertical illumination to reach the Spierer lens. A simpler way,
however, to use the Spierer lens alone as a dark-field lens, is to replace the
special condenser by an ordinary^ Abbe one.

The soundness of the theory on which the Spierer lens is built is prov(»n
by the results obtained. Spierer' has observed a fine granular structure in
dry collodion which is not revealed by any other type of optical system. I^he
cellulose and protoplasmic structure to be told of here is also not to be seen
except with the Spierer lens.

There can be no doubt that this new optical system of Charles Spierer will
bring a finer structure to view than we have yet been able to see in many
materials.

There are substances in which no more, in some instances less, can \ye
seen with the Spierer lens than with the ordinary- dark-field condenser. This
is likely to be true where the index of refraction of the material investigated
is suflficiently different from that of the surrounding medium to give pro-
nounced optical contrast; where the surrounding medium is of nearly the
same index of refraction as is the object viewed, then the Spierer lens reveals
contrast and structure not visible by any other lens or system of illumination.
For example, Pleurosigma angulatum has an index of refraction of 1.6 (that
of sihca); when this diatom is mounted in water (index = i) its structure is
fully visible, but when it is mounted in cedar oil, the index of refraction of
which is 1.5, nothing is to be seen with ordinary illumination except the
periphery. Lateral illumination (cardioid condenser) brings out considerable
structure but leaves part of the diatom blank and the rest unclear; the Spierer
lens reveals a much clearer structure and shows it throughout the diatom.
The index of refraction of water is very close in value to that of cellulose and
protoplasm; the Spierer lens should, therefore, show more of the structure of
these substances than do other optical systems.

The success of work with the Spierer lens is to a great degree dependent
on the nature of the medium in which the material to be studied is mounted.
It is well to try air, water, glycerin, balsam, and other media before giving
up, though often one is as good as another.

IRREGULAR PAGINATION

121

THE SPIERER LENS

There is another important difference between the Spierer lens and the
usual cardioid type of ultramicroscope ; this is the distinction between what is
light and what is dark in the two cases. If we view a bacterium with a cardioid
condenser, only the edges of the minute object are bright; the interior, the
bacterium itself, is dark. The same object viewed with the Spierer lens re-
verses things, the edges are dark and the bacterium light(since the material
is transparent); in other words, the Spierer lens gives, in a sense, a ''Roent-
genogram" of the interior^ while the cardioid only illuminates the object,
giving a ''photogram" of the whole.

Experimental
The observations to be reported here are restricted to the cellulose walls
of the dead pith cells of Samhucus (elder), the dead and living epidermal leaf-
cells of Allium (onion), the stalk of Apium (celery), and the protoplasm of

f-?-

Fig. 3
Cellulose wall of dead elder pith cells showing end-to-end orientation of linear micelles
in parallel striae, surface view.

the living cells of Allium. Some of the results were obtained in cooperation
with Mr. Spierer in his laboratory in Geneva. I am greatly indebted to him
for his technical assistance and continued interest in my work which involves

the use of his lens.

Since writing the first draft of the manuscript of this article I have learned
from Mr. Spierer that he too has continued our first investigations on the cellu-
lose walls of plant cells, and obtains confirming results on such other material
as the leaves of Cichorium and Plantago. With Mr. Spierer's kind permission I
shall incorporate some of his findings in with our original and my recent ones.

Cellulose. The ultramicroscopic^ structure of the cellulose walls of both
dead and living plant cells is characterized by Unear arrangement and dis-
continuity (Figs. 3 and 6). The linear arrangement is of parallel striae; the
discontinuity is due to the presence of small rod-shaped particles oriented
end-to-end (Figs. 3, 5, 6, and 9). The striae usually run parallel to each

2 In this article I shall use the word ultramicroscopic simply to mean what is seen with
the Spierer lens or other dark-field system without reference to size of structural units.

WILLIAM SEIFRIZ

122

Fig.

Cellulose (parallel lines, upper right), protoplasm (broken lines, lower left), and nucleus
(spotted circle) of living onion cell (nucleus is 20/i across).

other with only occasional irregularities. The general arrangement of the
striae is often strikingly symmetrical (Figs. 7 and 8). At times, the striae
run at right angles to or even cut diagonally across each other.

There is little difference to be seen in the ultramicroscopic structure of
the epidermal walls of elder, onion, and celery, nor does it matter (as regards
structure) whether the material is mounted in balsam (Fig. 3), in water (Fig.
4), or in air (Fig. 7). In some instances, the discontinuity of the striae, i.e.,

Fig. 5
Detail drawings of cellulose structure showing micelles and striae: A. from cell wall
shown m Fig. 3 (elder), B. from cell wall shown in Fig. 4, C. optical transverse section of
vertical walls at junction of three cells in onion epidermis (see Photo. 2 for same view with
dimensions).

123

THE SPIERER LENS

Fig. 6

Micellar structure of wall of elder pith cell seen in surface view (same view as lower half
drawing, Fig. 3.)

Fig. 7
Striated structure of the dry cellulose wall of a dead onion cell, surface view.

Fig. 8

Parallel striations of cellulose wall of onion cell, surface view: note lobbed appearance of
striae, the result of a micellar structure which is here less marked than in Fig. 6.

Fig. 9

Detail of transverse optical section of vertical walls at junction of three onion cells: the
first rod (compound micelle) from the junction in the tail of the " Y", is $tx, the second, 3/i,
and the third 2.3M long; the total thickness of a vertical wall, consisting of four striae, is 3m

(see Fig. 5 c for drawing of this same view)

WILLIAM SEIFRIZ

124

the presence of short rods in the cellulose, is lacking (Figs. 7 and 8), though
often the same cell may show continuous striae in one region (Fig 4) and di«<
continuous ones in another region (Fig. 5, c). Furthermore, though separate
rods are sometimes not present and the striae are unbroken lines yet such
striae are nearly always lobbed or undulating, as if built of distinct n-irticles
(Fig. 8). *• '

The striae average from 0.5 to 0.7^ in thickness. The rod-shaped particles
which build up the striae, vary in size, though they average rather close to i or
2M in length; some, however, are two or three times this size, and some are no
longer than broad (0.7/1). The longer ones are probably, like the long un-
broken striae, built up of several unit particles. The i/x rod-shaped units are
not of the order of magnitude which Nageli had in mind when he gave us the
term micellae. The units of these stiae are super-micelles.

A striated appearance in the cellulose walls of plant cells is no new ob-
servation. Strasburger^ in 1882, pictured fine striations in the walls of plant
cells. These markings are visible with an oil-immersion lens and direct
illumination. They have long been believed to be due to the rhythmic de-
position of cellulose by protoplasm. These linear markings seen by earlier
workers may be comparable to the striations described here but they first
lack the discontinuous feature, and second, have been described for trans-
verse optical sections of vertical walls and not for the surface view of horizontal
walls. There are, however, parallel markings to be seen on the surface of cell
walls when viewed with ordinary, direct, illumination. These are quite
another thing; they are twice as coarse as are the striae seen with the Spierer
lens. Spierer, in his continuation of our original joint work, has paid special
attention to the distinction between the coarser microscopic striations and
the finer ''ultramicroscopic'' ones. I shall, therefore, quote, in part, from him
(which IS done with his kind permission).

It is possible to pull the epidermis of plant tissue oflf in such a way so as
to have only a single layer of cellulose, i.e., the upper wall of the epidermal
cells. This cellulose membrane Spierer macerated for six months in water
sometimes sHghtly alkaline, sometimes slightly acid, and sometimes neutral!
It was thus freed entirely from cytoplasmic residues. The membrane was
also, in some cases, boiled. In every instance a structure composed of long
super-micelles was to be seen with the Spierer lens, but with no other system.
The markings which are visible with direct illumination have nothing to do
with the intimate structure of cellulose. These coarser markings result,
apparently, from a very fine folding (undulation) of the membrane. As a rule,'
they are transversal, (viz., perpendicular to the large axis of the cell), and if a'
pull is exerted on the membrane, these markings disappear entirely, or take
the opposite direction, becoming longitudinal on account of the pull. The
true micellar structure does not disappear when the membrane is stretched.
When the membrane is examined carefully by changing the adjustment of the
Spierer lens, the folds are recognized separately from the micellar striae.

'"Zellhaute" (1882).

123

THE tiPIEKEU LENS

WILLIAM SKIKHIZ

124

Fig. 6

Micollar structure of wail of older pith eeli seen in surface view (same view as lower half
drawing, F'm. 3.)

Fig. 7

Striated structure of the dry cellulose wall of a dead onion cell, surface view.

Fig. 8

Parallel st nations of cellulose wall of onion cell, surface view: note lobbed appearance of
striae, the result of a micellar structure which is here less marked than in Fig. 6.

Fig. 9

Detail of transverse optical section of vertical walls at junction of three onion cells: the
first rod (compound micelle) from the junction in the tail of the" V", is 5/x the second. 3^.
and the third 2.3)tx long; the total thickness of a vertical wall, consisting of four striae, is 3^

(see Fig. 5c for drawing of this same view)

t ir ,)r.s(^nco of short rods in the cellulose, is larkin^r , Fi^,.. - ,,,,, s, Uiouirh
often the same cell may show continuous s!ri:i(^ in nnr re-iori , Fi^r ^, .,,,,1 ^ij,
continuous ones in another region ( V^r, 5, ,■ ,. Furth(.rm(.re. thou^rh .eparite
rods are sometimes not present and the striae are unbroken line, yc^t .ueb
striae are nearly always lot,!),,! or undulating, as if built of distinct p-,rticlr<
(Mg. 8).

The striae average froiii 0.5 to 0.7^ in tliickness. The rod-shaped particle.
which build up the striae, vary in size, though tlu^v average^ nitber clo.(> to 1 or
2M in length; some, however, are two or three times this size, and some are un
longer than broad (0.7/1). The longer ones are probablv, like i\w long un-
broken striae, built up of several unit particles. The i^ rod-shaped unit, are
not of the order of magnitude which Xageli had in mind when he gave us the
term micellae. The units of these stiae are sui)er-mic(^lles.

A striated appearance in the cellulose walls of plant cells is no new ob
servation. Strasburger^ in 1882, pictured fine st nations in the walN of plant
cells. These markings are visible with an oil-immersion h^ns and dinrt
illumination. They have long been believed to be due to the rhvthmic de
position of cellulose by protoplasm. Th(^.(. linear markings seen by earlier
workers may be comparable to the striations descrilxHl here but they fir.t
lack the discontinuous feature, and second, have been described for trans-
verse^ optical sections of nrtical walls and not for the surfun view of hnriznnfal
walls. There are, however, parallel markings to be seen on the surface^ of cell
walls when viewed with ordinary, direct, illumination. These are quite
another thing; they are twice as coarse as are the striae sec^n with the Spierer
Ions. Spierer, in his continuation of our original joint work, has paid sj,ecial
attention to the distinction between the coarser microscopic striations and
the finer -ultramicroscopic" ones. I shall, therefore, (juote, in part, from him
(which IS done with his kind permission).

It is possible to pull the epidermis of plant tissue off in such .a way s(» a<
to have only a single layer of cellulose, i.r., the upper wall of the epiilerm:d
cells. This cellulose membrane Spierer macerated for six months in water
sometimes slightly alkahne, sometimes slightly acid, .ami sometimes neutrab
It was thus freed entirely from cytoplasmic residues. Th(. membrane was
also, in some cases, boiled. In every instance a structure composed of long
super-micelles was to be seen with the Spierer lens, but with no other system.
The markings which are visible with direct illumination have nothing to do
with the intimate structure of cellulose. These coarser markings result,
apparently, from a very fine folding ( undulation ) of the membrane. As a rule,
they are transversal, {viz., perpendicular to the large axis of the cell), and if a'
pull is exerted on the membrane, these markings disappear entirely, or taki'
the opposite direction, becoming longitudinal on account of the pull. The
true micellar structure does not disappear when the membrane is stretched.
When the membrane is examined carefully by changing the adjustment of the
Spierer lens, the folds are recognized separately from the micellar striae.

^"Zellhaute" (1882).

INTENTIONAL SECOND EXPOSURE

125

THE SPIERER LENS

WILLIAM SEIFRIZ

126

Question naturally arises over the actuality of these striae and rods, as,
indeed, of any structure "seen" with dark-field illumination; thus, Fierz-
David* working with a cardioid condenser, finds a ''parallel" structure in
artificial silk (cellulose) and natural silk. He observed a granular appearance
in one of his preparations and frankly states that he regards it as not real but
as an optical artifact. One cannot get an artifact without something to get
it from; while of course, this does not preclude having artifacts, yet one should
not be too ready to disregard them but rather be prepared to interpret them.
First, in regard to the striae, the evident criticism here is that these are
diffraction lines. Observations on Uving protoplasm, where the striae are in
motion, clearly oppose such a criticism. Furthermore, diffraction rings occur
at the edges of objects; the ultramicroscopic striae of cellulose cover the sur-
face. The following test finally ehminates diffraction phenomena as the
cause of the observed striae.

The Spierer lens contains an adjustable iris; diffraction lines are eliminated
when the iris is closed. In the classical test object, the diatom, Pleurosigma,
all detailed structure disappears when the iris is closed (the Pleurosigma sur-
face and contours remain visible from diffused light) ; the observable Pleuro-
sigma structure depends, therefore, mainly on diffraction effects. The
striated cellulose structure in plant cell walls, remains unaffected when the
lens iris is closed, proving that this structure is not entirely dependent on
diffraction but is and remains visible by diffused light.

As for the structural units of the striae, the super-micelles, continued
study of them under differing conditions leaves Uttle doubt of their reahty.
Cellulose would not show optical discontinuity without a structural back-
ground which possesses certain discontinuous features. Also, I find it difficult
by any change in illumination to eliminate the very marked broken appear-
ance of the parallel striations; nor does focusing up and down reveal a structure
of which the rods might be the "cut" ends. Personally, I do not doubt but
that the discontinuity in structure shown by the Spierer lens is a real picture
of the ultramicroscopic structure of the cellulose of plant cell walls.

It was my intention to compare photomicrographs of the same material
taken, first with direct illumination, then with the cardioid condenser, and
then with the Spierer lens, but the picture which direct light gives with an
oil-immersion lens is so pale, and the structure to be seen with the cardioid
condenser so blurred when compared with the beautifully clear-cut picture
given by the Spierer lens, that photographic comparison would be useless.
I can, however, state with emphasis that the micellar structure pictured in
the illustrations of this article are not to be seen with ordinary (direct)
illumination, nor with the cardioid condenser, but only with the Spierer lens.
Protoplasm. While observing the cellulose walls of living plant cells,
I often found it difficult to tell whether the lens was focused on wall or inner
protoplasm. This was difficult chiefly because the protoplasm seemed to
have essentially the same type of structure as the cellulose walls. How

^ Naturwissenschaften, 17, 703 (1929).

marked the resemblance may be is to be seen from Fig. ,^ of cellulose and the
lower half of Fig. 4 of protoplasm. The difficulty of distinguishing proto-
plasm from cellulose can be overcome by macerating the tissue and ridding
it of protoplasm, as did Spierer, or by plasmolyzing the cells (protoplasts),
that is to say, causing the protoplasm to shrink away from the cell walls by
dehydrating it in a hypertonic solution. This will leave part of the cellulose
walls free from protoplasm.

Partially surrounding the nucleus (the large sphere in the center of Fig.
4) are concentric layers of cytoplasm, each built up of short rod-shaped
particles. This is protoplasmic material. Where the protoplasm is stretched
into strands — protoplasm often forms stranrls within a cell extending from
wall to wall or from the nucleus to all the walls— the linear orientation of the
rodlets is very pronounced (Fig. 4, below the nucleus), so nuich so that it is
diflficult, on the basis of structure alone, to distinguish a protoplasmic strand
from cellulose.

The nuclear material (Fig. 4) shows no linear orientation of structural
units; we have to do here with a fine emulsion.

The linear arrangement of rods in living protoplasm is, under favorable
conditions, very marked. The rods are light gray in color (translucent) and
the background black. The linear masses are separated from each other by
distances shorter than their length, as in cellulose, and they retain their
relative positions, even while the protoplasm is streaming.

The question, What fills the interstices of a group of micelles? has been
considered by chemists in relation to cellulose, but, I believe, not definitely
answered. The same question arises in the case of protoplasm, but here it is
more easily answered, and with more certainty, though perhaps with no
more satisfaction to the chemist. The black and optically structureless back-
ground, which constitutes, in a sense, the dispersion medium of the visible
rod-shaped particles, is one of the constituents of the complex mixture which
we call protoplasm. Indeed, of these two main components, the visible gray
rods and the invisible black substratum, the latter is probably of primary
importance. I believe this to be true because foreign globules (of fat) are
carried by the ground substance quite independent of the visible rods. Scarth'
supports this in stating that only a portion of protoplasm moves (by which he
must mean that only a portion exhibits active movement). The rods, revealed
by the Spierer lens, move, as do foreign globules also, but their motion is a
passive one; they are carried by the active component, the black optically
empty background.

As in cellulose, the discontinuous character of the protoplasmic striae is
not always evident. The parallel lines are often continuous, but only in
quiescent protoplasm. Whether this, as in cellulose, is a true structural
feature, i.e., the semi-fluid protoplasmic rods have fused, or an optical im-
pression, cannot be definitely said.

' Protoplasma, 2, 189 (1927).

127

THE SPIERER LENS

We have, then, two main constituents of protophism, when viewed with
the Spierer lens, the one, bright illuminated rods, oriented end-to-end in
parallel lines; the other, an optically structureless material filling in the inter-
stices of the rods. Perhaps these two materials are comparable to Stras-
burger's distinction between trophoplasm, the less active nutritive component,
and kmoplasm, the active, irritable component.

We are still nmch in the dark over the ultramicroscopic structure of
protoplasm, and I hesitate to compare too closely the rods seen in proto-
plasm with those seen in cellulose. In form and general appearance, they are
almost identical, but in cellulose they are apparently the primary component
(the nature of the intermicellar substance in cellulose being in doubt), while
in protoplasm the rods are very likely of secondary importance. We must,
however, grant that the remarkable similarity in the structure of protoplasm
and of cellulose in living plant cells, as revealed by the Spierer lens, is a very
striking one.

Discussion
Historical. Nageli, in 1864, advanced the hypothesis that the structural
units of cellulose cell walls are linear, anisotropic, crystalline micelles. This
hypothesis, which was extended to include other colloidal systems of the
lyophilic (gelatin) type, has been critized and discarded from time to time; it
is probably not applicable to all lyophiles. As for cellulose, the Nageli
hypothesis is substantiated by the investigations presented here.

Full support of both the micellar and the striated structure of the cellu-
lose walls of plant cells as revealed by the Spierer lens, is to be found in a
brief survey of the literature.

As for the striae, in addition to the findings of Strasburger^ and Fierz-
David^ already referred to, Anderson,^ working more on the chemical side,
finds that the outer wall of the epidermis of plant tissues is built up of alternate
layers of cellulose and pectin.

The prevailing botanical opinion, that the lamellae of plant cell walls are
the result of a rhythmic deposition of cellulose by the protoplasm, may ex-
plain the presence of certain layers composing the walls of cells. 1 would
rather not question too critically so old a botanical hypothesis which may be
true in part; however, rhythmic deposition is not the cause of the striae illus-
trated here. It cannot be, since the striated structure is as pronounced when
seen in surface view, the plane of the supposed rhythmic deposition, as in

traverse view.

As for the micelles, in spite of the severe criticism to which their existence
has been subjected in recent years, subsequent work continues to bring sub-
stantial support to the micellar hypothesis of the structure of certain lyophilic
systems, notably cellulose.

The cellulose ''molecule" is a chain of some forty anhydrous glucose rings.
Sponsler and Dore^ were the first to indicate, by X-ray studies, a chain

« Jahrb. wiss. Botan., 69, 501 (1928).

^Colloid Symposium Monograph, 4, 174 (i92t>).

WILLIAM SEIFRIZ

128

structure of cellulose. These chains are bundled togetlier into fascicles, to
which Meyer and Mark^ have given the old Nageli term ''micellae," which
may, in cellulose, be regarded as "crystallites" since the internal arrangement
of the chains is orderly. We recall that Nageli found cellulose anisotropic.
Frey,'-* working on the submicroscopic structure of the cellulose walls of bast
fibers and other plant cells, finds them to consist of distinct micelles which
are strongly anisotropic, optically like rhombic crystallites, and seven times
as doubly refractive as quartz.

Clark,^" on the basis of X-ray studies, finds cellulose to be built up of
oriented colloidal micelles which are bundles of long primary valence chains.
On the basis of such a structure Clark gives a rational interpretation of the
physical and chemical properties of cellulose fibers.

The work of Sheppard and McNally,'^ and also that of Hatschek,'-
gives support to the possibility of a micellar structure in gelatin, the sul)-
stance which has been the center of the micellar controversy. Sheppard con-
cludes that the birefringence of gelatin films is due to the orientation of
symmetrical molecules or micelles, but whether the micelles are truly multi-
molecular or simply macromolecules cannot be stated. Seifriz^^ has carried
this conception of a fibrous structure of cellulose and related systems over to
living protoplasma.

Size of Micellae. Chemists regard the micelle as a bundle of chain mole-
cules. No limiting size is given to this bundle, nor can this be readily done,
any more than one can specify dimensions for a crystal (the micelle of cellulose
is a crystalHte). Yet, the cellulose micelle is assumed to be of ultramicroscopic
dimensions. Biologists, on the other hand, have characterized as micelles
particles well above the visual limit of the microscope whether or not they
have viewed these particles with an ultramicroscope. It might l)e well to
arrive at some understanding as to what the term micelle shall denote, even
though we may be forced to conclude that a micelle is no more capable of
precise limitation as to size than is a crystal.

The size of cellulose crystallites varies within limits generally placed at
100—200 A. U. long and 50-75 A. U. thick. An exceptional ma.xiinum length
is 600 A. U. for the crystalUtes of ramie fiber. Even this size is much smaller
than the i/x super-micelles visible in the cellulose walls of plant cells. We
must, therefore, be dealing with a larger unit in the visible (dark-field)
structure of cellulose. These super-micelles are probably aggregates of crys-
tallites. This idea is supported by Herzog'^ who states that a numl^er of
micelles may come together to build up larger structural units which he calls
secondary particles.

» Ber., 61, 593 (1928); see also Mayer: Xaturwissenschaften, 16, 7.S1 {\i)2i^).

9 Jahrb. wiss. Botan., 65, 195 (1926).

>" Ind. Eng. Chem., 22, 474 (1930).

1^ Colloid Symposium, 7, 17 (1929).

•- Kolloid-Z., 35, 67 (1924); 36, 202 (1925).

'•^ Am. NaturaHst, 63, 410 (1929).

'^ Z. angew. Chem., 34, 385 (1921).

THE SPIERER LENS
129

It is at present impossible to say whether the "secondary Particles" of
It IS ^; P^'J"; J:.^i,elles of microscopic dimensions descnbed and

micelle a contmuous and "°if°'^"™,,77''™. „^, answering this question
larger bundle of primary «'y«*^l\'*f . . ^'^ve th^^^^^

rrueoJ primary micelle is the crystallite of Meyer and Maj^ (- X^o A^ •) .
above this there may be a contmuous series from ^he se^'^ndary p
Herzog to the microscopic super-micelles described here (i X 0.5^)-

Summary

lens is that of tiny rods or super-micelles, arranged end-to-end to form long
and parallel striae.

Reprinted from Science, June 12, 1931, Vol. 73, Xo. 1902,

pages 648-649.

Department of Botany,
University of Pennsylvania,
Philadelphia, Pennsylvania.

JOVR: PHYS. CHEM.

THE STRUCTURE OF PROTOPLASM

Insufficient optical differentiation between the
constituent parts of protoplasm has greatly hampered
the advance of knowledge of protoplasmic structure.
Dark-field illumination with the cardioid condenser
has helped but little. The recent invention of a dark-
field oil-immersion objective by Charles Spierer^ is a
very successful forward step in indirect illumination
methods, especially when applied to the study of the
colloidal structure of living matter. The Spierer lens
reveals a structure in living protoplasm, as it does in
celloidin^ and in the cellulose walls of plant cells,*
which is not visible with any other optical system.

The Spierer lens is a Zeiss 1/12 inch oil-immersion
objective with a small platinum mirror electrolytically
deposited at the center of the upper surface of the
lowermost lens of the objective system. This mirror
reflects all direct light, thus producing a dark- field.
The scattered (colloidal) light from the object viewed
is picked up by the lens around the mirror. Increased
detail results because direct light is used instead of
the usual bilateral illumination of the older type of
ultramicroscope. The optical principles involved and
a fuller description of the lens are given in other
publications.^- *

When the hyaline protoplasm of living onion cells
is viewed through the Spierer lens, it is, under favor-
able conditions, seen to consist of two substances, one
brightly illumined, light gray in color, and very finely
granular in texture, and the other, an optically empty,

1 * * Un Nouvel Ultra-microscope k ficlairage Bilateral, ' '
Arch. Sci. Phys. et Natur. (Geneve) 8; 121, 1926.

2 * ' The Spierer Lens and What it Eeveals in Cellulose
and Protoplasm," Jour. Phys. Chem. 118: 35, 1931.

black background. In quiescent protoplasm these two
substances are intermixed as an emulsion and then
present a mottled appearance. Protoplasm under
tension, as it is when formed into a thread (Fig. 1, a),

'^N. «'T-~-«.'*

h

Fig. 1

or when streaming (Fig. 1, b), assumes a striated
appearance, due to the parallel arrangement of long
strands of the illuminated substance. These strands
may be continuous (Fig. 1, a) or discontinuous
(Fig. 1, b) ; in the latter case they are made up of
rods oriented end to end. The striated structure is
seen at its best in actively flowing protoplasm. In-
cluded particles occur, and appear as brilliant globules
imbedded in either the gray matter or the dark inter-
vening substance.

Without any attempt to characterize chemically or
vitally the two phases which make up this dark-field
structure of protoplasm, I propose calling the brightly
illuminated, gray-appearing, and at times discontinu-
ous, dispersed phase, the phaneroplasm, (phaneros =
evident), and the unilluminated, black-appearing,
optically empty background, or continuous phase, the
cryptoplasm (cri/p^os = hidden). In the accompany-
ing figure, the phaneroplasm is black and the crypto-
plasm (the background) white, which reverses what
is white and what is black in the actual material as
seen with the Spierer lens.

* ■ V

8

Both phaneroplasm and cryptoplasm flow, though
apparently not at the same rate, the phaneroplasm
being more sluggish in its movement. The crypto-
plasm is optically empty and can not, therefore,
actually be seen ; its streaming is, however, made evi-
dent by the movement of included particles. A rap-
idly moving particle may, where there is an irregu-
larity in the arrangement of the striae, strike against
the side of a strip of phaneroplasm; its forward
movement is thus retarded, but only for a few
moments while it is slowly pushed through the
phaneroplasm, first thinning it, and then breaking
through. This and other similar events indicate that
the cryptoplasm is the actively streaming component
of protoplasm.

The strands of phaneroplasm are from 0.3 to 0.4 (&
in thickness and 0.2 to 0.3 ji apart*

Except for the optical properties already referred '
to, there is little to be said concerning the physical,
ohemical and vital nature of the two substances which
make up the living hyaline diphasic system. The
phaneroplasm is brightly illuminated but the crypto-
plasm is not. Where the protoplasm, at rest, assumes
a m(?ttled appearance, the phaneroplasm becomes the
dispersed phase and the cryptoplasm the dispersion
medium of an emulsion ; where the protoplasm is -
striated but the striae broken, the structure is still
that of a (distorted) emulsion; but where continuous -
striae exist, there is no distinction between a discon-
tinuous and a continuous phase, that is to say, there
is no emulsion as ordinarily defined.

New terms are convenient if merely as temporary
handles to be discarded as knowledge of the subject
increases, but they add confusion if satisfactory old '
terms exist. Strasberger^ distinguished between kino-

s*<Ueber Cytoplaamsatructurem, " Jour. wiss. Boi, 30-
04 0,, 1S97< .

IRREGULAR PAGINATION

plasm (active substance) and trophoplasm (nutritive
substance), two terms which have been brought into
use again by Lloyd and Scarth;* but these terms are
not applicable to the cryptoplasm and phaneroplasm
described here if Strasberger's original description is
to be adhered to. He says that kinoplasm possesses a
fibrous structure and trophoplasm the structure of a
honey-comb, and that the two substances may be in
quite distinct regions of the cell. No such differences
are evident in the dark-field structure of protoplasm
as seen by the Spierer lens. This lens differentiates
not kinoplasm from trophoplasm, but two substances
which make up the kinoplasm alone.

There does not appear to be any satisfactory corre-
lation between the structure of actively flowing hya-
line protoplasm as revealed by the Spierer lens, and
structures seen and described before. The mottled
emulsion structure of the resting protoplasm is, in a
broad way, comparable to alveolar protoplasm as
described by BUtschli, and to numerous other pseudo-
alveolar, vacuolate and emulsion structures. The
striated structure of protoplasm when under tension,
has apparently no counterpart in the older literature.
The nearer we approach the ultimate structiAe of
protoplasm the less easy it is to differentiate between
the relative importance of its constituents, but the
idea that some substances in living matter are more
important than others is often entertained. If we
attempt to draw a distinction between the relative
"vital" significance of phaneroplasm and cryptoplasm,
then, frequent discontinuity in the former and active
streaming of the latter suggest that cryptoplasm is
the more fundamental of the two.

William Seifriz
Department of Botany,
University of Pennsylvania

4 * * The Role of Kinoplasm in the Genesia of Vacuoles, ' '
Science, 65: 599, 1927.

>

[Reprinted from Journal of Rheology.
Vol. 2, No. 3. July, 1931. J

THE EFFECTS OF SALTS ON THE EXTENSIBILITY OF PROTO-
PLASM

William Seifriz and Janet Plowe

Introduction

The life of a cell is intimately associated with its salt environment,
whether this is the sea, the soil water, or the body fluids. The classical
experiments of Ringer on the biological eflfects of salts initiated the ex-
tensive research which has been done on the relationship between cells
and their electrolytic environment. Our excuse for adding to the volumi-
nous literature on this subject is that we have chosen an as yet little studied
property of protoplasm, namely, extensibility, as the criterion by which to
judge the effects of the salts.

The measure of extensibility used in this work is the maximum
distance to which the protoplasm can be stretched. It is customary and
convenient to speak of the maximum stretching capacity of a substance
as the elastic limit; however, extensibility is not rightly called elasticity,
as the latter term is a measure of the tendency to recover from distortion!
and usually the greater this tendency the less is the stretching capacity;'
thus, steel is more elastic than rubber but rubber is more extensile than
steel.

Method

The method employed is that of micrurgy, which involves the micro-
manipulation of exceedingly fine glass needles controlled by a (Zeiss-
P^terfi) micromanipulator. The material used is the living protoplast,
the protoplasmic contents, of the epidermal cells of the onion (Allium cepii).
The procedure is that developed by Hofler and described by one of
us;« it involves exposing the naked plant protoplast in such a way as to
make the approach of a micro-needle possible without first penetrating the
heavy cellulose wall. The method consists in plasmolyzing (shrinking by
exosmosis of water) the contents of the cells of the epidermis of onion scale
leaves, and then cutting the tissue with a sharp razor. The end walls of a
number of cells are thus cut away leaving some of the plasmolyzed proto-
plasts untouched. The material is then mounted in a drop of the plasmo-
lyzing solution hanging from the under side of the glass cover of a moist
chamber. The protoplasts are reached with horizontal needles through
the open ends of the cells (Figure 1). Ordinarily, protoplasm is sufficiently
adhesive to adhere to the tip of a needle so that when the needle is with-
drawn the protoplasm is stretched to its elastic limit. It is this limit which
we have used as a measure of the effects of the salts. Only when the

2G4

Journal of Rheology

July, 1931

protoplasmic strand breaks along its length, and never when it is severed
from the needle, is the length of the stretched thread taken as the limit of

extensibility.

A stretched protoplasmic thread may be so fine as to be invisible under
high power, but evidence of its persistence is given by the fact that globules
of granular cytoplasm attached to it continue to move apart from one
another and away from the protoplast as the needle is withdrawn. If
stretching is continued, the thread eventually breaks and contracts,
moving so rapidly at first that it may double back on itself like a taut rope
suddenly severed, but later contracting more and more slowly. Finally,
both the hyaline surface protoplasm of the thread and the inner granular
protoplasm of the included globules, flow back into the original mass of the
protoplast. The globules along the strand also serve to indicate the
moment of breaking. Strands are often ten or more times the length of
the cell from which they are drawn, and usually become too slender to see
long before they are severed. In order to determine the length at which the
strand breaks, the needle is stopped at this point, and the distance between

Figure 1

protoplast and needle-tip measured with an ocular micrometer under low

power.

Dissectionists of living protoplasm are constantly confronted with the
task of knowing whether the material at the time of observation is living
and normal, but the question is by no means so serious a one as the un-
initiated are sometimes inclined to believe. Sudden changes in consis-
tency, discoloration, and rate of Brownian movement of granules are
among the numerous criteria for determining the physiological state
of the cell, but no proof more convincingly indicates that the protoplasm
is living and probably normal than does active streaming, when the rate
remains constant. Streaming remains undisturbed in the cell after
stretching parts of the protoplast twenty times or more.

Strips of the upper epidermis of the bulb scales oi Allium cepa were placed
in 0.05 M solutions of KNO3. NaNOa, Ca(N03)2, Mg(N03)2, and wSr(N03)2,
and in 0.01 M solutions of LiN03 and A1(N03)3. After fourteen hours in
these solutions the tissue was moved to a sucrose solution, usually 0.7 or
0.8 M, which would plasmolyze the protoplasts to one-half their original
volume. The material was then cut and mounted. Strands were pulled

Vol. 2, No. 3

Effects of Salts on Protoplasm

265

out from the protoplast with a microdissection needle, and the length at
which they broke measured. Ten successive strands from each protoplast
were measured, and the average of these measurements was used as the
reading for that cell. In some cases the cell was punctured by the needle,
that is, a hole was pierced through the protoplast which opened the vacuole
to the outside solution: in other cases the outer membrane ruptured,
resulting in death, before ten measurements had been made. It was
thought inadvisable to discard the first readings made from such cells
when they were still living and normal, lest the cells measured be an artifi-
cially selected group, consequently, readings from all cells for which four
or more measurements had been made were included in the final calcula-
tions. Table I gives the number of cells experimented upon and the
average number of strands measured per cell for each solution.

Table

I

Salt

No. cells

Av. no.
strands
per cell

Salt

No. cells

Av. no.
strands
per cell

KNO,

00

8.2

Mg(NO,),

79

7.6

NaNOj

64

9.1

LiNO,

61

9 4

Ca(NO,),

60

S.O

Al(NO,)3

64

8.1

Sr(N03)2

65

9.5

Sucrose

120

9 3

When readings for sixty or more cells had been obtained, the mean
reading for each group was calculated. Cells plasmolyzed in 0.7 or O.S M
sucrose without previous treatment were also measured as a control.
In Table II are enumerated the solutions, the mean length of strand, and
the difference between the mean for each solution and the mean for un-
treated cells in sucrose.

Salt

KNO3

NaNOa

Ca(N03)2

vSr(N03)2

Mg(N03)2

LiNOa

A1(N03)3

HNO3 at pH 3.7

Sucrose

Table

II

Concen-
tration
(molar)

Mean lenKth of

strand

(scale div.)

0.05

19

0.05

13

0.05

31

0.05

25

0.05

23

0.01

15

0.01

14
19

0.8

23

Differences between

av. strand salt and

av. strand sucrose

- 4
-10
-f- 8
+ 2

0

- 8

- 9
4

+

The difference between the strand length of cells in 0.8 and 0.7 M sucrose
is very slight, the latter being 21 as compared with 23 for the former.

Mere casual observations of cells plasmolyzed in salts is sufficient to
indicate that strands may be drawn out to much greater lengths from

266

Journal of Rheology

July, 1931

protoplasts plasmolyzed in Ca(N03)2 than from those plasmolyzed in
NaNOa or KNO3.

From Table II it will be seen that the elastic limit of the protoplasmic
surface layer is lowered by treatment with KNO3, NaNOa, and LiNOs,
and increased by treatment with Ca(N03)2. Sr(N03)2 produces a little
increase, although the difference from the control is too slight to be really
significant. Mg(N03)2 produces no detectable change. A1(N03)3 lowers
the elastic limit, as do the monovalent salts, but that this effect is due to
the aluminum ion is questionable without further evidence in view of the
acidity of this salt. Such evidence is presented below. It should be
remembered that the decrease produced by LiNOs is of particular signifi-
cance, because this salt was used in only one-fifth the concentration of
the other monovalent salts.

As the anion was identical in all cases, and all concentrations were hy-
potonic to the cell sap, we may assume that the effects recounted are not
osmotic, and are due to the cations of the salts used.

In order to avoid possible effects of prolonged plasmolysis in sucrose,
as well as the washing out of the salt by the plasmolyzing solution, fresh
material was taken from the salt solution and plasmolyzed every hour.
Within the period of an hour, plasmolysis causes no appreciable change
in the behavior of the protoplasmic strands. Other possible causes of
variation such as those due to differences among cells are rendered negligible
by the extent of the salt effect and the lack of any marked change in the
effect between fourteen and twenty- two hours of treatment. Average
strand lengths after treatment for varying periods, 14, 16, 18, 20, and 22
hours, show that no progressive effect, as the length of treatment is in-
creased, is detectable within these limits.

If the cations are arranged in the order of increase in average maximum
length of strand obtained, we find the series Ca > Sr > Mg > K > Li
> Na. The monovalent ions all fall below the control, the divalent, above.
Similar lyotropic series have been obtained for a number of reactions,
in both non-living (protein) systems and in the living system.

The evident question arises, how much of this effect, attributed to the
salt ions, is really due to ion specificity, and how much to change in
acidity ? For the salts so far put in series this question is answered by the
statement that the pK of the six solutions is practically identical, with
differences too slight to consider, in view of the extent of their influence
on the extensibility of protoplasm compared with that of acid used alone.

The case of aluminum is otherwise. Its salts are notorious producers
of an acid condition. However, further investigation reveals a definite
effect of the Al ion. The pH of the A1(N03)3 solution used was 3.7. Cells
treated with HNO3 at ^H 3.7 yield an average length of 18.75 scale units,
which is four units below normal (sucrose plasmolyzed) cells. Since the

Hl^

.1 ^k

Vol. 2, No. 3

Effects of Salts on Protoplasm

207

strand length in AKNO,), is 14 units, which is 9 units less than that of
untreated cells, then only 4 units of this decrease can be attributed to
acidity, i. e., to the H ion; the remaining 5 units must, therefore, be due
to the Al ion.

Discussion

A number of problems present themselves. First, there is the question
of the validity of the Hofmeister series. If we omit aluminum, we find that
there is a valence grouping of the ions, all monovalent cations lower the
stretching capacity of protoplasm and all bivalent cations increase this
capacity. This grouping is in agreement with the ideas of those who
oppose the existence of Hofmeister series; but within each valence group there
are differences greater than the minimum differences between the groups;
thus, Mg causes no detectable change while Ca increases the elastic limit

8 points above normal, and among the monovalent cations, Na lowers the
limit 10 points as compared with but 4 for K ; yet between the two groups
there are only 4 points between the bivalent cation Mg and the monovalent
K ion.

A second question is the influence of acidity on a Hofmeister series.
The pH values of the solutions differ too little to account for the marked
salt effect except in the case of A1(N03)3 and even here but 4 of the total

9 points are attributable to the H ion, 5 being due to the Al ion.

A further question is that of the nature on the property effected. We
have measured extensibility, stretching capacity, or elastic limit. What
determines this limit? Two evident factors which may influence extensi-
bility are viscosity and surface tension. While it is true that, within limits,
the more viscous or plastic a substance is, the greater is the distance it can
be pulled into strands before severing, other factors remaining the same,
yet, there is little theoretical reason for regarding a marked change in
elastic limit as due to change in consistency, Bingham' has frequently
pointed out that elastic deformation and viscous and plastic flow are in-
dependent variables. But if we should be inclined to so regard our results,
we would find trouble in substantiating that calcium decreases protoplas-
mic consistency and potassium and sodium increase it. Several workers
believe this to be true^ while others do not.^- ^

Another possible factor influencing if not determining extensibility is
surface tension. Surface tension can undoubtedly play an effective part
in the elastic limit of a substance, but, as with viscosity, this part does not
appear to be an important one. When a strand of protoplasm breaks,
it frequently doubles on itself; such behavior does not result in a decrease
in surface as a high surface tension would demand.

We come now to the third possibility, internal structure. If the elas-
ticity of protoplasm is the expression of structural features,^ that is to say,

268

Journal of Rheology

July, 1931

Vol. 2, No. 3

Kffects of Salts on Protoi'lasm

2(>9

if the elastic properties of jellies are due to an elastic framework of fibrous
units, then it is only natural to conclude that changes in the extensibility
of protoplasm are the result of changes in structural properties; either
there is a change in the arrangement of the fibrous units or a lowering in the
firmness of the bonds between them.

Sheppard^ has pictured the arrangement of fibrous micelles in cellu-
losic films of different optical character ; these or other arrangements might
well determine elastic properties.

The formation, both natural and artificial (by plasmolysis) , of proto-
plasmic threads within cells is of common occurrence. The number and
persistence of these threads have been used as indications of protoplasmic
consistency. There is no reason to believe that the viscosity of proto-
plasm is accurately indicated by the abundance or persistence of threads
formed within the cell any more than the extensibility of protoplasm
as here described is related to viscosity.

Scarth* emphasizes that adhesion (stickiness) and not cohesion
(viscosity) may be the determining factor in thread formation.

The significance of determinations of elastic values of protoplasm lies
in their direct bearing on such other physical properties of living matter
as imbibition, contractility, and structure, and their indirect bearing on
such biological processes as cell division, amoeboid movement and, con-
tractility in general.

The ability of protoplasm to contract, of an amoeba to withdraw a
pseudopod, of a muscle to tighten, is inherent in the elastic properties of
protoplasm, which, in turn are dependent upon an elastic framework.
The structural units of this framework are fibers of molecular or colloidal
dimensions, probably crystalline in nature, comparable to the anhydrous
glucose chains of cellulose.^ The tighter these linear units are inter-
locked, the more rigid, more elastic, and less extensile is the material;
the more loosely the units are interwoven, the less rigid is the material and
the greater is the elastic limit. Naturally, as fluidity increases there will
come a concentration at which the structure is no longer capable of holding
together with sufficient firmness to permit stretching, and the elastic limit
will again fall. This point has been determined for gelatin ; at a concentra-
tion of 0.7 per cent, gelatin can be stretched but 18/x (microscopically
determined), while a concentration of 0.8 per cent has an elastic limit of
130m; this latter value gradually falls as the gelatin becomes more firm
with increased concentration, until a value of IS/x is again reached for
1.8 per cent gelatin.^

Summary

1. Cells of the epidermis of Allium cepa previously treated with 0.05
or 0.01 M solutions of the nitrates of Ca, Sr, Mg, Li, K, Na, and Al, were

•z^

• !■■»

« ■ V

\ ^ 9

*m^

^f ♦

plasmolyzed in sucrose and the elastic limit of the protoplasm determined
by drawing out strands with a microdissection needle from the surface
layer of the protoplast until the strands broke; their length was measured
at the moment of breaking.

2. Statistical study of the results show that solutions of KNO3, NaNOs,
LiNOs, and A1(N03)3 decrease the length to which strands can be stretched
without breaking, while the bivalent salts raise the elastic limit; the
effect of Mg and Sr is very slight. The monovalent and bivalent ions
give the series, Ca > Sr > Mg > K > Li > Na.

3. The effect of the aluminum ion is specific in that about one-half
of the total effect of the salt is due to the aluminum ion and one-half to

increase in acidity.

4. The extensibility of protoplasm is only slightly influenced by vis-
cosity and surface tension; it is essentially a matter of structure.

Department of Botany
University of Pennsylvania
Philadelphia, Pa.

Bibliography

1. E. C. Bingham, "Plasticity and Elasticity," J. Franklin Inst., 197,99 (1924).

2. N. Cholodny, "Zur Frage uber die Beeinflussung des Protoplasmas durch
Mono-und-Bivalente Metallionen," Beihefte Z. Bot. Centralbl., 3Q, 1, 231 (1923).

3. Herbert Freundlich and William Seifriz," Uber die Elastizitiit von Solen und
Gelen," Z. physik. Chem., 104, 233 (1923).

4. G. W. Scarth, "Adhesion of Protoplasm to the Cell Wall and the Agents
Which Cause It," Trans. Roy. Soc. Canada, 17, 137 (1923).

5. G. W. Scarth, "Colloidal Changes Associated with Protoplasmic Contraction.
The Action of Cations on the Contraction and Viscosity of Protoplasm in Spirogyra,"
Quart. Jour. Exp. Physiol, 14, 99 (1924).

6. William Seifriz, "New Material for Microdissection," Protoplasma. 3, 191

(1927).

7. William Seifriz, "The Viscosity of Protoplasm. Molecular Physics m Relation

to Biology," Bull. Nat. Res. Council, 69, 229.

8. William Seifriz, "The Contractility of Protoplasm." Amer. Nat., 63, 410 (1929).

9. S. E. Sheppard and J. G. McNally, "The Structure of Gelatin Sols and Gels,
II. The Anistropy of Gelatin Gels," Colloid Sym. Annual, 7 (1929).

Discussion

Chairman Davey: I should like to point out. that some of the results given in Prof.
Seifriz' paper will fit in very nicely with a paper given by Martin Fischer before the Mayo
Foundation, and published in a little booklet by them, giving the lectures from the Mayo
Foundation. There again they found a sort of Hofmeister series, a series very much like
that given in this paper, except that they have gone a step further and investigated the
effect of the negative ions too. Martin Fischer insists that his results with soaps are
analogous to the action of proteins, since the proteins can be regarded as complicated
substituted soaps. If we adopt such a picture, we could predict that Seifriz would find
different elastic effects on these fibers, which would depend upon the speed with which

I

270

Journal of Rheology

July, 1931

he pulled them. If we could take into account Fischer's work and Du Nuoy's work at
the Rockefeller Institute, we see if we pull a fibre so as to increase its surface at a very
slow rate, more protein will diffuse through to the surface from the interior. We would
then have a continual mending of the protoplasm and the surface tension would be the
controlling or dominant factor. The change in the elastic effects would be smaller than
if a more rapid pull had been given. Studies in which ions were put on the sur-
face of the cells, and others in which ions were put in the interior of the cells, show
that the two processes give entirely different results. I am sure that reading that
lecture by Martin Fischer would contribute very much to an understanding of Prof.
Seifriz' paper. Incidentally I think there is a possible contribution here to the cancer
problem, because it seems to be the result of spectroscopic examination that what-
ever that thing is that is called a cancer cell, it has a different ion content than the
other thing which is called a normal cell. It seems that an increase in the calcium or
magnesium content would correspond more nearly to having a normal cell. If we could
have something which would introduce calcium or magnesium into a cell in the same way
that mustard gas can go into a cell and break up and introduce chlorine into a cell, then we
might possibly have an interesting agent for experimenting with cancer. As I under-
stand it, there is not now any substance known which can be made to diffuse inside of
cell and then chemically break up so as to deposit a bivalent positive ion. If such a sub-
stance could be discovered, it would open up an entirely new field in cell research, and by
cell research I mean not only cancer work but fluidity and plasticity and similar effects
in cells.

Dr. Bingham: I think we might draw the conclusion from that paper that we have
here presented a very nice problem for the rheologist to assist the biologist. It seems to
me that the field of fluidity of different salt solutions is one that the rheologist knows
nothing about today and it is extremely important. We know that when we have bivalent
ions in our condensation products, for example, phthalic acid condensing with glycerol,
we have the possibility of chains that polymerize almost indefinitely, making large mole-
cules of high viscosity. We know also that potassium is extremely feebly hydrative,
and calcium and magnesium still more so and aluminum very strongly hydrative. That
is just a clue to a sort of solution that might be offered.

Chairman Davey : That subject of hydration is extremely important and fits in
again with Fischer's picture, for he tried to determine the whole effect in terms of the effect
of these ions, positive and negative, and the degree of hydration of the colloids. He even
takes the Hofmeister series of positive and negative ions and shows why magnesium sul-
fate is a better cathartic than some other sulfates, and citrate still better. Mercury
ions are still better dehydrators. The only mercury compound that is safe to use, of
course, is insoluble, and that means calomel. So we can reduce a very large fraction of the
bag of tricks of the family practioner town to Physical Chemistry. Presumably aspirin,
or better strontium salicylate, acts as a chemical wringer on the cells of the body in much
the same way, causing the cells to drain outward. This is probably why it is one of the
favorite remedies of physicians for the treatment of colds.

f*m.

I „

r ^

A NEW AND RAPID METHOD FOR MAKING PERMANENT

ACETO-CARMIN SMEARS

William Campbell Steere
Botanical Laboratory, University of Pennsylvania, Philadelphia, Pa,

Abstract.— Smear the pollen mother cells of a single anther from
each flower bud on a clean dry slide, using a small scalpel. Flood the
slide with Belling's aceto-carmin and heat for a second over an alco-
hol flame. Examine under the microscope to determine the stage of
microsporogenesis. If the stage is satisfactory, smear the remaining
anthers in the same manner, but fix and stain them by immediate im-
mersion, face downward, in a petri dish full of hot (steaming ) aceto-
carmin for from 1 to 10 minutes. Then rapidly transfer thru the fol-
lowing mixtures: two parts 99^ [ (glacial) acetic acid plus one part
absolute ethyl alcohol; one part acetic acid plus two parts absolute
alcohol; and finally one part acetic acid plus nine parts absolute alco-
hol. The slides are then to be dehydrated completely by 1 to 2
minutes immersion in pure absolute alcohol, and cleared 2 to 3
minutes in a mixture of xylene and absolute alcohol in equal parts.
The preparations are then made permanent by mounting each with
balsam and a cover glass. The whole process takes from 5 to 15
minutes and is particularly recommended for chromosome counts.

For several years the author has been en^'aged in cytological and
genetical investigations in which it frequently has been found nec-
essary to determine the chromosome number of each of many individ-
uals resulting from the crossing of forms with different chromosome
numbers. The aceto-carmin smear method introduced to plant
cytology by Belling (19*21) has been found to he eminently suited to
such undertakings, when the chromosome number is of incidental
value only. If, however, as is usually the case, permanent records in
the form of slides are wished, the most serious fault of this method
appears. Since the aceto-carmin method, as described by Belling,
gives essentially temporary results, even when the preparations are
sealed, the author has heretofore been forced to rely upon much
slower methods for his permanent slides.

Belling (19^26) has casually mentioned making aceto-carmin mounts
permanent by passing them thru grades of acetic acid into })alsam.
McClintock (19'29) has improved upon Belling's suggested procedure
and has given details for an efficient method of making aceto-carmin
smears permanent. Her procedTire depends, however, on macerating

Stain Technology, Vol. VI, No. 3, July, 1931
107

IRREGULAR PAGINATION

108

STAIN TECHNOLOGY

the anthers in aceto-carmin, heating the preparation in order to affix
the pollen mother cells to the slide and cover glass, and then soaking
them apart with dilute acetic acid. Great care must be taken during
the process not to dislodge the pollen mother cells, and both the slide
and cover glass have to be carried thru the stages of glacial acetic
acid, then dehydrated and finally reunited in balsam.

Considering the time and patience required for the above proced-
ure, the permanent smear methods of Taylor (1924) and Mann (1924)
and the various modifications of the former have been found more
efficient, since they require about the same length of time and give
results much finer and more accurate in all ways. Newton's (1927)
modification of Taylor's smear method has been found particularly
useful in the past.

In order to secure large numbers of chromosome counts in the
shortest possible time, the author has devised a very simple and rapid
method for making permanent aceto-carmin smears.

Flower buds from the plants to be investigated are brought into the
laboratory from the field or greenhouse and kept fresh in moist paper
until ready for use. One anther is removed from a bud of approxi-
mately the right size, and the pollen mother cells smeared out on a
clean dry slide with a small scalpel, especially honed flat for this pur-
pose. The anther fluid and disintegrating tapetal cells are viscous
enough so that the sporocytes adhere rather firmly to the glass. The
slide is then flooded with Belling's aceto-carmin (45% glacial acetic
acid saturated with carmin,' then filtered) and heated over an alcohol
lamp until the liquid steams, when it may be examined microscopic-
ally. With some practice, the stage of microsporogenesis may usually
be determined immediately, using the low power, with which a cover
glass is not necessary. If the proper stage is discovered, the remain-
ing anthers are smeared and stained in the same fashion, otherwise
the test slide is wiped clean and the bud discarded. If the pollen
mother cells of the test slide tend to loosen up and float away when
the aceto-carmin is poured on, or when the slide is heated, an alterna-
tive method is recommended. Immediately after smearing, instead
of being flooded with aceto-carmin, the slides are immersed, smeared-
side downward, in the cover of a petri dish filled with hot aceto-car-
min. A very short piece of small glass rod or tubing cemented to the
bottom of the dish, at one side, will serve to hold one end of the slide
from the bottom and prevent the pollen mother cells from being

^Good resiilts were ol)tained with pre-war Grubler's carmin. with Coleman and Bell's
carmin and National Aniline carminic acid, but not with carmin Merck. Poor results
are sometimes due to using insufficient dye to saturate the solution.

^ w p

ACETO-CARMIN SMEARS

109

rubbed off. The cover of a 10 cm. petri dish allows room for two slides
side by side. This latter method has been found to solve the diffi-
culty in the frequent cases in which the sporocytes tend to loosen,
since the liquid meets the surface of the slide with equal force at all
points, and the heat apparently causes the cells to adhere more firmly.
The heat of the fixative may be maintained l)y keeping the petri dish
on a hot plate or substage microscope lamp.

The process of fixing and staining in the aceto-carmin will take
from one to ten minutes, depending on the material. The time re-
quired for the species under investigation is soon determined, and the
procedure can be standardized accordingly. If the chromosomes do
not stain deeply enough within this time, a small amount of ferric
hydroxide added to the stain will remedy the difficulty, as Belling
first discovered. A few iron filings in the aceto-carmin will do just as
well, or even the blade of a steel scalpel held in the staining solution on

the slide.

When a satisfactory stain has been secured, the aceto-carmin is
drained of! as completely as possible and the slides rinsed carefully in
a Coplin jar containing two parts 99^^ (glacial) acetic acid and one
part absolute alcohol. This wash usually removes all anther walls
and other debris. The slides should next be transferred rapidly thru -
a mixture of one part acetic acid and two parts absolute alcohol into
one part acetic acid plus 9 parts absolute alcohol, where they may be
left from thirty seconds to two minutes. A prolonged stay in the
first two acetic acid-absolute alcohol mixtures is liable to remove the
stain from the chromosomes as well as from the cytoplasm. Dehydra-
tion is completed by allowing the slides to remain 1 to 2 minutes in
absolute alcohol. Clearing is accomi)lished by transferring the slides
to a mixture of xylene and absolute alcohol in equal parts, leaving
them for 2 to 8 minutes. A cover glass is then placed on the smear
with a drop of balsam, and the permanent slide is complete.

The method of mounting in Canada balsam directly from the
mixture of absolute alcohol and xylene, as suggested by McClintock,
was found to be entirely satisfactory. It is also considerably more
rapid than running the slides thru the several stages of absolute al-
cohol and xylene which would be required to bring the fresh micro-
sporocytes into pure xylene without undue shrinkage.

The chromosomes in successful preparations will appear from dark
red to deep purple, while the cytoplasm will be practically colorless.
The slightly acid reaction of Canada balsam and the small amount of
acid carried over in the cells should tend to keep the stain from fading.
With care, a counterstain may be added in the absolute alcohol stage.

110

STAIN TECHNOLOGY

ACETO-CARMIN SMEARS

111

If the cytoplasm appears very dark, a shorter staining period is
recommended. Overstained slides may be differentiated singly by
flooding them with pure glacial acetic and warming moderately, until
the stain has reached the desired intensity. In routine work on mate-
rial which persistently overstains, a well of glacial acetic may be
interpolated between the stain and the first acetic acid-absolute al-
cohol mixture. It should also be kept in mind that heat applied dur-
ing the staining process will intensify the stain in the chromosomes
and usually keep the cytoplasm from staining too heavily. This effect
of heightened temperature on the stain's selectivity for chromatin
has been discussed previously by McClintock.

The following schedule is included for reference:
1. Secure fresh flower buds of the desired plant.

Smear the pollen mother cells from one anther on a dry slide.
Immediately flood the smear with aceto-carmin.
Heat carefully over an alcohol flame until the stain steams.
Examine under the low power of the microscope (a cover glass

is not necessary) after 1 to 5 minutes of staining.
Smear remaining anthers if the stage is satisfactory.
Stain these slides by either flooding with, or immersion in, aceto-
carmin, for 1 to 10 minutes.
Two parts glacial acetic acid plus one part absolute ethyl

alcohol. Rinse.
One part glacial acetic acid plus two parts absolute ethyl alco-
hol. Rinse.
One part glacial acetic acid plus nine parts absolute ethyl alco-
hol. ^ to 2 minutes.
Equal parts xylene and absolute ethyl alcohol. 2 to 3 minutes.
Mount in Canada balsam.
During the winter months of 1930-31, this method was tried out on
several different species, both monocotyledons and dicotyledons.
Good results were secured from Rhoeo^ Tradescantia and Velthemia,
while Nicotiana and Petunia gave entirely satisfactory results. The
method is now being tried on animal material.

The procedure, as outlined above, is recommended for securing
chromosome counts in pollen mother cells, as is often necessary in
genetical work. It may also be used for more detailed cytological
work, especially if the smears are made upon cover slips, which are
then treated in the same manner as slides. This method brings the
pollen mother cells into direct optical contact with the lens, thru the
medium of the immersion oil.

2.

3.
4.
5.

6.

7.

8.

9.

10.

11.
12.

^f\

The rapidity and simplicity of this method is due mainly to the
fact that the anthers are smeared on the slide before the aceto-carmin
is added, instead of being macerated into a drop of the stain. In the
latter case, it is rather difficult and tedious to secure a good permanent
mount without losing many of the sporocytes.

When working with a plant which responds well to this method,
permanent slides, ready for examination under the lower powers of
the microscope, may be prepared in from five to fifteen minutes, since
a satisfactory stain may usually be secured in from two to three
minutes, and only a minute or so is required in each grade of dehy-
drating and clearing solution. If material is abundant, a series of
slides may be kept going thru the various grades, in succession, so
that it is often possible to make a large number of permanent prepara-
tions in a short time.

This technic, then, appears to be one of the most rapid yet evolved
for the preparation of permanent slides demonstrating meiosis in pol-
len mother cells.

REFERENCES

Belling, J. 1921. On counting chromosomes in pollen mother cells. Amer. Nat.,
55, 573-574.

1926. The iron-aceto-carmin method of fixing and staining chromo-
somes. Biol. Bull., 50, 160-162.

McClintock, B. 1929. A method for making aceto-carmin smears [)ermanent.
Stain Techn., 4, 53-56.

Mann, M. C. 1924. A method of making permanent smears of pollen mother cells.
Science, 60, 548.

Newton, W. C. F. 1927. Chromosome studies in Tulipa an<I some related genera.
J. Linn. Soc. Bot., 47, 346.

Taylor, W.R. 1924. The smear method for plant cytologj-. Bot. Gaz., 78, 230-238.

y

IRREGULAR PAGINATION

Reprinted from the American Journal of Botany, XVII: 9^3-994, \^^^-i 1930

NUTRITION OF THE CULTIVATED MUSHROOM

J. Franklin Styer
(Received for publication May 15, 1930)

The art of mushroom growing has been passed on from generation to
generation without the development of exact methods. There is a great
and costly lack of understanding of the biological processes which influence
this culture. This situation must be charged to the fact that until the last
few years the principles of spore germination and spawn priKiuction were
poorly understood; and students who developed spore germination methods
usually entered the spawn business without carrying on further investiga-
tions. Although the attitude of the spawn producers toward ex j:>eri mental
work has been very helpful to the industry, they have in most cases lacked
laboratory facilities for the kinds of work that were needed. Fortunately
the development of the industry in America and Germany is leading to a
concerted attack on the problems of mushroom growing, by specialists in
the lines involved.

This study concerns the growth of the mycelium of the cultivated mush-
room on simple media of known composition. The natural habitat of this
mushroom is the sod of meadows and plains, but the medium used in mush-
room growing is a partially decomposed preparation of horse manure.
Chemical studies of this medium were begun by the writer along with this
investigation, but the complexity of manures is such that the value of any
such analyses is as yet uncertain. The record of growth on simple media
throws more light on the nutrition of the organism and forms a more
reliable basis for further work.

The literature on the subject of mushroom nutrition was reviewed in a
preliminary paper (10). The outstanding work of Duggar (2) was the
only one known to the writer in which a study of nutrition was the chief
aim. Duggar's cultures were made upon filter paper dampened by nutrient
solutions, and the physical nature of the media was supposed to resemble
that of manure. His results with a series of carbonaceous materials in-
dicated that the organism preferred starch as compared with sugars, and
proteins above all other compounds. Salts of organic acids were appar-
ently of no value. As sources of nitrogen the proteins were best, and growth
was uniformly better with ammonium salts than with nitrates. Calcium
hippurate seemed useful as a source of nitrogen.

The writer's first study (10) of this problem was carried out on the
same plan as Duggar's. The ratio by weight of li(|uid to paper in Duggar's
flasks was about 4 to i, or 4(30 percent moisture, while the ratio in the
writer's was 2 to i, or 200 percent, and there was better growth by reason

983

i'

984

AMERICAN JOURNAL OF BOTANY

[Vol. 17,

of better aeration. The importance of aeration as a factor will be shown
later in the present paper. The results of this series agreed in general with
those of Duggar's, but it was found that the paper base alone could serve
as a nutrient. It was also shown that while ammonium salts may be fair
sources of nitrogen, the total nitrogen in that form in the medium must be
small, about o.i molar. Complex organic compounds could be added
safely in much larger amount. In both nitrogenous and carbonaceous
series the more complex and less soluble nutrients gave the best results.

Careful examination of these results and comparison with Duggar's led
the writer to suspect that the poor growth of the cultures upon soluble carbo-
hydrates and nitrogenous compounds might have been due to the higher
concentration of those media, rather than to their unsuitability as nutrients.

Concentration of Media

In the preliminary study the solution of nutrient salts with which the
paper was dampened had a concentration of 0.172 M. All cultures in which
other soluble materials were added to the extent of o.i M or over, failed
completely. Duggar's solution A was made up of salts totaling 0.16 M,
He added to this solution other soluble materials; for example, i}4 percent
dextrose, the concentration of which is 0.08 M. In both these series the
salts were highly dissociated, further raising the concentration from the
osmotic point of view.

A new series of cultures was made to ascertain the effect of concentration,
with graded concentrations from 0.1 AI to 0.3 M, and with more than one
nutrient balance. It was found that the total concentration must be
below 0.2 M for vigorous growth, and that the partial concentrations of
individual nutrients may be varied without effect on this maximum point.
Dissociation of salts, however, must be considered, and in a solution of 0.2 M
concentration highly dissociated salts may be permitted up to 0.1 AI only.

This result affords a good reason for the apparently greater value of the
less soluble nutrients. The addition at the beginning of soluble nutrients,
for example dextrose, might raise the osmotic effect of the medium to a
point where the mycelium would not be able to grow at all. The insoluble
materials, for example starch, could presumably be added in larger quantity,
and be made available gradually by enzym action. This affords also a
possible leason for the better growth obtained by Duggar in washed manure.
It is a practice of spawn producers to wash the manure used for pure cultures.
From the same considerations it would be reasonable to expect that the
addition of soluble nutrients to manures used in mushroom growing would
harm the spawn. This would depend upon the concentration of the manure
solution and the time when the application is made. In Duggar's experi-
ments the results of such additions were negative. Experiments upon
manures and other media for mushroom growing, which are becoming
numerous among the growers, should be carried out with careful consider-
ation of this important factor.

Dec, 1930I

stver

MUSHROOM NUTRITION

985

Moisture Context

The growth of the mycelium is indifferent in liciuids. The species seems
to be intensely aerobic, and too much moisture in fibrous media such as
manure will invariably produce an adverse effect upon growth. The danger
point in moisture content appears to be the degree of saturation at which
free movement of air is cut off.

To determine this degree a number of test-tubes were tightly packed to
a depth of 10 cm. with manure which had been washed with distilled water,
and distilled water added so as to obtain a series with moisture content
ranging from 125 percent to 250 percent of dry weight. The latter figure
was the limit of moisture content so that aeration was cut off except through
water. The tubes were inoculated at the surface of the medium, and the
growth downward was measured. Growth was best at 125 percent, fell off
very slowly to 200 percent, and at 22^ percent dropped to less than half.
With a total volume of manure of 25 cc, the air included at 125 percent
moisture was 11 cc, and at 225 percent was reduced to about 3 cc. At
saturation no downward growth occurred.

A second experiment in flasks with filter paper and the nutrient solution
used in the preliminary investigation (10) gave similar results. This series
covered a range from 50 percent to 350 percent moisture content, (irowth
failed below 100 percent, after a feeble start; it was good up to 233 percent,
and then fell off sharply. Aeration was better in the flasks than in the tubes,
having been actually cut off only at 300 percent. Aeration in a mushroom
bed six inches deep should be good at 200 percent, if, as is usual, the surface
is allowed to dry.

It is worthy of note that the nutrition experiments of Duggar discussed
above seem to have been carried out with 25 cc. of solution added to each
6 gms. of paper, which would be a condition very unfavorable to growth.
Many of his cultures grew only at the surface. The results would probably
have been clearer if the moisture content had been better adjusted.

All the cultures affected by high moisture content develoi:>ed excessive
numbers of mycelial strands. These have recently been described at length
by Hein (6), and were attributed by him to the stimulus of moist conditions.
He stated the least number of strands were produced at a point between
45 percent and 80 percent moisture content; and although the impression
was given that the best growth occurred at this point also, that was probably
not his intention. There is a tendency to formation of strands in old
cultures, and spawn producers are often forced to discard a strain which
becomes "stringy." Certain organic nutrients cause strand growth as will
later be shown. But if other conditions remain the same, the writer agrees
with Hein that increases in moisture result in increases in the number and
size of the strands.

Since the strands form profusely in the wetter media, and since they are
also essential to fruiting through the translocation of food materials, a

986

AMERICAN JOURNAL OF BOTANY

[Vol. 17.

connection between aeration or respiration and fruiting is suggested. The
grower maintains his beds dry up to a certain point, and then induces the
production of sporophores by watering. It seems probable that while
the initial growth is favored by perfect aeration, fruiting is brought about by
reduction of aeration, and the addition of moisture is a means of reducing
it. Many other factors enter into crop development, but a study of respir-
ation in this connection seems to the writer to have great possibilities.

Silica Gel Medium

After the preliminary study brought out the fact that the cellulose of
filter paper could support growth, and after agar was found to have inherent
nutrient qualities, the writer turned to silica gel as an inert solid medium.
It is an ideal medium for this type of work, for with surface growth aeration
ceases to be a factor requiring consideration. The mycelium can also be
observed with ease and comparisons can be made more exactly. The
development of this medium has been described in a recent paper (11).

The medium was prepared by mixing two solutions; a potassium silicate
solution 0.2 M with respect to K2O, and an acid solution consisting of
H3PO4. 0.135 M; MgS04, 0.015.1/; CaCls, 0.006 i/; and FeS04, trace.
For the series of non-nitrogenous nutrients NH4N()3 was added to the acid
solution in 0.15 M concentration. The alkaline solution was poured into
the acid solution until a pH of 6.6 was reached, the optimum for this species
according to Frear, Styer, and Haley (5), and water added until the volume
was one and one-half times the volume of acid solution used. Thirty cc. of
this mixture were poured into each plate and allowed to gel. The plates
were dialysed in a dishpan full of tap water until about half the soluble
materials were removed. The nutrient gel was then of the following com-
position: MgS04, 0.005 Af; CaClo, 0.002 M; FeS04, trace; potassium
phosphates, 0.045 M. The gel framework was a 1.2 percent content of
Si02, hydrated. When NH4NC)3 was present its concentration was 0.05 M.

After the silica gel plates had been dialysed, the substances whose nu-
tritive value was to be tested were added in finely powdered form. The
plates were then sterilized in the autoclave, and inoculated in the center
with a small amount of a pure culture on manure. Within reasonable
limits no difference in growth has been observed from different sized pieces
of inoculum. During sterilization and subsequent standing the soluble
substances diffused through the gel, while the others remained as even
surface films. The amount of any insoluble material used was adjusted so
that the film was not too thick to allow microscopic observation.

Spore Germination

The cultures used were pure spore cultures, germinated and growing in
washed fresh manure. A distinct strain of mushroom called "Snow-white"
was used in all cultures. This strain originated in the beds of a Pennsyl-

Dec, 1930]

STYER

MUSHROOM NUTRITION

987

vania grower; it may be recognized by its failure to develop pigments when
grown in light. It is probably a strain of Agarkus campestris. No dif-
ficulty was met with in germinating the spores, after they had been col-
lected under aseptic conditions. A brief description of the method follows.

Pieces of tissue with gills attached were removed aseptically from
vigorous mushrooms before the rupture of the veil, and placed on match
sticks in dry sterile Petri dishes. The sporophores naturally contain foreign
organisms which may develop on the gills and cut surfaces of the pieces,
and may contaminate the spore prints. This was prevented by permitting
air to circulate under and around the pieces, which were cut between one
and two centimeters square. Plates in which the tissue became damp or
moldy were discarded, and the pieces were removed from all plates within
60 hours.

Small squares of sterile filter paper were drawn through the sjK)re prints
until heavily coated with spores, and then placed in tubes of sterile washed
manure. The tubes were incubated at 25° C\, and a mycelium was observed
growing out from the spores in from six to ten days.

The quickness of germination and growth in these tubes is a matter of
especial interest. The question arises whether the same principles are in-
volved in this and the other known methods of stimulating germination of
these spores.

The early work in this field, adequately reviewed by Miss Ferguson (4),
failed to make known any dependable method of germination. Miss
Ferguson found that the presence of mycelium or previously germinated
spores of the same species served to hasten germination in hanging drops
and to increase the number germinating to almost 100 percent. Duggar
(2) found that this effect was not produced by sporophore tissue, but only
by active mycelium. He obtained 10 percent germination in five days in a
few solutions containing various acid phosphates. The reaction of these
solutions was not discussed.

Richard and Olga Falck (3) advanced the hypothesis that one or more
organic acids, produced by the fungous flora which normally precedes
Agaricus campestris in outdoor manure beds, were the specific stimulants
aiding germination. They obtained stimulation by the use of 0.25 percent
succinic acid in malt and manure extracts and distilled water, the germina-
tion amounting to 50 percent in nine days. Other acids failed to stimulate
germination, although they caused the same pH changes. Bechmann (i),
in a series of tests with careful pH control, obtained the best germination,
100 percent in 19 days, at pH values between 5.0 and 6.0. At this reaction,
attempts at stimulation with the aid of several enzym preparations and
autolyzed tissues, including the tissue of Agaricus campestris, failed.

Bechmann considered the sugars in the malt extract used by him to have
been the greatest factor in germination. Lambert (7) used a mixture of
sucrose, maltose, and dextrose in agar and obtained plentiful germination.

988

AMERICAN JOURNAL OF BOTANY

[Vol. 17,

ll!'
i

Hein (6) obtained 75 percent germination in 14 days in distilled water

alone.

These results lead to uncertainty as to what are the stimuli which cause
germination. It is certain that the spores vary in their quickness to
respond. Most of the investigators reported the germination of the spores
in two groups several days apart. Miss Ferguson's explanation of this
phenomenon is that the second group germinates only as a result of mycelial
growth from the first. She cited the prompt germination of both these
groups in drops containing mycelium from the beginning. The writer has
obtained the same results in distilled water in test tubes.

This effect seems to have been obtained by the writer not only from
mycelium, but from spores alone when massed. In making the stock
cultures referred to above, the spores were in thick masses on the squares of
filter paper, and an easily visible mycelium was produced within ten days.
When the same approximate number of spores was distributed through the
manure, or when the paper carried less spores, the mycelium appeared more
slowly and sometimes did not appear at all.

It seems possible that carbon dioxid causes the germination of the more
passive group of spores, either directly or through some influence on pH
or oxidation-reduction potential. Apparent necessity for carbon dioxid
in germination has been reported by Platz, Durrell, and Howe (8) for
Ustilago zeae. Germination in a given time increased from 10 to over 50
percent by saturation of the solution with carbon dioxid. Unfortunately
this was accompanied by a great change of pH and their cultures were not
compared with cultures in which the pH was changed by other means.
Rippel and Bortels (9) found that the removal of the carbon dioxid from the
atmosphere in which the spores of Aspergillus niger were placed increased
the time required for germination, and reduced the number germinating.
When the few germinated spores produced a mycelium the rest germinated
at once, a phenomenon quite like that observed by Miss Ferguson for
Agaricus campestris. They added the following very interesting statement:
"Now if germination did occur here and there over potassium hydroxid at
so early a time, this was the case at only those places in which the spores lay
together in thick heaps." The parallel between these cultures and those of
the writer is striking.

If carbon dioxid is not a factor, some substance such as an acid or
enzym, or some effect more difficult to detect, must emanate from the
spores and mycelium to cause germination.

Nutrition Experiments

The growth of the mycelium on the plates to which organic materials
were added is recorded in table i. The list of these materials was chosen
so that it would represent all the classes of organic constituents of undecom-
posed and decomposing plant tissues as found in manure. Some types of
soil organic matter were also included.

Dec. 1930I

STYER

MUSHROOM XUTRITIOX

989

•f I

The diameter, density, and strand development of the mycelium were
recorded after 18 days growth at 25° C. Growth continued in all the
vigorous cultures until the plates were full, but the results were recorded at

Table i. Mycelial Growth of Agaricus campestris After iS Days at 2^° C. Upon Silica Gel

Plates With a Variety of Organic Nutrients. Plates Contained XlfiXO:^, 0.0$ M,

Except Where the Organic Material Contained Available Nitrogen

Class

Acids .
Sugars

Starch group.

Lecithin

Cholesterol . . .
Gums, pectins.

Hemicelluloses

Cellulose

Ligno-complexes

Humus

Proteins, etc.

Blank

Material

Amount

Succinic acid
Citric acid. . .
Arabinose . . .

Xylose

Dextrose . . .

Diam-
eter

(cms.)

Dcnsitv

Galactose
Maltose. .

Mannite

Starch from wheat

("ihcogen

Inulin

0.0,^ M
0.02 M
0.04 M
0.04 M
0.04 M
0.12 M
0.20 M
0.04 \f
0.02 M
o.io M
o.io .1/
0.04 M
0.25 gm.
0.25 gm.
0.25 gm.
0.25 gm.
0.25 gm.
0.25 gm.
i.(K) gm.
2 cc.
0.30 gm.

0-55 K"i-

Gum arabic

I\'ach gum, from wound
"Certo," commercial pectin

X\ Ian

Agar

Coconut, after ether extrac

tion

Wheat bran

Wheat straw

Shredded filter paper ...

Ground cellulose |o.2S gm.

j<>-^5 K'H-

0.32 gm
0.42 gm
0.32 gm
0.25 gm.

Liriodendron wtjod

Cierman granulated jx'at
moss, commercial "G.P.-

M.

W'cx)d, cellulose remo\ed l)y

rot, 80 percent lignin

2 percent Xil^OH extract of

rich soil

2 [xrcent XH,()H extract of

rich s<3il

"JIunms" from Xew Jersey

defxjsits

Xucleic acid from yeast

Xucleoprotein from yeast . .

IVptone

( iliadin

Glutenin

Casein

Albumin

Xo organic material

0.25 gm
0.36 gm,

0.20 gm,

0.25 gm.

0.25 gm.

0.15 gm.

0.20 gm.
0.30 gm.
O.IO gm.
0.30 gm.
0.30 gm.
0.20 gm.
0.30 gm.
0.30 gm.

30
0.0

4-5

3-0

.V5
2.0

1-5

4-5

5-5
0.0

3.0
30
0.0

4.0
0.0
i.o

50

4.0

2.0

5-5
4-5
1.0

2.5

2.5
7.0

5.0

9.0

7.5
30
1.0

2.5

4-5
4.0

3-5
3.5
6.5
4-5
50
0.0

Sparse

Medium
Dense

Medium

Thin

-Medium

IX'Use

Medium

Dense

Thin

Den.->e

Very dense

Dense

Thin

Dense

\'er\- dense

I )ense

Thin

Dense

Medium

Dense

\'ery thin

Dense
Thin

Wry thin

Dense

\'ery dense

Dense
44

it

Strands

Many

Few

Few

Many

Few

Few

Few

Many

990

AMERICAN JOURNAL OF BOTANY

[Vol. 17.

Dec, 1930]

STYER — MUSHROOM NUTRITION

991

18 days because after that time strands began to appear in all the plates.
The strands were recorded as *'few" when appearing only in the center,
and as "many" when extending within yi cm. of the tips of the hyphae.

The pH of the gel was changed to 4.0 by the addition of succinic and
citric acids. As this is far from the optimum pH, the results should be
compared only with each other.

It is necessary to note that the materials containing hemicelluloses and
lignin were quite impure chemically, and that their classification is only
approximate. For example, wheat straw contains cellulose, gums, pectic
substances, lignin, and hemicelluloses. Wheat bran was more correctly
classified with the hemicelluloses. Woods of the class to which Liriodendron
belongs contain about 50 percent cellulose, 20 percent pentosans, and 20
percent lignin. The granulated peat moss is much higher in lignin content.
The black peaty soil from ancient bogs of New Jersey, sold under the name
of "humus," may have been produced mainly from the lignin fraction of the
original organic matter, but, like real humus, is quite different from lignin.
For the selection and classification of these complex materials the writer
has followed the principles of Waksman (12).

The growth of the mycelium upon plates with various nitrogen sources
is recorded in table 2. It was necessary in the two series to supply every

Table 2. Mycelial Growth of Agaricus campestris After 18 Days at 25° C. Upon Silica Gel

Plates With a Variety of Nitrogen Sources. Plates Contained Dextrose, 0.035 M;

Maltose, 0.035 M; and Starch, o.i gm.. Except Where the Nitrogenous

Compound Contained a Carbohydrate Group

Class

Salts

Urine nitrogen
Amino-acids . .

Material

Proteins, etc.

Blank

(NH)2S04

NH4NO3

NaNOa

Urea

Creatine

Creatinine

Cilycine.

Glutamic acid

Arginine carbonate

Leucine •

Nucleic Acid (Also in table

I) ;

Nucleoprotein (Also in table

I)

Peptone (Also in table i). .

(iliadin " " " " • •

utenin

C-> • <

.asem

Albumin '

No nitrogen

Amount

0.03
0.05
0.03
0.05
0.05
0.05
0.05
0.05
0.03
0.03

M
M
M
M
M
M
M
M
M
M

Diameter

Density

n

0.30 gm

o.io gm
0.30 gm
0.30 gm
0.20 gm
0.30 gm
0.30 gm

2.5
4.0

1-5

1-5

2.5

30

3.5

1.5
2.0

4.0

4-5

4.0
3.5
3-5
6.5
4-5

50
4.0

Medium
Dense
Thin
Dense

Medium

Very dense

Dense

Thin

Very thin

Dense
V^ery dense

Dense

i<

(<
Sparse

Strands

Many

culture with all nutrients except the one to be tested. This was done in
the case of the organic series by adding ammonium nitrate whenever the
organic nutrient did not contain nitrogen in a form already proved available.

A

In the nitrogen series, when the nitrogen source did not contain carbohydrate
groups, dextrose and maltose were added, each in 0.035 molar concentration,
along with about o.i gm. of starch. The compounds are grouped in the
table according to complexity, with a separate group for nitrogen forms of
urine.

The growth was dendritic and the mycelium was irregular in outline in
the cultures on sugars. The peach gum and pectin cultures were quite
different, spreading evenly and entirely in hyphal form, with many aerial
hyphae. The aerial hyphae of the pectin plates presented a powdery ap-
pearance, due to great numbers of crystals on their surfaces. The nncelium
changed most of the materials classed with the hemicelluloses into semi-
liquid pastes. Xylan and fat-free coconut when thus liquefied practically
submerged the mycelium, and may have interfered with its growth; the
growth was dense, however, in advance of the liquefied zone. The growth
on wheat bran, straw, and all the lignin-bearing materials was much the
same as the typical dendritic growth on maltose.

The protein cultures resembled the xylan cultures in that the growth
was dense, and accompanied by active digestion. The hyphae were
straight, pointed directly outward, and were surprisingly equal in length;
their tips were observed emerging from the liquefied protein mass un-
branched and equidistant, like the teeth of a comb. This type of growth
was quite different from the dendritic growth of the maltose cultures, and
the enlargement into strands was much slower. Active leading growth in
the maltose plates was focused into a few hyphae, and strands were built
up from those hyphae after the plates became full. This was not the case
in the protein plates.

The growth on nucleic acid and nucleoprotein was extremely thin, with
long, unbranched hyphae. This effect was not noted in other plates.
The mycelium had a silky appearance, with no strands.

The absence of nitrogen except in the inoculum resulted in the production
of many aerial hyphae. The growth was sparse but spread quickly;
evidently the nitrogen of the inoculum could be made use of even when
spread out quite thinly. The cells of these plates were very commonly
bulbous at both ends.

Discussion*

The results of these experiments indicate that the organism attacks a
great variety of substances. Duggar's conclusion that sugars fail to
support it is shown to be erroneous; his culture solution was probably made
too concentrated by the addition of sugars. The growth with xylose,
dextrose, and maltose supports the opinion that the mycelium grows upon
complex organic matter by reason of the production of sugars. It would be
interesting to attempt a culture upon cellobiose, which is supf)osed to be
produced by organisms which break down cellulose.

Cellulose supported only a little growth when spread out on silica gel.

992

AMERICAN JOURNAL OF BOTANY

[Vol. 17.

Dec, 1930]

There was more opportunity for the products of hydrolysis to diffuse away
from the mycelium than in the preliminary paper base cultures, where the
paper was packed in tightly. In a recent experiment the writer corroborated
the earlier evidence of cellulose breakdown, by measuring the loss in weight
of small pieces of filter paper attacked by the mycelium. The loss averaged
about 4 percent, and the mycelium was necessarily weighed with the paper.
The fibers were swollen and frayed at the ends. In spite of this it seems
most likely that the cellulose in a material such as manure is broken down
more slowly than some other constituents.

The surprising growth on peach gum and commercial pectin adds two
more classes of materials to the list of available foods. The results with the
hemicelluloses tested are not very conclusive. The lignin-bearing materials
produced the fastest growing cultures of the series. It is important that
this mushroom seems to belong with the lignin-destroying organisms,
although this must not be considered established by this very limited study.
According to Waksman (12) the decomposition of organic matter is usually
accompanied by the accumulation of lignin or substances of the same
nature, due to the inability of most organisms to attack it. The final
residue of such decomposition is of a humus or peaty nature. In support
of the theory that Agaricus campestris can break down and utilize lignin
it may be noted that when the spent manure from mushroom beds is applied
to the soil it soon disappears and does not seem to add to the humus of the
soil. It would seem that lignin had not accumulated in the mushroom bed.
The growth upon the proteins was not quite typical of the organic series.
It was dense and not dendritic. Strands did not appear in most cases until
the medium became exhausted. Whether considered from the point of
view of organic or of nitrogenous nutrition, it appears that the proteins are
very suitable nutrients. From the researches of Waksman (12) we learn
that a considerable change of nitrogen from urea and ammonium salts to
protein takes place when manure is composted. This process would
probably favor the nutritional preferences of the mushroom, and probably
takes place in the composting of manure for mushroom growing. The
ammonium salts and amino-acids served well as nitrogen sources, as was to
be expected, for proteins must be broken down into some of these compounds
before the organism can utilize them.

The occurrence of strands in certain plates has no special significance
with reference to nutrition, so far as the writer has been able to observe.
The appearance of crystals on the hyphae was noted in all the cultures.
More crystals were produced in some than others; the pectin cultures were
thickly covered with them. Hein (6) observed crystals which he stated
were calcium oxalate. Since more crystals appeared on the aerial hyphae
than on the others, and since some of these were soluble in water, it seems
possible that some other organic salt or acid was also produced. The pH
of the medium is invariably lowered by the spawn in commercial mushroom

STYER — MUSHROOM NUTRITION

993

Ak*

rl*

production, finally arriving at a value of about 4.5. The same effect was
produced in the cultures of Frear (5), with nutrient solutions on a paper
base. This pH change may have a profound influence upon the further
life of the mycelium; and it may possibly be prevented by chemical means
with great benefit to the mycelium. The further study of this matter is
being undertaken with hope of useful results.

It should now be possible to make silica gel and also larger scale cultures
upon many materials, including composts and fractions prepared from them,
and to correlate the results with analytical studies. There should result
many improvements in the handling of manures for mushroom production,
as well as better understanding of the principles involved. The micro-
biology of manures will most certainly be found of profound importance.
After finding that the organism can obtain nutrients from so many classes
of organic compounds, the writer sees no good reason why many other
materials can not replace manure in mushroom production.

Summary

1. The mycelium of the "Snow-White" variety of Agaricus campestris
does not tolerate a medium of nutrient salts and soluble organic substances,
in which the total concentration is much above 0.2 M.

2. The mycelium is intensely aerobic. In fibrous media it will not toler-
ate the addition of water if aeration is cut off thereby. Mycelial strands
develop profusely in the wetter media, and as their formation necessarily
precedes fruiting, a connection between aeration changes, mycelial strands,
and fruiting is suggested.

3. The spores of Agaricus campestris germinate more quickly in large
masses than when separated. The germination stimulus developed in the
spore mass appears to be the same as that by which the living mycelium
hastens germination.

4. The mycelium was grown on silica gel plates containing materials
representative of the main classes of substances in manure and decomposing
organic matter. Growth was good with xylose, dextrose, maltose, peach
gum, commercial pectin, wheat bran, wheat straw, Liriodendron wood, and
commercial granulated peat moss; and also on a sample of wood from
which nearly all but lignin had been removed by wood-rotting fungi.
Several proteins were excellent sources of nitrogen and organic food. Cel-
lulose failed to support vigorous growth. The organism probably can make
use of nearly all the substances in manure including lignin. Cellulose,
however, would probably not be among the first attacked.

5. The characteristic mycelial growth is dendritic with strand formation.
On the proteins the dense hyphal growth is uniforn in outline and few
strands are produced.

The guidance and assistance of Dr. Rodney H. True throughout the
progress of the work is acknowledged with the greatest pleasure. The

994

AMERICAN JOURNAL OF BOTANY

[Vol. 17,

writer has also been fortunate in receiving most useful suggestions, as well
as samples of some of the nutrients tested, from Dr. Selman A. Waksman.

Botanical Laboratory,

University of Pennsylvania

LITERATURE CITED

I . Bechmann, E. Untersuchungen uber die Kulturf ahigkeit des Champignons. Zeitschr.

Bot. 22: 289-323. 1929. . , , ,.„

2 Duggar, B. M. The principles of mushroom growmg and mushroom spawn makmg.

U. s' Dept. Agr. Bur. PI. Ind. Bull. 85, 1-60. 1905. .

3. Falck, Richard and Olga. Uber die Sporenkeimung des Champignons. Mykol.

Unter. und Ber. 1 : 1-60. 1924. . . r u t

4 Ferguson, Margaret C. A preliminary study of the germmation of the spores of

Agaricus campestris and other basidiomycetous fungi. U. S. Dept. Agr. Bur. 1 1.

Ind. Bull. 16, 1-43, 1902. . ^- „

5. Frear, D., J. F. Styer, and D. E. Haley. A study of the effect of H-ion concentration

on 'the growth of Agaricus campestris. Plant Physiol. 3: 91-94. 1928.

6. Hein, I. Studies on the mycelium of Psalliota campestris. Amer. Jour. Bot. 17:

197-211. 1930. C A •

7. Lambert, E. B. Production of normal sporophores in monosporous cultures of Agaricus

campestris. Mycologia 21: 333-335- 1929.
8 Platz, G. A., L. W. Durrell, and Mary F. Howe. The effect of carbon dioxide upon the
germination of the chlamydospores of Ustilago zeae (Beckm.) Ung. Jour. Agr.

Q Rippel*, A.; Ind h! Bortels! Vorlaufige Versuche uber die allgemeine Bedeutung der
Kohlensliure fur die Pflanzenzelle. (Versuche an Aspergillus niger). Biochem.

Zeitschr. 184: 237-244. 1927. . , , a

10. Styer, J. F. Preliminary study of the nutrition of the cultivated mushroom. Amer.

Jour. Bot. 15: 246-250. 1928.

II . A simplifted silica gel. Amer. Jour. Bot. 17: 636-637. 1930.

12! Waksman, S. A. The nature and origin of the soil organic matter or soil humus HI.

Soil Sci. 22: 323-333. 1926.

/

0 ^1

A synopsis of the marine algae

of Brazil

Wm RANDOLPH TAYLOR

INTRODUCTION

™,r; ^"'l ^^^ "? c' ^t\'''^^ knowledge as has been acquired of the
manne algae of South Amenca exists in the form of scattered original
hsts wh.ch resulted from the study and naming of plants from many
con.parat.vely small collections. In rare cases the incentive of I

Sitv tL "T ''^ """u "^, ^^™*"^ ^°™ --« p-^-'-

eTated to ?hi,'"l r?.' ^^^^^^^^^^o bring together the lists
Scihwi *'* d.str.ct. to consohdate them with his own. and so to
fac. htate greatly all subsequent study in this area. Brazil was thus
treated by Martens .n 1870 but in nearly 60 years algal classifiLttn
has undergone such rev.s.on that this list is largely obsolete. For South
Amenca we have a few modern compilations of this type. but. while
hey are of the utmost value, studies in algal distribution require that
these listed areas be jo.ned by similar surveys of the flora of the inter-
tr'Kltr'Sn* present the more notable synopses include studies of
the Falldand Islands by CoTTON. of the American Antarctic and the

H^r. "'"°'' S°"'^A™"ican shores by Gain, and of Peru by
HOWE Jeav.ng most of the mainland in a very unsatisfactory state.
When the algal collections of the « Hassler y> Expedition came

MARINE ALCAE OF BRAZIL

Wm. RANDOLPH TAYLOR

into the writer's hands for study several years ago, the need for a
consoHdation of the Hterature was urgently felt, and during the study
of Brazilian algae brought back by this expedition, by the « Alba-
tross » and by Dr. W. L. SCHMITT, an attempt at a new catalog
was made. It was decided to accept Martens* « Conspectus »
(1870) as a fair interpretation of the earlier records, and to this the
principal subsequent lists have been added. The fundamental diffi-
culty with all preliminary floras is the doubt which rests on many
of the early determinations, and this can only be removed by a res-
tudy of the original materials. As this was generally impracticable in
the present instance, the opinion of DE ToNi (Sylloge Algarum) was
accepted to give a working basis, and such changes made as seemed
necessary. Where a given species is one appropriate to the district
or repeatedly reported no question has, as a rule, been raised as to
the record. On the other hand where the identity of the plant is in
doubt because of inadequate description, uncertain synonymy, or
extraordinary and unconfirmed extension of range, a sign of interro-
gation has been prefixed. In some measure this a matter of personal
opinion, and often the matter is, in the absence of the original speci-
mens, quite debatable. However, as this is distinctly a preliminary
catalog or Prodromus, expressly for the purpose of attracting attention
to the limits of our present knowledge, such faults can readily be
excused now, and later recognized and rectified. Some genera, such
as Cladophora, Padina and Celidium are in very doubtful condition,
having been crowded with determinations of very uncertain value, and
an inspection of the names listed will give little help to a modern
student. In citing the original names cognizance has been taken of the
varying old custom of giving sometimes either the describer or the
author of the revised combination, but not both. Here the name is
followed by both authors as is now habitual, probably without intro-
ducing important errors. MARTENS and MoBlUS often do not give
definite stations for the plants in their lists, but most of their material
was collected by Dr. Glaziou near Rio Janeiro. Where a definite
station was available from some author, citations of « Brazil » alone by
others were omitted from the list of stations. Mention by name of the
various towns in the immediate vicinity of flio Janeiro was also not
usually attempted. It was difficult to locate some of the stations, but
with these exceptions the arrangement is approximately from upper

4»w

m» »

* I «

Brazil southwards, the unlocated stations following the others after a
semicolon.

In view of Setchell's suggestions upon the distribution of ma-
rine algae (Ann. Missouri Bot. Card.. 2 : 287-305 1915) it is
appropriate to iiiquire into the temperature range prevalent in the waters
along the Brazilian coast below the Equator. Detailed temperature re-
cords from shallow waters directly in the region of algal growth being
lacking. It is of some consequence to examine the records of off-shore

iqIt!' '^"' '*'"'" ** '^^"^ ^^^ Marine Observer, vol. 4.

IV^/) It appears that there is comparatively little variation along the
north coast below the Equator, the average monthly temperature ran-
ging from 26" C. to 29.3" C. generally about 27.5" C. always well
within the tropica! limits. The isotherm of 26.6 ' C (80" F ) shifts
along the coast from about 20" S. L. in February, to pass out of range

/7nu^^x '^-',"'''' '■^'"'■"'"g '■" September. The isotherm of 2 ! " C.
(70 F.) in February is at about 36"30' S. L., below the Rio de la
h^Iata. and in August-September is near C. St. Thome at 22" S. L
1 he water off Ilha Sao Francisco ranges from about 25.5" C in
summer to 17.7" C. in winter, and this station represents the southern-
most from which collections are available. By Setchell's criterion
of a summer temperature of 25" C. this would be near the boundary
between Tropical and Subtropical zones, but would probably support
a tropical flora and such seems to be the case. The upper Subtropical
sumnier temperature limit of 25" C. is found somewhat south of Ilha
Sao Francisco at about 28" S. L., and the lower limit of 20 C lies
north of C. Corrientes in northernmost Argentina. Since the shoal
water close to shore will generally be warmer than the off-shore water
It is to b^ expected that the boundaries will actually be south of these
points J he Subtropical zone, by Setchell's criteria, will therefore
probably be narrow, including a short stretch of Brazilian coast, Uru-
guay, and a little of Argentina. The northern Argentina coast will
otherwise lie in the South Temperate belt to 45" S. L.; below this
the Argentina-Patagonian coast to the northern shore of Tierra del
huego apparently lies in the Lower Austral, and the southern shore
and eastern end of Tierra del Fuego in the Upper Austral zone.

It becomes of immediate interest to determine the general charac-
ter of the Brazilian algal flora and the floristic source of its more
important elements. It is obvious that so far as we know it the charac-

IRREGULAR PAGINATION

MARINE ALCAE OF BRAZIL

Wm. RANDOLPH TAYLOR

ter is overwhelmingly tropical, its affinities, as might be expected,
connecting with the West Indies. A list of the more important species
will help to explain this. A selection was made of those which ranged
over a coast of at least 5° latitude, on the basis of repK>rts by at least
3 authors covering 3 or more well separated stations, so\ presumably
the common species, and it is significant that these are also almost
without exception important elements in the West Indian flora, and
apparently none represent a non-tropical element incursive from the
south :

Ertteromorpha compressa (L.) Grev.
Ulva fasciata Delile.
Ulva Laciuca (L.) Le Jolis.*
Chaiomorpha media (C. Ag.) Kg.
Anady^omene stellata (Wulf.) C. Ag.
Chamadoris Peniculum (Sol.) Ktze.
Dr\^posis permaia Lamx.
Caulerj>a crassifolia (C. Ag.) J. Ag.
Caulerpa prolifera (Forsk.) Lamx.
Caulerpa racemosa (Forsk.) J. Ag.
Caulerpa seriularioides (Gmel.) Howe
Codium tomentasum (Huds.) Stackh.
Halimeda Opuntia (L.) Lamx.

Dict^oia Bariayresi'i Lamx.*
Dict^ota dentata Lamx.
Dict}fota dichotoma (Huds.) Lamx.
Newocarpus delicatulus (Lamx.) Ktze
Neurocarpus Justii (Lamx.) Ktze.
Neurocarpus plagiogrammus (Mont.)

Ktze.
Padina VicI^ersicR Hoyt.*
Zonaria variegata (Lamx.) C. Ag.*
Zonaria zonalis (Lamx.) Howe.
Sargassum c}fmosum C. Ag.*
Sargassum vulgare C. Ag.

Calaxaura marg'mata (E. & S.) Lamx.
Calaxaura oblongata (E. & S.) Lamx.
Calaxaura rugasa (E. & S.) Lamx.
Celidium corneum (Huds.) Lamx.
Celidium rigidum (Vahl.) Grev.
Cracilaria cervicomis (Turn.) J. Ag.
Cracilaria confervoides (L.) Grev.
Cracilaria ferox J. Ag.
Cracilaria lacinulata (Vahl) Howe.
Hypnea musciformis (Wulf.) Lamx.
Hypnea spirtella (C. Ag.) Kg.
Chr];s}fmenia Uvaria (L.) J. Ag.
Acanthophora muscoides (L.) Bory.
Acanthophora spicifera (Vahl.) Borg.
Amansia muliifida Lamx.
Bry^othamnion Seaforthii (Turn.) Kg.
Br}foihamnion triquetrum (Gmel.)

Howe.
Lxiurencia obtttsa (Huds.) Lamx.
Laurencia papidosa (Forsk.) Grev.
Vidalia obtusiloba (C. Ag.) J. Ag.
Centroceras clavulatum (C. Ag.)

Mont.*
Cryptonemia crenulata J. Ag.
Cryfptonemia luxurians (Mert.) J. Ag.
Crateloupia filicina (Wulf.) C. Ag.
Amphiroa brasiliana Dene.
Corallina subulata E. & S.
Janla rubens (L.) Lamx.

<»^

rl •

•*>•

Of this group 3 are not recorded from much south of the Abro-
Ihos Ids. (18°S. L.) — Caulerpa crassifolia, C. prolifera, Vidalia
obtusiloba — but all the rest reach to Clabo Frio (23** S. L.), where

the coast turns westward toward Rio Janeiro. Similarly, 3 are not
recorded on the Brazilian coast north of Bahia (13° S. L.), but they
are all found in the West Indies. — Chtztomorpha media, Hypnecf
spinella, Acanthophora muscoides. A considerable incursion from the
south is not greatly to be expected. Coming westward from outside
of the Gulf of Guinea a broad belt of warm tropical water (the South
Tropical stream) divides off Cap San Roque, the north stream branch
driving up the north coast of Brazil past the Antilles into the Gulf
of Mexico while the southern branch turns down the eastern Bra-
zilian coast (Wiss. Ergebn. Schwedische SUdpolar Exped. 1901-
1903, Bd 4, I, fg. 6, map). This tends to equalize the coastal water
temperature and offsets the natural gradation from warmer water in
the north to cooler water in the south temperate belt. It also suggests
the possibility and a manner of the introduction of modern algal species
from African waters and the eastern Atlantic islands, though this
seems to have happened to a very trifling extent, if at all. However,
a drift of cold water from between Cape Horn and the South Shet-
land Islands comes about the Falkland Islands and up along the
coast, wedging between Patagonia and this warm tropical current
from the north, so that we must look to it as an explanation for condi-
tions favorable to any incursive south temperate floral elements. That
it can bring little direct to Brazil is to be inferred from the fact that
is greatly reduced opposite Uruguay and disappears off the southern-
most part of Brazil, Rio Grande and vicinity. Brazil shows the tro-
pical typ>e of algal flora dominant as far south as adequate collections
have extended, but we really know little of the flora south of 28" S. L.
Of course, since most active collecting has centered about Pernam-
buco, Bahia and Rio Janeiro it is not surprising that a sharp decrease
occurs in the lists of species from stations south of the last-named
place, with the dropping out of species certainly to be ex|>ected to
occur throughout Brazil and which will probably appear in subsequent
more complete collections. In the list given above the species starred(*)
go to the latitude of the Ilha Sao Francisco neighbourhood at 27°S.L.
Since this area represents the southernmost that has received any
considerable amount of collecting it is worth while to note the species
reported from it. It will be seen that the flora is still very strikingly
tropical, although including a number of cosmopolitan species of wide
warm-temperate as well as tropical distribution :

MARINE ALCAE OF BRAZIL

Calaxaura marginata (E. & S. ) Lamx.
Calaxaura oblongata (E. &S.) Lamx.
Catendla impudica (Mont.) J. Ag.
Cracilaria ferox J. Ag.
Hypnea musciformis (Wulf.) Lamx.
Hypnea spinella (C. Ag.) Kg.
Acanthophora muscoides (L.) Bory,
Amphibia radicans (Mont.) Ktze.
Laurencia papillose (Forsk.) Grev.
Centroceras clavulatum (C. Ag.)

Mont.
Criffithsia radicans Kg.

Enter omorpha Linza (L.) J. Ag.
Viva Lactuca (L.) Le Jolis.
Cladophora fascicularis (Mert.) Kg.
Caulerpa racemosa (Forsk.) J. Ag.
Gaulerpa sertularioides (Gmel.) Howe

Sphacelaria tribuloides Menegh.
Colpomenia sinuosa (Roth.) D. & S.
Hydroclathrus clathratus (Bory)

Howe.
Dictyota Bartayresii Lamx.
Dictyota cervicornis Kg.
Neurocarpus delicatulus (Lamx.) Ktze.
Padina Viclfersics Hoyt.
Zonaria variegata (Lamx.) C. Ag.
Sargassum cymosum C. Ag.

Finally, it is necessary to consider what may be the importance
of the numerous doubtful species in relation to temperate floral ele-
ments from the south or elsewhere. Many are species only well known
from the Cape of Good Hope region; others belong to the Pacific
coast ; others are European species. Among these there is little encou-
ragement for the citation of a temperate element, because the proba-
bility that the specimens have been incorrectly named is great, and
it is likely that a review of the materials on which tiiese records were»
made will result in the renaming of most of them as species in accord
with the general character of the flora. Among the plants first (and
inadequately) described from Brazil there may well be incursives
from the little-known temperate flora to the south, but since such
records mostly come from the better known stations within the truly
tropical area they probably do not represent a very important tempe-
rate element, and most of them will probably turn out to be plants
familiar under other names. However, in this connection an inspection
of the types of most of the plants which Greville described as new
from St. Hilaire's Brazilian collections showed (HoWE & Taylor
1930) that almost all had rightly been considered novelties, and that
they were not plants better known under other names. In spite of
all this, it is to be expected that there will eventually be recognized
a small number of plants representing members of both the north and
the south temperate floras which have succeeded in establishing them-
selves along the Brazilian coast.

Wm. RANDOLPH TAYLOR

With the fact in mind that algae of deep water are under more
equable conditions and exposed to lower maximum temperatures than
those of the littoral, the lists of species collected by dredging off
Pernambuco, the Abrolhos and Cabo Frio are worthy of inspection :

PERNAMBUCO 20 Fa., 8' S. L.

Anadyomene stellata (Wulf.) C. Ag.
Cham&doris Peniculum (Sol.) Ktze.
Cauhrpa Ashmeadii Harv.
Caulerpa lanuginosa J. Ag.

Dictyota dentata Lamx.

Zonaria variegata (Lamx.) C. Ag.

Zonaria zonalis (Lamx.) Howe.

Bryothamnium triquetrum (Gmel.)

Howe.
Chondria floridana (Collins) Howe.

ABROLHOS IDS 18" S. L.

Anadyomene stellata (Wulf.) C. Ag.
Cham&dedoris Peniculum (Sol.) Ktze.
Valonia ventricosa J. Ag.
Caulerpa crassifoHa mexicana (Sond.)

J. Ag.
Halimeda Tuna (E. &S.) Lamx.
Udotea cyathiformis Dene.

Sporochnus Bolleanus Mont.
Neurocarpus Justii (Lamx.) Ktze.
Padina Sanctce-Crucis Borg.
Zonaria variegata (Lamx.) C. Ag.

Cracilaria conjervoides (L.) Grev.
Cracilaria mamillaris Howe.
Bryothamnion Seaforthii (Turn.) Kg.
Dasya sertularioides Howe & Taylor.
Spyridia filamentosu (Wulf.) Harv.
Halymenia Floresii (Clem.) Ag.

CABO FRIO 23" S. L.

Dasycladus vermicularis (Scop.)

Krasser.
Dictyosphceria favulosa (Ag.) Dene.
Chamcsdoris Peniculum (Sol.) Ktze.
Codium decorticaium (Woodw.)

Howe.
Halimeda Tuna (E. &S. ) Lamx.
Rhipilia tomentosa Kg.

Sporochnus Bolleanus Mont.
Sporochnus pedunculatus (Huds.) C.

Ag.

Dictyoia Bartayresii Lamx.
Dictyota cervicornis Kg.
Dictyota dentata Lamx.
Dictyota divaricata Lamx.
Neurocarpus Justii (Lamx.) Ktze.
Zonaria variegata (Lamx.) C. Ag.
Zonaria zonalis (Lamx.) Howe.

Caaxaura cylindrica (Sol.) Lamx.
Celidium rigidum (Vahl.) Grev.
Callophyllis microdonta (Grev.) Falk.
Wurd^mannia setacea Harv.
Rhodophyllis gracilarioides Howe &

Taylor.
Cracilaria mamillaris Howe.
Chrysymenia Enteromorpha Harv.
Chrysymenia planifrons (Melv.) J.

Ag.
Chrysymenia pyriformis Borg.
Fauchea Hassleri Howe & Taylor.
Nitophyllum odontophorum Howe &

Taylor.
Nitophyllum uncinatum (Turn.) Grev.
Laurencia lata Howe & Taylor.
Hakymenia Integra Howe & Taylor.
Halymenia rosea Howe & Taylor.
Halymenia vmacea Howe & Taylor.
Platoma tenuis Howe & Taylor.

8

MARINE ALCAE OF BRAZIL

It will be seen from the first locality that all are common West
Indian forms : to be sure, the list is short and unquestionably incom-
plete as representing the deep water flora. Off the Abrolhos Ids. —
depth unknown — the short list includes common West Indian forms
only, with one exception. The Cabo Frio collection is much the largest,
and approaches most nearly the south Subtropical zone. The flora
is predominantly tropical, of West Indian affinities still, except for
Callophyllis, and new species of Rhodoph^llis, Fauchea, Haly-
menia and Platoma. Perhaps these are forms that will appear in
the south Subtropical or Temperate floras when these are better
known. At any rate there is more of novel character here than was
collected in the Ilha Sao Francisco neighbourhood a few degrees of
latitude to the south, but primarily from the littoral vegetation, and
so exposed to seasonally higher water temperatures.

CATALOG OF SPECIES RECORDED

MYXOPHYCEi^

It IS certain that even the few records which exist of marine Myxophyceae
from Brazil are of very doubtful value, and therefore no attempt to reduce
the names to modern usage is made here, or to organize the report as in the
other groups of algae. The lists include the following : JInahaena variabilis
Kg., Cabo Frio, MoBius (1889); Calothrix aeruginea (Kg) Thuret,
MoBius (1890); Calothrix ptlosa Harv., MoBius (1890) Chroococcus
iurgidus (Kg.) Nag., De Toni (Sylloge Algarum 5; 1907); Leibleinia
aequalis Kg., Martens (1870); L. .seriala Kg., Pemambuco, Piccone
(1889); Lyngbya aesluar ii Qur^.) Liebm., MoBius (1890); L. confer--
voides C. Ag. ex Com., Conceicao de Itanhaen, Taylor (1930 b) ; L. no-
ronhae Dickie, Fernando Noronha, DiCKiE (1875); L. semiplena J. Ag.
ex Gom., De Toni (Sylloge Algarum 5; 1907) iMicrocoUus chthonoplastes

Wm. RANDOLPH TAYLOR

(Fl. Dan.) Thuret, MoBlus (1980): Scxtonema ocellatum Lyngb., De
Toni (Sylloge Algarum 5; 1907) ; Trichodesmium Thiebautii Com., Bahia,
Taylor (1930).

CHLOROPHYCE/E

PROTOCOCCACEiE

?Protococcus affinis Dickie. — Dickie (1884 b). St. Paul Rocks.

ULOTRICHACEiE

?Hormospora pellucida Dickie. — Dickie (1875). Fernando Noro-
nha Ids.

ULVACEiE

Enteromorpha clathrata (Roth) Grev. — Martens (1870); £. da-
thrata (Roth) J. Ag., MoBius (1890). Saguarema, near Rio Janeiro.

Enteromorpha compressa (L.) Grev. — MbBius (1890); Schmidt
(as fa. lingulata Hauck) (1924); ? E. compressa Link and ? E. complanata
Kg., Martens (1870). Mucuripe, Cabo Branco, Macahe, Rio Janeiro, Ja-
quaranga.

Enteromorpha erecta (Lyngh.) J. Ag. — Schmidt (1924). Mucuripe,
Jaquaranga.

Enteromorpha intestinalis (L.) Link. — Martens (1870), Zeller
(1876); Martens (as fa. capillaris and fa. tubulosa Kg.) (1871). Rio
Janeiro.

Enteromorpha lingulata J. Ag. — Taylor (1930, 1930 b). Rio
Janeiro, Conceicao de Itanhaen, Guaruja.

Enteromorpha Lima (L.) J. Ag. — Schmidt (1924); Phycoseris
Linza L., Martens (1870). Jaquaranga, Itapacorohy.

?Enteromorpha minima Nag. — De Toni (Sylloge Algarum 1 : 125).
St. Paul Rocks.

Enteromorpha prolifera (O. F. Muller) J. Ag. — Howe (1928).
Bahia, Nictheroy, Rio Janeiro.

?Monostroma obscurum(Kg.) J. Ag. — Ulva obscura Kg., Martens
(1870). Rio Janeiro.

10

MARINE ALCAE OF BRAZIL

U Iv a Tasciata DelWe. — Taylor (1930, 1930 b); Phyfcoseris fasciata
Delile Martens (1870, 1871) ; P. lobata Kg., Zeller (1876) ; P. nema^
(oidea (Bory) Mont., Grunow (1870); Ulva fasciata lobata (Kg.), PlC-
CONE (1886); ?U. lobata Kg., DiCKIE (1875). Fernando Noronha, Ma-
cahe, Nictheroy, Rio Janeiro, Sao Vicente, Conceicao de Itanhaen, Guaruja.

Ulva Lactuca (L.) Le Jolis. — Martens (1870), Mobius (1889,
1890, 1892, 1895), Howe (1928). Bahia, Rio Janeiro, Ponta da Copaca-
bana, Paqueta, Itajahy.

?tilva Lactuca fa. laciniata (Wulf.) J. Ag. — Phyco&eris australis
Kg., Martens (1870). Rio Janeiro.

Ulva Lacluca\air. latissima (L.) D. C. — Taylor (1930); U. la-
tissima L., Martens (1870). Rio Janeiro.

Ulva Lactuca var. rigida (C. Ag.) Le Jolis. — Taylor (1930);
U. Lactuca fa. r'lg'ida (Kg.) Le Jolis, ScHMIDT (1924) ; Phycoseris rigida Kg.,

Martens (1870); Ulva rigida C. Ag., Howe (1928). Mucuripe, Cabo

Branco, Bahia, Nictheroy, Rio Janeiro, Jacquaranga, Ilha Sao Sebastiao, Ilha
Sao Francisco.

?Ulva plicata MuIIer. — ?Phycoseris plicata Kg., Martens (1871).
Rio Janeiro.

PRASIOLACEit

? Prasiola minuta Dickie. — Dickie (1884 b). St. Paul Rocks.

CHAETOPHORACEiE

Endoderma minuta (Reinke) Lagerh. —
MoBlus (1889. 1890). Near Rio Janeiro.

CLADOPHORACEi*

Entocladia viridis Reinke,

?Cladophoropsis,

?Aega^rophila membranacea caespilosa C. Ag. -
Martens (1870). Pemambuco.

Chaetomorpha claoata (C. Ag.) Kg. — Martens (1870, 1871).
Rio Janeiro.

Chaetomorpha gracilis Kg. — MoBius (1890). Near Rio Janeiro.

Chaetomorpha Linum (Miill.) Kg. — C. chlorotica Kg., Mobius
(1890). Near Rio Janeiro.

Wm. RANDOLPH TAYLOR

II

Chaetomorpha Ltnum fa. aerea (Dillw.) Collins. — C. aerca Dillw.,
Dickie (1875); C. Dubyana Kg., Piccone (1889). Pemambuco. Barro
Grande.

Chaetomorpha media (C Ag.) Kg. — Mobius (1889, 1890), Howe
(1928), Taylor (1930); C. antermina Kg., Martens (1871), Zeller

(1876) ; ?C. media C. Ag., MarTENS (1870). Bahia, Macahe, Rio Janeiro,
Ponta da Copacabana, Sao Vicente, Conceicao de Itanhaen.

?Chaeiomorpha pachynema Mont. — Zeller (1876). Rio Janeiro.

?Chaetomorpha saccata Kg. — Martens (1876). Jacuhy near
Porto Allegre.

?Cladophora alhida (Huds.) Kg. — MbBlus (1889). Near Rio
Janeiro.

Cladophora brasiliana Mart. — Martens (1870), Mobius (1889,
1892). Rio Janeiro, Lagoa Rodrigues, Lagoa de Rodrigo de Frietas.

?Cladophora cornea Kg. — Mobius (1892). Rio Janeiro.

?Cladophora Echinus (Bias.) Kg.— MoBlus (1889). Ponta da Co-
pacabana.

Cladophora fascicularis (Men.) Kg. — Martens (1870, 1871),
Schmidt (1924). Taylor (1930). Mucuripe, Cabo Frio, Jaquaranga, Ilha
Sao Sebastiao.

Cladophora heteronema (C. Ag.) Kg. — BoRGEsen (1913). « Bra-

zil

».

?Cladophora Bilarit Grev. — Martens (1870). Aldea Vclha.

?Cladophora minuta Dickie. — Dickie (1875). Fernando Noronha.

Cladophora nilida Kg. — ?C. trichotoma Kg., MoBius (1890). Near
Rio Janeiro.

Cladophora pellucida Kg. — Martens (1870). Macahe; Villa Boa.

Cladophora prolifera (Roth) Kg. — Martens (1871), Mobius
(1890); C. fruticulosa Kg., Martens (1871); C. catenata L., Martens
(1870). Macahe, Rio Janeiro.

ICladophora subvaricosa Diche. — Dickie (1875). Fernando No-

ronha.

Cladophora utriculosa Kg. — Piccone (1886). Rio Janeiro.

12

MARINE ALCAE OF BRAZIL

Microdiciyon BorgeseniiSeicheW. — M. umbilicatum (Velley) Zan.,
Gepp (1905). Dredged between Bahia and Rio Janeiro.

?Microdictyoncahdiclyon Mont. — Dickie (1875). Barro Grande
near Pernambuco.

DASYCLADACEiE

Jlcicularia Schenckiii^'o^) Solms. — MoBius (1889). Cabo Frio.
Dasycladusvermicularis (Scop.) Krasser. — Taylor (1930). Cabo

Frio.

VALONIACEi^

Anadyomene stellata (Wulf.) C. Ag. — Martens (1870), Schmidt
(1924), Howe (1924), Taylor (1930). Mucuripe, Pernambuco, Abrolhos
Ids., Bahia, Jaquaranga.

Chamaedoris Peniculum (Sol.) Ktze. — Schmidt (1924), Taylor
(1930); C. annulata Mont., Martens (1870), DiCKIE (1875), ZellER
(1876). Cabo Branco, Pernambuco, Barro Grande, Abrolhos Ids., Cabo Frio,
Rio Janeiro.

Dictyosphaeria favulosa (Ag.) Dene. — Taylor (1930). Cabo
Frio.

Ernodesmisverliciliata (Kg.) Borg. — Schmidt (1924); Valonia
verticillata Kg., Martens (1871), var. major Mart., Martens (1870).
Mucuripe, Rio Janeiro.

Valonia AegagrophilaC. Ag. — Schmidt (1924). Mucuripe, Jaqua-
ranga.

Valonia ventricosa J. Ag. — Taylor (1930). Abrolhos Ids.

BRYOPSIDACEi*;

?Bryopsis caespitosa Suhr. — Zeller (1876). Rio Janeiro.

Bryopsis pennata Lamx. — Gepp (1905). St. Paul's Rocks.

Bryopsis penndfa var. secunda (Harv.) Collins. — B. Harveyana
J. Ag., Howe (1928) ; B. Leprieurii Kg., Zeller (1876) ; B. plumosa var.
Leprieurii (Kg.) Borg.. ScHMIDT (1924). Mucuripe. Bahia. Rio Janeiro.

Bryopsis plumosa (Huds.) C. Ag. — MoBius (1890). Near Rio
Janeiro.

Wm. RANDOLPH TAYLOR

13

Bryopsis ramulosa Mont. — Murray (1889). « Brazil >.
?Bryopsis spinescens Zeller. — Zeller (1876). Rio Janeiro.

CAULERPACEiE

Caulerpa Ashmeadii Harvey. — Taylor (1930). Pernambuco.

Caulerpa crassifolia fa. mexicana (Sender) J. Ag. — Taylor
(1930) ; C. mexicana Sender. DiCKIE (1875) ; C. p'mnaia mexicana Weber-
V. Bosse, Schmidt (1924). Fernando Norohna Ids., Cabo Branco, Abrolhos
Ids.

Caulerpa crassifolia isi. pechnata (Kg.) Collins. — Schmidt (1924)
Cabo Branco.

Caulerpa fastigiata Mont. — Howe (1928); Herpochaete fastigiata
Mont., Martens (1870). Nictheroy, Rio Janeiro.

Caulerpa lanuginosa J. Ag. — Taylor (1930). Pernambuco.

Caulerpa Murray i^cher-y. Bosse. — Gepp (1905). Dredged bet-
ween Bahia and Rio Janeiro.

Caulerpa paspaloides (Bory) Grev. — Chauvinia paspaloides Bory,

Martens (1870). « Brazil ».

Caulerpa paspaloides var. phleoides (Bory) J. Ag. — ChauVmia
phleoides BORY. MARTENS (1870). « Brazil ».

Caulerpa peltata (Turn.) Lamx. — Chauvmia peltata Kg., MarTENS
(1870, 1871). Rio Janeiro.

Caulerpa prolif era (Forsk.) Lamx. — Dickie (1875). Piccone

(1889). Schmidt (1924). Mucuripe, Barro Grande, Abrolhos Ids.

Caulerpa pusilla (Kg.) J. Ag. — Siephanocoelmm pusillum Kg.,
Martens (1870). Pernambuco.

Caulerpa racemosa(FoYsk.) J. Ag.— C. Chemnilzia (Esp.) Lamx.,
Piccone (1886, 1889); Chauvinia Chemnitzia Kg., Martens (1870).
Pernambuco, Bahia.

Caulerpa racemosa var. clavifera (Turn.) Weber-v. Bosse. —
Schmidt (1924), Taylor (1930); C. clavifera C. Ag., Dickie (1875);
?C. clavifera Kg., /a., MoBIUS (1890). Fernando Noronha Ids.. Mucuripe.
Barro Grande, Nictheroy, near Rio Janeiro.

14

MARINE ALCAE OF BRAZIL

Caulerpa racemosa var. laefevirens (Mont.) Weber-v. Bosse —
Gepp (1905), Taylor (1930). St. Paul's Rocks, Ilha Sao Sebastiao.

Caulerpa racemosa var. Lamourouxii (Turn.) Weber-v. Bosse —
Chauvinia clcwifera Lamourouxii Kg., Martens (1870). Pernambuco, Rio
Janeiro.

nc^S^'i^''^" ''"^^""''^ ^"- occidentaltsQ. Ag.) B6rg. — Schmidt

(1924), Taylor (1930). Cabo Branco, Scarahy.

„r.Sf"'^''P'' '■'''^«'"^*'' ^^i-- UviferaiTnxu.) Weber-v. Bosse. — Gepp
(1905). Schmidt (1924), Taylor (1930). Between Bahia and R.o Janeiro
Jaquaranga, Ilha Sao Sebastiao.

^Caulerpa sedoides (R. Br.) C. Ag. - Chauvmia scdoide. Kg..
Martens (1870). Bahia. ^

(M^il^^t'I^V^!^^" ^- ^^- ~ ^^°"^''"''' ^'^"^" Bo^y. Martens
(lo/O). Aldea Velha.

Caulerpa sertularioides (Gmel.) Howe.— Taylor (1930) • C plu-
mosa (Forsk.) C. Ag.. ScHMIDT (1924). Jaquaranga. Ilha Sao Sebist.ao.

Caulerpa sertularioides i^.longiseta(]. Ag.) Sved. — ?C. phmaris
var. Bor^ana Kg.. Martens (1870). Maranhao.

Caulerpa verticillata j. Ag. - Howe Bahama Flora. 1 920

« Drazil ».

Cattlerpa Wehbiana Mont. — Dickie (1875). St. Paul's Rocks.

hernando Noronha Ids.

CODUCE/E

Avrainvillea nigricans Dene. — Borgesen (1913). « Brazil ».
,,^,^f'^J!'"'/^^<"'f'^''f"'n('^ood^) Howe. — Howe (1924), Taylor
r D- T ''""«"'"'" ^^- Martens (1870. 1871). Mobius (1890) Cabo
^no, R,o Janeno. Juparayha, Scarahy. Ilha Paqueta. Ilha Sao Sebast.ao. Ilha
oao rrancisco.

Codium intertextum Collins & Hervey. — Howe (1924) • C adhae-
rens (Cabr.) Ag.. Martens (1870), Mobius (1890). Schmidt (1924)
Bahia. Jaquaranga, Rio Janeiro.

?Codium lineare C. Ag. _ De Toni (Sylloge Algarum 1 :495).
« Drazil ». o ^/.

Wm. RANDOLPH TAYLOR

15

?Codium Pilgeri O. C. Schmidt. — Taylor (1930). Ilha Sao Se-

bastiao.

Codium tomentosum (hiuds.) Stackh. — Martens (incl. C. decum-
bens Mart, and vars. of C. (omentosum). (1870, 1871), DiCKlE (1875),
MoBius (1890), Gepp (1905), Schmidt (1924). C:\bo Branco, Barro

Grande, Bahia, Rio Janeiro, Jaquaranga. Probably in part best referable else-
where.

HaiimedaOpuntiaiL.) Lamx.— Martens (1870), Dickie (1875).
Zeller (1876), Mobius (1889), Piccone (1889), Howe (1928). Fer-
nando Norohna Ids., Barro Grande, Bahia, Abrolhos Ids., Rio Janeiro.

Balimeda Opunfia var. trilola (Dene.) Bart. — Schmidt (1924).

Cabo Branco.

Tialimeda Tuna (E. & S.) Lamx. — Dickie (1875), Taylor

(1930). Barro Grande, Abrolhos Ids., Cabo Frio.

J{hipilia tomentosa Kg. — Taylor (1930). Cabo Frio.
Udotea congluHnata Umx. — Dickie (1875). Barro Grande.
Udotea cyathiformis Dene. — Taylor (1930). Abrolhos Ids.
Udotea flabellum (E. & S.) Howe — Schmidt (1924). Cabo

Branco.

Udotea occidentalis Gepp. — U. Halimeda Kg., Martens (1870),
Dickie (1875). Bahia.

PHAEOPHYCE/E

SPHACELARlACEiC

Cladostephus verticillatus (UghiL) C. Ag.— C. m}frioph\fUum Ag..
Martens (1870). Cabo Frio.

Sphacelaria hrachygonia Mont. — Martens (1870), Sauvageau
(1900-04). Sta. Catherina.

Sphacelaria trihuloides Menegh. — Mbeius (1892), Schmidt
(1924). Rio Janeiro, Jaquaranga.

ECTOCARPACEit

Ectocarpus confervoides (Roth) Le Jolis. — Schmidt (1924). Ja-
quaranga.

16

MARINE ALCAE OF BRAZIL

Wm. RANDOLPH TAYLOR

17

.^^^^^^^^'^''P^^ f^'^^^^^^^^i^s yar. refractus (Kg.) Ardis«. — Schmidt
(1924). Jaquaranga.

?Ectocarpus Glaziovii ZtWer. — Zeller (1876). Rio Janeiro.
Ectocarpus minutuius Mont. — Zeller (1876). Rio Janeiro.
Ectocarpus siliculosus (DiWw.) Lyngb. — Martens (1870); ?E.
confervoides var. siliculosus Hauck. MoBIUs (1890). Near Rio Janeiro.'

Pylaiella fulvescens (Schousb.) Bomet. — Schmidt (1924) Jaqua-
ranga. Howe (Bahama Flora 1920) suggests that this may be a synonym of
P. Antillarum (Grun.) De Toni.

Pylaiella linoralis (L.) Kjellm. — ? Ectocarpus iittoralis (Dillw.)
Harv. var. brasiliensis Grunow (1870). Rio Janeiro.

CHORD ARUCEit

?Mesogloia brasiliensis Mont.— DeToni (Sylloge Algarum 3 : 427).
« Brazil ».

ASPEROCOCCACEiE

Chnoospora fasUgiafa y^r. atlantica ]. Ag. — Dickie (1884 b)
M6BIUS (1889). St. Paul's Rocks; Ponta da Copacabana.

Colpomenia sinuosa (Roth) D. & S. — Howe (1928), Taylor
(1930) ; Hydroclathrus sinuosus (Roth) Zanard.. MoBIus (1892). Nictheroy,
Rio Janeiro, liha Sao Sebastiao.

Bydroclathrus clathratus {Boxy) Howe. — Taylor (1930)- En-
coelium ciaihratum C. Ag., Martens (1870); Hy^droclaihrus cancdlatv
Bory, Martens (1871), MoBIUS (1890. 1892). Rio Janeiro, Scarahy, Ilh
oao oebastiao.

Tiosenvingea intricafa (J. Ag.) Bbrg. - Encoelium inlrlcaium Uebm..
Martens (1870) ; Asperococcus intricaius J. Ag.. Dickie (1875). Fernando
Noronha Ids., Bahia.

SPOROCHNACEiE

SporochnusBolleanusMoni. — Taylor (1930). Abrolhos Ids., Cabo

us
a

Frio.

?Sporochnus pedunculafusiHuds.) C. Ag.— Taylor (1930), Cabo
Frio.

s >

«'■ "

LITHODERMATACEi^E

Lifhoderma fatiscens Aresch. — Mobius (1890). « Brazil ».

DICTYOTACEi^:

Dictyota Bar lay resit Lamx. — Martens (1870), Zeller (1876),
Dickie (1875), Mobius (1889). Howe (1928), Taylor-? (1930). Ma-
ranhao, Fernando Noronha Ids., Aldea Velha, Cabo Frio. Nictheroy, Ilha
Gamboa.

Bictyota cervicornis K%. — Zeller (1876). Taylor (1930); D.
Fasciola Lamx., Martens (1870). Maranhao, Cabo Frio, Rio Janeiro, Ilha
Sao Sebastiao.

Biclyota ciliolafa Kg. — D. ciliata J. Ag., Dickie (1875), Mobius

(1890); D. crenulata J. Ag., Martens (1871). Fernando Noronha Ids.,
near Rio Janeiro.

?Dictyota cuneata Dickie. — Dickie (1884 b). St. Paul's Rocks.

Dictyota dentata Lamx. — Martens (1870), Dickie (1875),
Schmidt (including forms) (1924), Howe (1928), Taylor (1930);
D. Mertensii Kg. and D. Brongrtiartii J. Ag. Martens (1870) ; D. pirma-
tifida Kg., Schmidt (1924). Mucuripe, Cabo Branco. Pernambuco, Barro
Grande, Bahia, Cabo Frio, Jaquaranga.

Dictyofa dichotoma (Huds.) Lamx. — Martens (1870), Grunow
(1870), Dickie (1875), Mobius (1890), Gepp (1905), Schmidt (1924).
Maranhao, Cabo Branco, Barro Grande, Bahia. Rio Janeiro. Probably in part
best referable elsewhere.

Dictyota divaricata^mx. — Taylor (1930); ?D. Barla'^resii var.
divaricata J. Ag.. Martens (1870). Bahia, Cabo Frio.

T)ictyota indica Sonder. — Martens 1870. Rio, in Praia Grande.

Dictyota pardalis Kg. — Schmidt (1924). Mucuripe.

Dilophus allernans J. Ag. — Taylor (1930). Ilha Govenador, near
Rio Janeiro.

?Dilophus gutneensis (Kg.) J. Ag. — Howe (1928); ?Diciyola
Antigua Sonder. Zeller (1876). Bahia, a fragment only; Rio Janeiro.

?Gymnosorus collaris (C. Ag.) J. Ag. — Zonaria collaris C. Ag..
Martens (1870).

18

MARINE ALCAE OF BRAZIL

?(J\eurocarpus) Hatiseris Areschougii J. Ag. — J. Agardh (Sp..
Gen. & Ord. Alg. 1 : 119.1848). Pernambuco.

JMeurocarpus delicatulus (Lamx.) Ktze — Taylor (1 930. 1 930 b) ;
Haliseris deUcaiulus (Lamx.) J. Ag.. Martens (1870). DiCKiE (1875)'
PiCCONE (1886. 1889); Dictyopteris delicatulus Lamx.. SCHMIDT (1924)!
D. Hauckianus Mob.. MoBlus (1889); Neurocarpus Hauckianus (Mob.)
Ktze., Howe (1928). Fernando Noronha Ids., Cabo Branco, Olinda, Per-
nambuco, Bahia. Nictheroy. Rio Janeiro, Ilha Sao Sebastiao. Guaruja.

Meurocarpus Justii (Lamx.) Ktze. — Taylor (1930); Haliseris
Jusiii Ag.. Martens (1870); Dici^opteris Jusdi Lamx., Schmidt (1924).

Cabo Branco, Pernambuco, Bahia, Abrolhos Ids., Cabo Frio, Rio Janeiro.

?Meurocarpus membranaceus (Stackh.) Ktze. — De Toni (Sylloge
Algarum 3 : 254) ; Haliseris pol-^podioides (Desf.) Lamx., J. Agardh (Sp..
Gen. & Ord. Alg 1 : 1 1 7.1848). « Brazil ».

J^eurocarpus plagiogrammus (Mont.) Ktze.— Howe (1928); Hali-
seris plagiogrammus Mont., Martens (1870) ; DiCKIE (1875) ; Dictyopteris
plagiogramma Mont., MoBlus (1889). Fernando Noronha Ids., Barro Grande,
Bahia, Rio Janeiro.

?Padma Commersonii Bory. — Schmidt (1924). Mucuripe. Cabo

Branco. Jaquaranga.

?PaJina Durvillaei (Bory) Harv., fa. obscura Piccone. — PiccoNE
(1886. 1889). Rio Janeiro.

?Padina Fraseri (Grev.) J. Ag. — Zonaria Fraseri Grev., Martens
(1870). Bahia.

PaJina gymnospora (Kg.) Vickers. — Schmidt (1924), Zonaria
gVmnospora Kg.. Martens (1870), Zeller (1876). Rio Janeiro, Jaqua-
ranga.

?Padina Pavonia Lamx. — ?P. Pavonia Gail!., DiCKiE (1875);
Zonaria Pavonia Ag.. Martens (1870) ; Z. tenuis Mont.. Martens (1870).'
Zeller (1876). Fernando Noronha Ids.. Bahia. Rio Janeiro. Probably refe-
rable elsewhere.

Padina Sanctae-Crucis Borg. — Howe (1928), Taylor (1930).
Abrolhos Ids.. Bahia.

PadinaVickersiaeHoyt. — Howe (1928), Taylor (1930)- P
variegata (Kg.) Hauck, MoBlus (1889, 1895) ; P, Howeana Borg.. Schmidt

Wm. RANDOLPH TAYLOR

19

(1924). Cabo Branco. Rio Janeiro. Jaquaranga, Ilha Sao Sebastiao, Ilha
Gamboa.

Spatoglostum Areschougii J. Ag. — S. Schroederi Kg.. Martens
(1870). Piccone (1886). Pernambuco.

Spatoglossum Schroederi (Mert.) J. Ag. — Schmidt (1924). Cabo
Branco.

Zonaria flava (Clem.) Ag. — Sfypopodium flavum Kg., Martens
(1870). Macahe.

Zonaria variegata (Lamx.) C. Ag. — Howe (1928). Taylori

(1930); Z. variegata C. Ag., Martens (with var. discolor) (1870),
Piccone (1889); Cy^mncsorus variegatus J. Ag., Gepp (1905); Zonaria
variegata Lamx., DiCKIE (1875); Z. variegata (Lamx.) Mart., MoBIUS
(1889), Schmidt (1924). Mucuripe, Olinda, Pernambuco, Barro Grande.
Bahia. Abrolhos Ids., Cabo Frio, Rio Janeiro, Jaquaranga, Itapacorohy.

Zonaria zonalis (Lamx.) Howe. — Taylor (1930); Z. lohata
(C. Ag.) J. Ag., Dickie (1875), Schmidt (1924); Sty^popodium fuUgi-

nosum Kg., & Spatoglossum versicolor Kg., MarTENS (1870); Stypopodium
lohatum Kg., Gepp (1905). Cabo Branco, Pernambuco. Barro Grande. Cabo
Fno. Rio Janeiro.

FUCACEi«

?Jlscophytlum nodosum (L.) Le Jolis. — De Toni (Sylloge Alga-
rum 3 : 210). « Brazil ».

?Tucus vesiculosus L. — De Toni (Sylloge Algarum 3 : 206).

« Mari Brasiliense ».

?Sargassum Chamissonis K^. — Zeller (1876). Rio Janeiro.

Sargassum cymosum C. Ag. — Martens (1870. 1871), Zelleji
(1876), Piccone (1886, 1889), Mobius (1889, 1890, 1892). Howe
(1928). Taylor (1930, 1930 6); 5. c}fmosum var. stenophyllum (Mert.)
Grun., Schmidt (1924); 5. chierifoUum Kg., MoBIUS (1889); 5. Esperi
C. Ag., Piccone (1886), Mobius (1889) ; 5. integri folium Kg., Martens
(1870). Piccone (1886); 5. ramifolium Kg., Martens (1870), Zeller
(1876) ; 5. rigidulum Kg., MARTENS (1870,1871), MoBIUs (1889, 1890).
Schmidt (1924). Mucuripe, Cabo Branco, Pernambuco, Bahia, Abrolhos
Ids., Nictheroy, Rio Janeiro, Scarahy, Copacabana, Jaquaranga. Ilha Grande,
Ilha Sao Sebastiao. Guaruja. Ilha Gamboa.

20

MARINE ALCAE OF BRAZIL

Sargassum Tilipendula C.Ag.— Howe (1928), Taylor (1930);
S. athe C.Ag., Zeller (1876). Rio Janeiro, Copacabana.

Sargassum lendigerum (L.) C.Ag. — Martens (1870), Grunow
(1870). Rio Janeiro.

1 Sargassum Liebmannii ] . Ag. — Piccone (1886). Pemambuco,
Abrolhos Ids.

?Sargassum^aximiliani (Schrad.) Mart.— Martens (1870). Near
Bahia.

Sargassum natans (L.) Meyen.— 5. bacciferum C.Ag., Martens
(1870). « Brazil ».

Sargassum platycarpum Mont.— Mobius (1889) ; Carpacanthus pla-
t^arpus Mont., Martens (1870). Pemambuco, Bahia.

Sargassum polyceraHum Mont. — Howe (1928) ; S. hahiense Kq.,
Martens (1870) ; 5. pol^ph^llum Mert., Martens (1870) ; 5. spinulosum
var. ciliatum Grun., Piccone (1889). Pemambuco. Bahia, Abrolhos Ids.

Sargassum vulgare C. Ag. — Martens (with vars.) (1 870 1871)
Dickie (1875), Gepp (1905), Schmidt (with vars.) (1924). Mucuripe!

Fernando Noronha Ids., Cabo Branco. Pemambuco. Barro Grande. 18« 24*
S. L.; 37°58* W. L.. Abrolhos Ids., Rio Janeiro, Jaquaranga. Probably in
part best referred elsewhere.

Turbinaria turbinata (L.) Ktze.- T, irialata Kg., Martens (1870)
« Brazil ».

RHODOPHYCE/E

BANGIACEiC

Brythrocladia subintegra Rosenv. — Taylor (1930 fc). Conceicao
de Itanhaen ; M. Jundiahy. Sao Vicente.

?Porphyra laciniata (Lightf.) C. Ag. — Martens (with var. umhi-
licata Ag.) (1870). Zeller (1876). Piccone (1886, 1899). Maranhao.
Rio Janeiro.

Porphyra J{oseana Howe. — Howe (1928). Cabo Frio.
Porphyra Sp?. — Taylor (1930 b). Guaruja.

Wm. RANDOLPH TAYLOR

21

NEMALIONACEiC

JlcrochaeHum barbadense (Vickers) Borg.— ?Calliihammon pedimcit-
hatum Kg., Martens (1870). « Brazil >.

AcrochaeHumluxurians(H3iTy.) J. Ag., fa. ^^cwn^^ /a (Lyngb.)TTiur.
— Chantramia secundata (Lyngb.) Tliur., PiCCONE (1889). Pemambuco.

? JlcrochaeHum Savianum (Menegh.) Nag. — Calliihamnion Posido-
nia Zanard., Grunow (1870). « Brazil ».

hiagora ceranoides Lamx. — Howe (Bahama Flora, 1920).
« Brazil ».

?Liagora distenta (Mert.) C. Ag. — ?L. disienta Lamx., Zeller
(1876). Rio Janeiro.

CHi^ETANGIACEiC

Galaxaura cylindrica (Sol.) Lamx. — Martens (1870), Dickie
(1875), Taylor (1930). Fernando Noronha Ids., Bahia, Cabo Frio.

?Galaxaura dichotoma Lamx. — Martens (1870). Aldea Velha.

Galaxaura marginata (E. &S.) Lamx. — Dickie (1875), Mobius
(fa. I'lneam Kg.) (1889), HowE (1928), Taylor (1930); C. canaliculata
Kg., Martens (1870), Zeller (1876). Fernando Noronha Ids., Bahia.
Cabo Frio, Rio Janeiro, Ilha Paqueta, Ilha Sao Sebastiano.

Galaxaura moniliformis Kjellm. — De Toni (Sylloge Algarum 6
141). « Brazil ».

Galaxaura oblongata (^ it S.) Lamx. — Dickie (1875), Howe
(1928), Taylor (1930). Fernando Noronha Ids., Bahia, Ilha Sao Sebas-
tiao.

Galaxaura obfusafa (E. &S.) Lamx. — Martens (1870), Howe

(1928). Bahia, Rio Janeiro.

?Galaxaura ramulosa Kjellm. — De Toni (Sylloge Algarum. 6 :

114). Pemambuco.

Galaxaura rugosa (E. &S.) Lamx. — Dickie (1875) ; G. armulata
Lamx., Martens (1870); G. plicata Kg., Martens (1871). Fernando

Noronha Ids., Bahia, Rio Janeiro.

?Galaxaura umbellata (Esper) Lamx. — Martens (1870). Bahia.

22

MARINE ALCAE OF BRAZIL

GELlDIACEi^E

r

?Cautacanthus rigidus Kg. — Zeller (1876). Rio Janeiro.

?Caulacantkus usfulatus (Men.) Kg. — Cauhacanthus fasiigktus Kk
Martens (1870). MoBIUs (1889). ?Rio Janeiro. Copacabana.

?Gelidium coarctatum Kg. — Martens (1870). Pemambuco.

.^n^^lt^''""^'"'"^''"* (""^'-^ Lamx. — ?Martens (var. mtidum Ag..
1870). Zeller (1876). Mobius (1889). Gepp (1905). Schmidt (1924)
Howe (1928). Taylor (1930). Mucuripe. near Bahia. Cabo Frio. N.cthe^
Toy, Rio Janeiro.

IGelUium corneum y^t. pi„„atum (Huds.) Turner. — ?C. comeurn
n ATm^o'^'T '''"''^' ^"'"-^ ^"™— ^''''^'P^' crinalis Kg.. Martens

Uo/U). Kio Janeiro.

K. f^^^'^'"^ mumfidum Grev. — Martens (1870). Aldea Veiha
Macahe, Rio Janeiro.

?Gelidium parvulum Grev. — Martens (1870), Zeller (1876)
Kjo Janeiro. ^'

Gelidium pusitlum (Stackh.) Le Jolis. — Howe (1928). Rio Janeiro.

rig^m'^'fi^"'"'*'^^^^^ ^'''' - ^'^^^^^ (^««^' ^^S^), Taylor

Wk n ''^'f"'" "'•' Martens (1870); Celidiopsis rigida (Vahl)

Weber-v. Bosse, ScHMIDT (1924) Mucurine C^Un R,o D l

r u JT • T VI ^^t;. mucuripe, <^abo branco, Pernambuco,

v^aDo rno, Jaquaranga.

?Gelidiumsupradecomposnum Kg.— Martens (1870). Rio Janeiro.

.^.J^^^^'^^^^^^^^^^osum Kg. _ Martens (1870), Piccone (1886,
1089). Pernambuco; near Rio Janeiro.

buco''^^^''^'"'" ^^nVx^/Ze (Grev.) Schmitz. - PiccoNE (1886). Pernam.

GIGARTINACEi^

?Catophyllis discigera J. Ag. — Martens (1871). Rio Janeiro

li'in. RANDOLPH TAYLOR

23

Caliophyilis microdonta (Grey.) Falk. — Howe & Taylor (1930),
Taylor (1930); Odonlhalia microdoniu Grev., MarTENS (1870), MoBIUS
(1890). Cabo Frio.

?Chondrus divaricatusGrev. — Martens(1 870). Near Rio Janeiro ?

Endoctadia vernicata J. Ag. — Acanihobolus brasiliensis Kg., Mar-
tens (1870), Zeller (1876). Rio Janeiro.

Gigartina acicularis (Wulf.) Mont. — Martens (with var. p'mnaia
Mont ?) (1870). Macahe. Itapacorohy.

?Gigartina ChamtssoiiMen.) Mont. — G. Chamissoi Kg., Martens
(1870, 1871), Zeller (1876). Rio Janeiro.

?Gigarfina elegans Grev. — Martens (1870). Aldea Velha.

?Gigartina nana (C. Ag.) J. Ag. — Spharococcus nanus C. Ag..
Martens (1870). « Brazil ».

Gigartina Teedii (Roth) Lamx. — Martens (1870), Mobius
(1890). Maranhao, near Rio Janeiro.

Gymnogongrus Griffithsiae (Turn.) Mart. — Martens (1870).
MoBIUS (1890, 1892), Taylor (1930). Rio Janeiro, Ilha Paqueta.

RHODOPHYLLIDACEi*:

Cafenelta impudica (Mont.) J. Ag. — C. impudica Kg., Mobius
(1889, 1895). Joinville, Itajahy.

ICystoclonium difficile (C. Ag.) J. Ag. — Sparococcus difficiUs
C. Ag., Martens (1870). « Brazil ».

Eucheuma echinocarpum Aresch. — Mobius (1889). Olinda.

?Eucheuma Gelidium J. Ag. — Taylor (1930). Ilha Paqueta.

T^hodophyllis gracilarioides Howe & Taylor. — Howe & Taylor
(1930), Taylor (1930). Cabo Fno.

Wurdemannia sefacea Harvey. — Howe (1928), Taylor (1930).
Cabo Frio, Nictheroy, Rio Janeiro, Ilha Govenador.

SPHiEROCOCCACEiE

?Calliblepharis juhata {G. & W.) Kg. —
Pernambuco.

Piccone (1886. 1889)

24

MARINE ALCAE OF BRAZIL

?(Gracilaria) Plocaria aculeata (K^) Crn. — De Toni (Sylloge
Algarum 4 : 397; 6 : 275). « Brazil ».

?Gracilaria armata {C. A%.) J. Ag.— Dickie (1875)), Mobius
(1890), Schmidt (1924). Mucuripe, Fernando Noronha; near Rio Janeiro.

Gracitaria cervicornis (Turn.) J. Ag. — Piccone (1886, 1889),
Mobius (1889, 1890, 1892), Schmidt (1924); (incl. fa. acanthophora
(Kg.) O. C. Schmidt), SphcEfococcus acanthophorus Kg., Martens (1870);
5. cervicornis C. Ag., Martens (1870). Mucuripe, Cabo Branco, Olinda,
Pernambuco, Bahia, Aldea Velha, Rio Janeiro, Jaquaranga.

Gracilaria chondroides (Kg.) Crn. — Maze & Schramm (1870-
11). « Brazil ».

Gracilaria compressa (C. Ag.) Grev.
Jaquaranga.

Schmidt (1924). Mucuripe.

Dickie (1875), Mobius
Sphcerococcus confervoides

Gracilaria confervoides (L.) Grev. —
(1889), Schmidt (1924), Taylor (1930)

C. Ag., Martens (incl. var. setaceus Ag.) (1870, 1871). Mucuripe, Olinda,
Barro Grande, Abrolhos Ids., Aldea Velha, Rio Janeiro.

Gracilaria cornea J. Ag. — Piccone (1886, 1889) Pernambuco.
Gracilaria cuneata Aresch. — De Toni (Sylloge Algarum 4 : 451).

Pernambuco.

? Gracilaria denfafa J. Ag. — G. rangiferina (Kg.), Piccone (1886) ;
Sph<£rococcus rangiferinus Kg., MARTENS (1870). Pernambuco.

?Gracilaria divergens (C. Ag.) J. Ag. — Spharococcus divergent
C. Ag., Zeller (1876). Rio Janeiro.

Gracilaria ferox J. Ag. — Grunow (1870), Piccone (1886,
1889), Taylor (1930). Pernambuco, Rio Janeiro, Scarahy. Ilha Sao Sebas-
tiao.

Gracilaria lacinulata (Vahl) Howe.— G. lacinulala (Kg.), PiccoNE
(1886), G. lacinulata (Vahl) Borg., ScHMIDT (1924); G. multipartita
(Clem.) J. Ag., Dickie (1875), Piccone (1886, 1889), Mobius (1889) ;
G. multipartita Harv., DiCKIE (1875); Sphcerococcus multipartitus C. Ag.,
Martens (? with varieties) (1870). Fernando Noronha, Mucuripe, Cabo
Branco, Olinda, Pernambuco, Macahe, Bahia, Aldea Velha, Rio Janeiro,
Jaquaranga.

«

IVm. RANDOLPH TAYLOR

25

Gracilaria mamillaris Howe.— Taylor (1930). Abrolhos Ids., Cabo
Frio. ^

Gracilaria ornata Aresch. — De Toni (Sylloge Algarum 4 : 450;
6 : 271). Pernambuco.

?Gracilaria SalzmanniBomeL — Mobius (1889). Olinda.

?Hypnea alopecuroides Kg. — ?H. dwaricata Grev., Martens
(1870). Macahe, Aldea Velha, Rio Janeiro.

Hypnea cervicornis J. Ag. — Martens (1870), Taylor (1930).

Rio Janeiro.

IJiypnea hamulosa Lamx. — Martens (1870). Near Rio Janeiro.

hypnea musciformis (Wulf.) Lamx. — Martens (1870, 1871).
Zeller (1876), Piccone (1886, 1889), Mobius (1889, 1890). Schmidt
(1924), Howe (1928), Taylor (1930); H. Esperi Bory, Martens
(1871); H. Rissaana J. Ag., Martens (1870), Zeller (1876). Cabo

Branco, Pernambuco, Bahia, Aldea Velha, Copacabana, Rio Janeiro, Ilha
Sao Sebastiao, Itapocorohy; Aldea dos Reys Magnos.

Bypnea pannosa J. Ag. — Martens (1870). Rio Janeiro.
Tiypnea spinella (C. Ag.) Kg. - Martens (1870), Howe (1928).
Taylor (1930). Bahia, Rio Janeiro, Scarahy, Ilha Sao Sebastiao.

?Sphaerococcus dendroides Kg. — Martens (1870). « Brazil >.
?Sphaerococcus dumosus Martius. — Martens (1870). Between

Bahia and Rio Janeiro.

RHODYMENIACEi^

?Champia compressa Harv. — C. Vidhrdli Kg., Zellfr (1876).
Rio Janeiro.

Champia parvula (C. Ag.) Harv. — Schmidt (1924); Lomentaria
pan^ula Gaill, Zeller (1876). Rio Janeiro, Jaquaranga.

Chrysymenia Dickieana J. Ag. — J. Agardh (Analecta Algol. 1 :
90. 1892"). Bahia.

Chrysymenia Enteromorpha Haryey. — Dickie (1875), Taylor
(1930). Fernando Noronha Ids., Abrolhos Ids.

Frio.

Chrysymenia planifrons (Melv.) J. Ag. — Taylor (1930). Cabo

26

MARINE ALCAE OF BRAZIL

Chrysymenia pyriformis Borg. — Taylor (1930). Cabo Frio.

Chrysymenia Uvaria (L.) J^ Ag. — Piccone (1886, 1889),
Schmidt (1924); Castroclonium uvuria Kg., Martens (1870), Mucuripe,
Cabo Branco, Pernambuco, Bahia, Jaquaranga.

Tauchea HassUri Howe & Taylor. — Howe & Taylor (1930),
Taylor (1930). Cabo Frio.

?Gastroclonium ovale (Huds.) Kg. — Martens (1870). Bahia.

?Lomentaria claveilosa (Turn.) Thuret. — Chondroihamnium clavello-
sum Kg., Martens (1870). Maranhao.

Lomentaria uncinata Menegh. — L. reflexa Chauv. var. uncinata
J. Ag.. Piccone (1889); L. uncinata Mert., Martens (1870). Pernam-
buco.

Plocamium brasiliensis (Cire\.) Howe & Taylor. — Howe & Taylor
(1930). Taylor (1930) ; Thamnophora brasiliensis Grev., Martens (1870,
1871). Zeller (1876). Rio Janeiro.

Plocamium coccineum (Huds.) Lyngb. — Zeller (1876), Mobius
(1890, with fa. Binderiana Hauck, 1892), Taylor (1930); P. coccineum
Grev., Martens (1870). Macahe, Rio Janeiro.

?Piocamium membranaceumSuhr. — P. latiusculum Kg., Martens
(1871). Rio Janeiro.

T'Rhodymenia acanthophora Grev. — Mobius (1889). Olinda.

T{hodymenia Palmafa (L.) Grev. — J. Agardh (Sp., Gen. & Ord.
Algarum 2 : ^Ib-^ll . 1852, Brazil, on authority of Greville).

?J{hodymenia Palmetfa (E^per) Grev. — Mobius (1890, 1892).
Rio Janeiro.

DELESSERIACEiC

Cotfoniella filamentosa (Howe) Borg. — Howe (1928). Brazil.

CoHoniella sanguinea Howe. — Howe (1928). Rio Janeiro.

?Delesseria spathulata Sond. — Dickie (1884). St. Paul's Rocks.

?l^itophyllum acrospermum J. Ag. — Dickie (1884 b). St. Paul's
Rocks.

?JSIitophyUum fimbriatum Grev. — Cr^piopleura fimbriata Kg.. MAR-
TENS (1870). Aldea Velha.

Wm. RANDOLPH TAYLOR

27

i

mophyllum laceratum (Gmel.) Grev. - Crypiopfeura laccrala Kg..
Martens (1870). Southern Brazil.

?mtophyllum monanthos J- Ag. - Mobius (1890). Near R.o
Janeiro.

mtophyllum odontophorum Howe & Taylor. — HowE u Taylor
(1930) ; Taylor (1930). Cabo Frio.

mtophyllum uncinatum (Turn.) Grev. - De ToNI (Sylloge Algarum
4 : 650, 6 : 326), Taylor (1930). St. Paul's Rocks, Cabo Fno.

?Sarcomenia miniata (C. Ag.) J. Ag. - De Ton. (Sylloge Algarum
4 : 735, 6 : 359). « ?Brazil ».

BONNEMAISONIACEiC

Jlsparaeopsis taxiformis (Delile) Collins u Hervey. - A. Ddiki
Mont., Martens (1871). Zeller (1876), MiiBius (1890). R.o Janeiro.

RHODOMELACEiiE

Jfcanthophora muscoides (L.) Bory. - Grunow (1870). Taylor
(1930): A. muscoides Grev.. MARTENS (1870. 1871). Zeller (1876).
MoBIUS (1890). Bahia, Rio Janeiro, Scarahy, Uha Paqueta, Ilha Sao Sebas-
tiao.

Jlcanthophora spicifera (Vahl) Borg. - Schmidt (1924) ; A. Thierii
Lamx., Martens (1870), Zeller (1876), Dickie (1875). ?Schmidt

(1924). Fernando Noronha Ids., Mucuripe, Bahia, Rio Janeiro. ? Jaquaranga;
Villa Velha.

Jlmansia multifida Lamx. - Martens (1871), Piccone (1886.
1889), Schmidt (1924), Howe (1928) ; Epineuron multifidum Kg.. Mar-
tens (1870). Mucuripe, Cabo Branco. Pernambuco. Bahia. Abrolhos Ids..
Rio Janeiro.

JJmphibia radicans (Mont.) Klze. — Boslr},chia radicans Mont., with
fa. brasiliana Mob., MoBIUS (1889, 1890, 1895). Near Rio Janeiro, Join-
ville, Itajahy.

JJmphibia Sertularia (Mont.) Howe. — Boslrj,c/iia SerlJaria Mont.,
Martens (1871), Mobius (1890). Rio Janeiro.

Amphibia tenella (Vahl) Ktze. — Taylor (1930); Boslry^chia (e-

28

MARINE ALCAE OF BRAZIL

nella (Vahl) J. Ag.. MoBIUS (1890); B, Viellardi Kg., ZellER (1876).
Rio Janeiro, Sao Vicente.

Bryocladia thyrsigera (J. Ag.) Schmitz. — Taylor (1930). Sao
Vicente.

Bryothamnion Seaforthii (Turn.) Kg. — Martens (1870), Pic-
cone (1886), Dickie (1875), Schmidt (1924), Howe (1928), Taylor

(1930). Maranhao, Mucuripe, Cabo Branco, Pernambuco, Bahia, Abrolhos
Ids., Jaquaranga, Santos; Manguinos.

Bryothamnion triquetr urn (Omel) Howe. — Schmidt (1924),
Howe (1928), Taylor (1930); B. triangulare Kg., Martens (1870).

Maranhao, Mucuripe, Cabo Branco, Pernambuco, Bahia, Jaquaranga.

Chondria atropurpurea Harv. — Chondriopsis atropurpurea Harv.,
Dickie (1875). Bahia.

Chondria fioridana (Collins) Howe. — Taylor (1930). Pernam-
buco, ?Cabo Frio.

Dasya sertulanoides Howe & Taylor. — Howe & Taylor (1930).
Taylor (1930). Abrolhos Ids.

- Howe (1928); D. Wulfeni Kg..

Digenia simplex (Wulf.) Ag. -
Martens (1870). Bahia.

Dipterosiphonia dendrihca (Ag.) Falk. — Schmidt (1924); Pol}f-
siphonia dendritica Ag., MARTENS (1870). Cabo Branco, Jaquaranga.

'Enantiocladia Duperryi (C. Ag.) Falk. — Amansia Duperryi J. Ag..
Dickie (1875). Fernando Noronha Ids.

?Halopity s pinastroides (Omel.) Kg.- Schmidt (1924). Jaquaranga.

?Laurencia dendroidea J. Ag. — Martens (1870). « Brazil ».

?Laurencia hyhrida (D. C.) Lenorm. — Martens (1870), Zeller
(1876). Rio Janeiro.

Laurencia lata Howe & Taylor — Howe & Taylor (1930), Tay-
lor (1930). Cabo Frio.

Laurencia ohtusa (Huds.) Lamx. — Martens (1870, ?incl. var.
gracilis C. Ag.), MoBius (1889, 1890). Schmidt (1924). Mucuripe, near
Bahia, Aldea Velha, Macahe, Rio Janeiro, Copacabana, Jaquaranga.

Laurencia papillosa (Forsk.) Grev. — Martens (1870), Piccone
(1886, 1889), Schmidt (1924), Taylor (1930). Mucuripe, Cabo Branco.

<m V

«" ■ »

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Wm. RANDOLPH TAYLOR

29

Pernambuco. Aldea Velha, Macahe, Rio Janeiro, Jaquaranga, Ilha Sao Sebas-
tiao.

?Laurencia pinnatifida (Cmel) Howe. — Martens (1870). Ma.
ranhao, Aldea Velha.

Laurencia Poitei (Lamx.) Howe. — Schmidt (1924); L. brasiliana
Mart., Martens (1871), Zeller (1876). Rio Janeiro, Jaquaranga.

Laurencia scoparia J. Ag. — Martens (1871), Piccone (1886).

Pernambuco.

?Laurencia thyrsoidea Mont. — Martens (1870). Bahia.

?Lophosiphonia neglecta (Harv.) De Toni. — Polysiphoma neglecta
Harv., Zeller (1876). Rio Janeiro.

?Polysiphonia dichotoma Kg. — Martens (1870). « Brazil ».

?Polysiphonia elongata (Huds.) Harv. - P. dongaia Ag., Martens
(1870), Zeller (1876). Rio Janeiro.

Polysiphonia suhtilissima Mont. — Mobius (1890). « Brazil ».

?Polysiphonia violacea iP^o\\\) Grev. — Martens (1870), Mobius
(1890). « Brazil ».

?(Yidaliaj J{yHphlaea Tiilariana Grev. — Martens (1870).
« Brazil ».

i Vidalia ohfusiloha (Ag.) J. Ag. — Piccone (1886, 1889), Mobius
(1889), Schmidt (1924). Howe (1928); Ryilphloea obtusiloba C. Ag..

Martens (1870). Mucuripe, Cabo Branco, Olinda, Pernambuco, Bahia,
Aldea Velha.

WrightiellaTumanowtczii (Cany) Schmitz. — Das}fa Tumamwiczii
Harv., Dickie (1875). Bahia.

CERAMlACEiC

Callithamnion corymbosum (Sm.) Lyngb. — Phkbothamnion versi-
color Kg., Martens (1870). Bahia, Cabo Frio.

Callithamnion dasytrichum Mont. — De Toni (Syiloge Algarum
4 : 1362); Collins. Holden & Setchell, P. B.-A. 697. « Brazil ».

Centroceras clavulatum (C. Ag.) Mont.- Piccone (1886), Mobius
(1889, 1890), Schmidt (1924), Howe (1928), Taylor (1930b); C.
leptacanthum Kg.. MarTENS (1870); C. macranthum Kg.. MartENS

30

MARINE ALCAE OF BRAZIL

Wm. RANDOLPH TAYLOR

31

(1870); C. micranthum Kg., Martens (1870, 1871). Zeller (1876);
C. clavulatum var. mkranihum (Kg.). PiCCONE (1886, 1889). Mucuripe,
Cabo Branco, Pernambuco, Macahe. Rio Janeiro, Copacabana, Sao Vicente,
Conceicao de Itanhaen, Guaruja, Itapacorohy.

?Ceramium ciliatum (EH) Duel — Echinoceras ciliatum Kg., MAR-
TENS (1870). « Southern Brazil ».

?Ceramium diaphanum (Lightf.) Roth. — Hormoceras diaphanum
Kg., Martens (1870), Zeller (1876). Rio Janeiro.

Ceramium Luetzelhurgii O.C. Schmidt. — Schmidt (1924). Cabo

B

ranco.

Martens (1870). « Central

?Ceramium ohsolefum C. Ag. -
Brazil ».

?Ceramiuni ruhrum (Huds.) C. Ag. — Martens (1870). « Sou-
them Brazil ».

Ceramium sfrictum Grev. & Harv. — MoBIUS (1890). « Brazil ».

Ceramium tenuissimum (Lyn^h.) C. Ag. - MoBIUs (1889), Schmidt
(1924). Cabo Branco.

?Ceramium vimineum J. Ag. — C. ruhmm virgaium C. Ag., Mar-
tens (1870). « Southern Brazil ».

Griff ithsia radicans Kg- — Martens (1870), Schmidt (1924),^

Taylor (1930). Jaquaranga, Ilha Sao Sebastiao.

Jialoplegma Duperryi Mont. — Piccone (1886). Pernambuco.

Spyridia aculeata (Schimp.) Kg. — Taylor (1930). Rio Janeiro.

Spyridia complanata J. Ag. — Murray (1889). « Brazil ».

Spyridia filamentosa C^uU.) Harv. — Schmidt (1924), Taylor
(1930)1 S. arcuata Kg., MoBIUs (1890). Cabo Branco, Abrolhos Ids., Pa-
queta.

GRATELOUPlACEi?:

Cryptonemia crenulata J. Ag. - Dickie (1875), Piccone (1886),
MoBIUS (1889) ; Acrodlscus crenulalus (J. Ag.) De Toni. ScHMIDT (1924) ;
Ph\)llophora crenulata J. Ag., MARTENS (1870). Cabo Branco, Olinda, Per-
nambuco, Bahia, Rio Janeiro.

Cryptonemia iuxurians (Men.) J. Ag. — Martens (1870, 1871),

*v ^

«■ >

^ I ^

Piccone (1886). Gepp (1905), Schmidt (1924). Mucuripe, Pernambuco,

Rio Janeiro, Jaquaranga.

Grafeloupia cuneifolia ]• Ag. - Martens (1871), Mobius (1890).
Rio Janeiro.

Grateloupia filicina (Wulf.) C. Ag. — Martens (1870), Piccone
(1886), Schmidt (1924). Mucuripe, Cabo Branco, Aldea Velha, Rio Ja-
neiro, Jaquaranga.

Jialymenia Tloresia (Clem.) Ag. — Dickie (1875). Taylor
(1930). Bahia. Abrolhos Ids.

Jialymenia integra Howe & Taylor. — HowE & Taylor (1930).
Taylor (1930). Cabo Frio.

?Balymenia ligulata (Woodw.) C. Ag. — Mobius (1890). « Bra-
zil ».

Jialymenia rosea Howe & Taylor. — Howe & Taylor (1930).
Taylor (1930). Cabo Frio.

Jialymenia vinacea Howe & Taylor. — Howe & Taylor (1930).
Taylor (1930). Cabo Frio.

NEMASTOMACEi^E

Platoma tenuis Howe & Taylor. — Howe & Taylor (1930). Tay-
lor (1930). Cabo Frio.

SQUAMARIACEi^E

?Jiildenbrandtia expansa Dickie. — Dickie (1884 b). St. Paul's
Rocks.

?Jiildenhrandtia prototypus Nardo. — H. Nardil Zanard.. ZelleR
(1876). Rio Janeiro.

?Jiildenbrandtia rosea Kg. — Zeller (1876). Rio Janeiro.

Peyssonnelia DubviCm.— Dickie (1875). Fernando Noronha Ids.

?PeyssonneliaimbricataKg. — Zeller (1876). Rio Janeiro.

?Peyssonnelia squamaria (Cmel) Dene.— Martens (1870). « Bra-

zil ».

CORALLINACEiC

Jlmphiroa Beauvoisii Lamx. — Piccone (1886). Schmidt (1924) ;

32

MARINE ALCAE OF BRAZIL

Wm. RANDOLPH TAYLOR

33

Amphlroa exilis Harv., MARTENS (1870, 1871). MbBIUS (1889). Rio Ja-
neiro, Jaquaranga.

Amphiroa brasiliana Dene. - Martens (1870). Zeller (1876).
M6BIUS (1889). Howe (1928). Taylor (1930. 1930 b). Olinda, Bah.a.
Nictheroy. Rio Janeiro, Guaruja.

Amphiroa fragiUssima (L.) Umx. - Taylor (1930). Nictheroy.

Amphiroa variabilis Harv. - Martens (1870). Cabo Frio.

?Cheilosporum anceps (Kg.) Yendo. - Corallka anceps Kg.. MoBius
(1889. 1890). Copacabana.

?Cheihsporum cultratum (Harv.) Aresch. - Martens (1870),
MoBIUS (1890). Rio Janeiro.

?Cheihsporum palmatum (E. ^ S.) Yendo. - Arlhrocardia pdmcla
Aresch.. Martens (1870) ; var. Filicula (Umx.) Yendo. {Corallma Fxbcula
Lamx.) Zeller (1876). Rio Janeiro.

?Cheilosporum planiusculum (Kg.) Yendo. - CoraUha Pi^r,iuscuh
Kg. (and forms). Martens (1871), P.ccone (1886), Mobius (1889,

1890). Pernambuco, Rio Janeiro, Botofago.

?Cheilosporum sagittatum (Lamx.) Aresch., fa. minor. — PChei-
losporum sagilMum (Umx.) Harv.. MoBIUS (1889. 1890). Cabo Fr.o.

?Corallina carinata Kg. — Zeller (1876). Rio Janeiro.
?Corallina ceratoides Kg. - Dickie (1875). Fernando Noronha

Corallina cubensis (Mont.) Kg. emend. Borg. — Jania cubemls Mont..
Martens (1871). Dickie (1875). Fernando Noronha Ids.. Rio Janeiro.

Corallina officinalis L. - Martens (1870. 1871). Zeller
(1876). Schmidt (1924). Mucuripe. Bahia, Rio Janeiro, Jaquaranga.

Corallina subulata Ell. a. Soil. - Martens (1870), P.ccone
(1886). Howe (1928) ; C. Cuvieri Umx.. var. subulaki (E. & S.) Aresch.,
Schmidt (1924). Mucuripe, Abrolhos Ids., Bahia, Jaquaranga.

Dermatolithon pustulatum (Umx.) Fosl. - Melobesia pusiulala
Umx., PicCONE (1886). Pernambuco.

Goniolithon mamillare (Harv.) Fosl. — Mehbesia mamillaris Harv..
Martens (1870). Bahia.

Jania aJhaerens Umx. — Martens (1870). Taylor (1930). Rio

Janeiro, Scarahy.

■-4

Jania capillacea Harv. — Howe (1928). Taylor (1930). Bahia.
Nictheroy, Guaruja.

?Jania fastigiata Harv. — Mobius (1890), « Brazil ».

Jania rubens (L.) Lamx. — Martens (1871). Piccone (1886),
MoBIUS (1890, 1892). Schmidt (1924); Corallina rubens L.. MoBIUs
(1889). Mucuripe, Abrolhos Ids., Rio Janeiro, Copacabana. Jaquaranga.

hifhothamnium brasittense Fosl., and fa. heteromorpha Fosl. —
FOSLIE (1900). Sao Sebastiao.

?Lithothamnium erubescens Fosl. — Foslie (1900): L. maml'Jarc
Harv., Dickie (1875). Fernando Noronha Ids.

?Ufhothamnium fasciculafum (Lamk.) Aresch.— Mobius (1889).
Bahia.

?Lithothamnium Lenormandi (Aresch.) Fosl. — Liihophyllum Le-
normandi (Aresch.) Rosan., MoBIUS (1890). « Brazil ».

? Lithothamnium lichenoides (F. & S.) Heydr. — ?Melohesia liche-
noides Kg., Dickie (1884b). St. PauFs Rocks.

Lithothamnium membranaceum (Fsp.) Fosl. — Melobesia membra-
nacea (Esper.) Lamx.. PiCCONE (1886), MoBIUS (1889). Pernambuco,
Abrolhos Ids.

?Lithothamnium polymorphum (L.) Aresch. — Dickie (1884 b),
MoBIUS (1889, 1890). St. Paul's Rocks, Cabo Frio.

?Lithothamnium scabiosum (Harv.) Fosl. — Mehobesia scabiosum
Harv., Martens (1870). Bahia.

Melobesia farinosa Lamx. — Piccone (1886). Pernambuco.

Melobesia Lejolisii Rosan. — MoBIUS (1890). <c Brazil ».

POSITION UNCERTAIN

lHormotrtchum vtrtdtfuscum Mont. — Martens (1870). Macahe.

IJialonema obscurum Dickie. — Dickie (1875). Fernando Noronha
Ids.

IDictyostphon charotdes Zeller. — Zeller (1876). Rio Janeiro
(PSlictyosiphon.)

ISporacanthus cristatus Kg. — Martens (1870). Pernambuco.

ISpongites verrucuhsaZeWer. — Zeller (1876). Rio Janeiro.
( ? Lithothamnium.)

34

MARINE ALCAE OF BRAZIL

BIBLIOGRAPHY

BoRGESEN, F. — The marine algae of the Danish West Indies. Reprinted
from Damk Bot. Ark'iv. i, Chlorophyceae, 1913. ii, Phaeophyceae, 1914.
iii, Rhodophyceae, with supplement, 1915-1920.

Dickie. G. — Enumeration of algae collected by H. M. Moseley M. A..
Naturalist to H. M. S. Challenger. Jour. Linn. Soc. Bot. London, 14 : 363-
365, 375-377. 1875.

Dickie, G. 1884. See W. B. Hemsley.

Gepp, a. & E.-S. — Atlantic algae of the « Scotia ». Jour. Bot.
43: 109-110. 1905.

Grunow, a. — Reise der osterreichischen Fregatte « Novara » um die
Erde in den Jahren 1857-1859 : Algae. Bot. TeiL 1 : 1-104. 1870.

Hemsley, W.-B. — Report on the botany of the Bermudas and various
other islands of the Atlantic and Southern oceans. Rept. Sci Res. Voy.
H.M.S. Challenger 1873-76. (Bot.) I, 1 : 1-128; II. 1 : 1-281. (1884)
1885.

Howe, M.-A. — Notes on some marine algae from Brazil and Barbados.
Jour. Washington Acad. Sci.. 18 : 186-194. 1928.

Howe, M.-A. & W.-R. Taylor. — Notes on some new marine algae
from Brazil. Brittonia /. 1930.

Martens, G. — Conspectus Algarum Brasiliae haectenus detectarum.
Kjob. Vidensk. Meddel. 1870:279-314. 1870.

Martens, G. — Algae Brasilienses circa Rio de Janeiro a dar. A. Gla-
ziou, horti publ. direct., botanico indefesso, annis 1869 et 1870 coUectae.
Kjob, Vidensk. Meddel. 1871 : 144-148. 1871.

Martius, K.-F.-P. von. F.-G. Eschweiler & C.-G. Nees von
E3ENBECK. — Algae, Lichens, Hepaticae exposuerunt in Martius Flora Bra-
tiliensis. 1 (1) :i-iv, 1-390. 1883.

Maze, H. & A. Schramm. — Essai dc classification des algues de la
Guadeloupe. Basse-Terre. 1870-77.

MoBius, M. — Bearbeitung der von H. Schrenk in Brasilien gesam-
■lelten Algen. Hedxvigia 28 : 309-347. 1889.

■ ( .

Wm. RANDOLPH TAYLOR

35

\

■i-

■i

r 1 ♦

MoBius, M. — Algae Brasilienses a cl. Dr. Glaziou coDectse. Notarisia
5 : 1065-1090. 1890.

MoBius, M. — Ueber einige brasilianische Algen. Ber. deutsch. boL
Cesell. 10 : 17-26. 1892.

MoBius, M. — Ueber einige brasilianische Algen. Hedrvigia 34 : 1 73-
180. 1895.

MoNTAGNE, C. — Cryptogamse brasilienses sen Plants ceDulares quas
in itinere per Brasiliam a celeb. Auguste de Saint Hilaire coUectas recensuit
observationibusque nonnuilis illustravit. Ann. Sci. Nat., Bot., ii, 12 : 42-44.
1839.

Murray, G. — Catalogue of the marine algae of the West Indian^
region. Jour, of Bot., 26 : 193-196, 237-243, 303-307, 331-338; 27 : 237-
242, 257-262, 298-305. 1888-1889.

PiccoNE, A. — Alghe del viaggio di circumnavigazione della « Vettor

Pisani ». Genoa, 1886, 87 pp., 2 pi.

PiccoNE, A. — Nuove alghe del viaggio di circumnavigazione della
« Vettor Pisani ». Reale Acad. Lincei (286) 4", 6 : 64 pp. 1889.

Schmidt, O.-C. — Meeresalgen der Sammlung von Leutzelburg aus
Brasilien. Hedwigia 65 : 85-100. 1924.

Taylor, W.-R. — Algae collected by the Hassler, Albatross and Schmitt
expeditions, I. Marine algae from Brazil. Amev. Jour. Bot. 17. — 1930.

Taylor. W.-R. — Algae from Sao Paulo, Brazil, Amer. Jour. Bot. 17.
— 1930.

Zeller, G. — Algae Brasilienses circa Rio Janeiro a Dr. A. Glaziou,
horti publ. dir., collect*. Kjob. Vidensk. Meddel. 1876 : 426-432. 1876.

(Received for Publication 20-2-1 930.)

Extrait de la Revue. Algologique (Vol. V, fasc. 3-4). Publicc au Laboratoire de Crypto-
gamie du Museum National d'Histoire Naturelle, a Paris.

r

Reprinted from American P^'ern Journal, Vol. 21, No. 3,

July-Septeinber, 1931.

4 b >

4 19

H>

*m f

^ I ^

ASPLENIUM BRADLEYI ErRONKOUSLY ReI'ORTKD OX LIME-
STONE AGAIN. — In carrying out studies on the soil reac-
tion preferences of ferns, I have repeatedly tested the
soils supporting Bradley's Spleenwort, and have in-
variablv found them to be decidedlv acid.^ This has led
me to inspect critically reports of the findings of the spe-
cies on limestone, where the soils are normallv alkaline,
and in all cases the data have proved open to doubt.
As the idea still seems to prevail in many phices that
this fern is 'Mime-loving," however, further discussion
of the matter seems called for.

Asplenium bradleiji was collected near Xewburgh, New
York, "on limerock," by Bumstead and Eaton- in 18()4;
as the rocks exposed near that place are dominantly
siliceous, however, there would appear to be slron*:
probability that the rock supporting the fern was
wrongly identified. It was later found in the Shawan-
gunk Mountains by Clarence Lown.-^ The compiler of
the Flora of the Vicinity of New York^ stated it to be
''known in our area only from the predominantly lime-
stone region in the Shawangunk Mountains," although
reference to any geological map would have shown him
that these mountains are made up almost entirely of
sandstone rocks instead.

^Vhen statements of this sort get into the literature,
however, their copying from one comj)ilation to another

1 Wherry, Am. Fern J. 10: 17, 47. 1920. If).- 47. 1J»25. IS:
62. 1928.

2 Eaton, Ferns N. A. 2: 40. 1893.

3 Davenport, Bull. Torr. Bot. Club 10: 6. 1893.
■» Taylor, Fl. Vie. N. Y. (U). 191.').

41 »

112

American Fern Journal

seems to be inevitable. So, in the Annotated List of the
Ferns and Flowering Plants of New York State'^ we find
the above cited statements paraphrased as ''On rocks,
preferring limestone," and these words were faithfully
repeated in the Fern Lover's Companion/'

Recently there has been published an elaborate book
on the Wild Flowers of the Alleghanies,' including a
check list of the ferns of that region. The latter con-
tains data as to the habitats of the several species,^ and
Aaplenium hrodleyi is alleged to grow ''usually on lime-
stone rocks." Unfortunately, in acknowledging my aid
in correcting the proof of that work, its writer inad-
vertently failed to mention that considerable material,
including that on ferns, was added after I had seen the
proof. This omission places me in the awkward posi-
tion of seeming to sponsor the very statement as to the
soil preferences of this fern which I have been trying
so hard to correct for the past ten years !

For the benefit of future c(mipilers of data on plant
habitats, then, I am once more stating the situation with
reference to Asplenium hradleyi: As far as authenti-
cated records go, this fern never occurs on limestone at
all, but is strictly limited to sandstone, quartzite, mica-
gneiss, and other siliceous rocks, in the crevices of
which decidedly acid soils have accumulated. — Edgar T.
Wherry, Department of Botany, University of Penn-
sylvaniu.

r. House, Bull. N. Y. St. Mus. No. 254: 27. 1924.
«Tilton, Fern Lover's Comp. 87. 1922.
7 Harned, Wild Fl. Alleghaiiics 624. 1931.

s These seem to have been compiled from more or less untrust-
worthy sources, as many of them are inaccurate or misleading.

..

i >

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IRREGULAR PAGINATION

Reprinted from Journal of Thb Washington Academy of Sciences

Vol. 21, No. 9, May 4, 1931

MAY 4, 1931

wherry: leatherflowers

195

BOTANY. — The eastern short'Stemmed leatherflowers.^ Edgar T.
Wherry, University of Pennsylvania.

In current botanical manuals two species of short-stemmed broad-
leaved leatherflowers are listed as present in the eastern United States,
Clematis ochroleuca Alton and C. ovata Pursh. Field studies of this
group of plants in the Appalachian shale-barrens^ and elsewhere have
indicated that their relationships and ranges have been to some ex-
tent misinterpreted, as the data here recorded may serve to show.

The principal diagnostic features of the plants in question are pre-
sented in the accompanying key. Certain characters often regarded
as significant have proved to be inconstant and variable, and have ac-
cordingly been omitted from consideration. For instance, leaf -termina-
tions range from obtusish to acutish or even acuminate from one
branch to another on a single plant. Again, the violet tinge on the
outside of the sepals may be intense on one individual and almost
lacking on another growing beside it, with intermediates elsewhere in
the vicinity. Finally, dimensions of sepals and of achenes vary by
25% within any large clump, depending on the degree of maturity
attained by the particular branch on which they are borne.

1 Contribution from the Botanical Laboratory of the University of Pennsylvania.
Received March 2, 1931.

* This Journal 20: 46. 1930.

t

Key to the Eastern Short-Stemmed Leatherflowers (Clematis spp.)

Plant sparingly branched and small leaves relatively few; head of fruit tend-
ing to be spherical, about 6 cm. in diameter; achenes nearly symmetrical.
Under side of leaves glabrate to moderately pubescent ; hairs of achene-
appendages deep, or exceptionally pale, yellow; range chiefly at
altitudes below 1000 feet, mostly in Piedmont. . . .C. ochroleuca ovata
Under side of leaves moderately to densely pubescent; hairs of achene-
appendages pale, or exceptionally deep, yellow; range chiefly at alti-
tudes above 1000 feet, mostly in Blue Ridge. . . .C. ochroleuca sericea
Plant copiously branched and small leaves relatively numerous; leaves glab-
rate.
Head of fruit nearly spherical, about 5 cm. in diameter; achenes fairly

symmetrical, their appendage-hairs brown C. viticaulis

Head of fruit spheroidal, about 4 cm. high and 6 cm. broad; achenes
rather unsymmetrical, their appendage-hairs whitish. . . .C. albicoma

Clematis ochrolevca Aiton. — This plant varies in a number of respects from
one clump to another, but the only features in which such variation shows
any recognizable geographical relationships are those enumerated in the key.
Two extreme variants with respect to degree of leaf-pubescence have received
specific names, but in view of the complete gradation between them only
varietal distinction seems justified. It is accordingly here proposed to divide
this species into two varieties, as follows:

Clematis ochroleuca ovata (Pursh) Wherry, status novus'

C. ovata Pursh, not of current manuals
C. integrifolia a ochroleuca Kuntze.

The specimen on which Pursh based his specific name is preserved in the
Sherard Herbarium at Oxford University, having been collected by Catesby
and labelled by him with a citation from Plukenet, followed by the words
"negroes head." The latter has been regarded as a locality,* but as Catesby
did not in general add place-names to his labels, and as '^nigger- head" —
in allusion to the globular mass of kinky plumes — is the term universally
applied to the leatherflowers by laymen in the south, it is believed to repre-
sent a common name instead.

In his work on the Natural History of Carolina, etc., Catesby 'did not men-
tion this plant. Pursh^ supposed it to have been obtained in South Carolina,
Small® in that state or Georgia. It could equally' well have come from Vir-
ginia, which was also visited by Catesby, as shown by the following quotation:'

"In the Year 1714 I travelled from the lower Part of St. James's River in
Virginia to that Part of the Apalatchian Mountains where the Sources of that

' ^'Status novas'' is believed to express the situation more accurately than the more
frequently used ^'comhinatio nova."

* Britton, Mem. Torr. Bot. Club 2: 28, footnote. 1890; Small, Flora Southeastern
U. S. 439. 1903.

» Pursh, Flora Amer. Sept. 2: 736. 1814.

• Small, loc. cit.

' Catesby, Nat. Hist. Carolina, etc. 1: v. 1731.

IRREGULAR PAGINATION

196 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 21, NO. 9

River rise, ... At the Distance of twelve Miles from the Mountains we
left the River, and directed our Course to the nearest of them."

The specimen in question was examined by Asa Gray, who stated^ that it
''appears to be C. ochroleuca, Ait." It was later compared by Messrs. Vines
and Druce with material from a West Virginia shale-barren sent to England
for the purpose by Dr. N. L. Britton,^ and was concluded by them to be iden-
tical with this. Their interpretation has been adopted in current manuals,
but as there is no evidence that Catesby ever reached the shale-barren region,
its reconsideration seemed desirable. Accordingly, at my suggestion, Dr.
and Mrs. Francis W. Pennell kindly obtained data on the specimen in Octo-
ber 1930, and their description of it indicates clearly that it does not repre-
sent the shale-barren plant after all. In lacking small-leaved branches, and
having relatively large leaves and a spherical head of achenes with pale yellow
appendage hairs, it corresponds exactly to the plant of the Virginia and
Carolina Piedmont.

The range of this variety is indicated by the following county records :^^

GEORGIA.

DeKalb: Stone Mt., AshCj not dated (N)

SOUTH CAROLINA.

Abbeville: Abbeville, Porcher, Aug. 1898 (U)

NORTH CAROLINA.

Alamance: Graham, Biltmore, May 26, 1902 (U)
Caldwell: Lenoir, Biltmore, May 17, 1902 (U)
Davie: Farmington, Biltmore^ Aug. 27, 1895 (U)
Guilford : High Point, Canhy, June 1868 (N)
Halifax: Weldon, Williamson, April 16, 1897 (N, P)
Iredell: Statesville, Hyams, June 1898 (N, U)
Polk: Lynn, Peattiey April 19, 1919

Randolph: , Ashe, June 1895 (N)

Rowan: Salisbury, many collectors and dates (N, P, U).

VIRGINIA.

Arlington: many localities north of Alexandria (G, N, P, U)

Dinwiddie: Petersburg, Tuomey, not dated (P)

Fairfax: many localities south of Alexandria (G, N, P, U)

Fauquier: Buckland, Meredith, May 25, 1922 (P, U)

Greensville: Belfield (now Emporia), Heller, June 19, 1893 (G, N, P, U)

Henrico: Richmond, Burk, July 25, 1887 (P)^^

Spotsylvania: Fredericksburg. Ward, May 3, 1872 (U).

• Gray, Curtis's Bot. Mag. [3] 37; pi. 6594. 1881.

• Britton, loc. cit.

*» The following abbreviations are used for names of herbaria: G, Gray Herbarium;
.N, New York Botanical Garden; P, Academy of Natural Sciences of Philadelphia;
U, U. S. National Herbarium.

" This specimen is especially similar in leaf outline and pubescence to the type of C.
ovata; as Catesby would have started his 1714 trip in the vicinity of what is now Rich-
mond, it is quite possible that they came from the same station.

MAY 4, 1931

WHERRY : LEATHERFLOWERS

197

PENNSYLVANIA.

Chester: London Grove (old record; no specimens seen).

NEW YORK.

Queens: Brooklyn, Carey, May, 1841 (G, X)

Kichmond: many localities on Staten Island (G, X, P, U).

Clematis ochroleuca sericea (Michaux) Wherry, status novus.

C. sericea Michaux.

C ochrolema jS Torrey and Gray.

C. integri folia a ochroleiica 2 tomentosa Kuntze.

Fig. 1. Clematis albicoma Wherry
West of Covington, Virginia, June 10, 1930.

This is the variety of C. ochroleiica which occurs at the higher elevations.
It is characterized by its tendency toward tomentose pubescence on the leaves
and pale hairs on the achene appendages. The most extreme material seen
is that from Botetourt county, Virginia; the remaining records here cited
are of specimens more or less transitional to the other variety.

GEORGIA.

Stevens: 5 miles west of Toccoa, Wherry, April 7, 1930 (P).

north CAROLINA.

Forsyth: Salem, Schwcinitz, not dated (P).

196 JOURNAL OF THE WASHINGTON ACADEMY OP SCIENCES VOL. 21, NO. 9

River rise, ... At the Distance of twelve Miles from the Mountains we
left the River, and directed our Course to the nearest of them."

The specimen in question was examined by Asa Gray, who stated'^ that it
'^appears to be C, ochroleucaj Ait." It was later compared by Messrs. Vines
and Druce with material from a West Virginia shale-barren sent to England
for the purpose by Dr. N. L. Britton,^ and was concluded by them to be iden-
tical with this. Their interpretation has been adopted in current manuals,
but as there is no evidence that Catesby ever reached the shale-barren region,
its reconsideration seemed desirable. Accordingly, at my suggestion. Dr.
and Mrs. Francis AV. Pennell kindly obtained data on the specimen in Octo-
ber 1930, and their description of it indicates clearly that it does not repre-
sent the shale-barren plant after all. In lacking small-leaved branches, and
having relatively large leaves and a spherical head of achenes with pale yellow
appendage hairs, it corresponds exactly to the plant of the Mrginia and
Carolina Piedmont.

The range of this variety is indicated by the following county records :^^

GEORGIA.

DeKalb: Stone Mt., Ashe, not dated (X)

SOUTH CAROLINA.

Abbeville: Abbeville, Porrher, Aug. 1898 (U)

NORTH CAROLINA.

Alamance: Graham, Bilimore, May 26, 1902 (U)
C\aldwell: Lenoir, Biltmore, May 17, 1902 (U)
Davie: Farmingtcm, Biltmore, Aug. 27, 1895 (U)
Guilford: High Point, Canby, June 1868 (N)
Halifax: Weldon, Williamson, April 16, 1897 (N, P)
Iredell: Statesville, Hymns, June 1898 (N, U)
Polk: Lynn, Pentiie, April 19, 1919

l^andolph: , Ashe, June 1895 (N)

Rowan: Salisbury, many collectors and dates (N, P, U).

VIRGINIA.

Arlington: many localities north of Alexandria (G, N, P, U)

Dinwiddie: Petersburg, Tuomey, not dated (P)

Fairfax: many localities south of Alexandria (G, N, P, U)

Fauquier: Buckland, Meredith, May 25, 1922 (P, U)

Greensville: Belfield (now Emporia), Heller, June 19, 1893 (G, N, P, U)

Henrico: Richmond, Burk, July 25, 1887 (P)^^

Spotsylvania: Fredericksburg. Ward, May 3, 1872 (U).

8 Gray, Curtis's Bot. Mag. [3] 37: pi. 6594. 1881.

• Britton, loc. cit.

^° The following abbreviations are used for names of herbaria: G, Gray Herbarium;
.N, New York Botanical Garden; P, Academy of Natural Sciences of Philadelphia;
U, U. S. National Herbarium.

" This specimen is especially similar in leaf outline and pubescence to the type of C.
ovata; as Catesby would have started his 1714 trip in the vicinity of what is now Rich-
mond, it is quite possible that they came from the same station.

41 «

*m*

MAY 4, 1931

WHKHUY : LKATHKKKLOWKKS

197

PEWSYLVANIA.

Chester: London Grove (old record: no specimcMis seen).

NEW YORK.

(Jueens: Hrooklyn, Cahkv, May, ISll ((1, X)

Hichniond: many localities on Staten Island ((1, X, P, 1).

Clematis ochroleuca sericea (Michaux) Wherry, status novus

C. sericea Michaux.

C. (x-hrolenva p' Torrey and Gray.

C. integrifoUn a orhroleura 2 ((tmcntosa Kuntz(\

Fig. 1. Clematis alhUomn Wherry
West of Covington, Virginia, June 10. 1930.

This is the variety of C. ix'hroleuca which occurs at the hijjjlKT elevations.
It is characterized by its tendency toward tonientose pubescence on the leaves
and pale hairs on the achene appendages. The most extreme material seen
is that from Botetourt county, Virginia; the remaining records here cited
are of specimens more or less transitional to the other variety.

GEORGIA.

Stevens: 5 miles west of Toccoa, Wherry, April 7, 1930 (P).

NORTH CAROLINA.

Forsyth: Salem, Schwcinitz, not dated (P).

INTENTIONAL SECOND EXPOSURE

198 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 21, NO. 9

VIRGINIA.

Botetourt: Eagle Rock, LeiviSj April 27, 1929 (P, Va. State Herbarium).
Koanoke: Roanoke, E. 6^. BriUonund Vail, May 16-27, 1892 (G, N, P, U).

Clematis viticaulis Steele' ^
This species is known thus far only from the type locality, where it occurs
on shale slopes along the railroad west of the tunnel. The characters listed in
the key amply differentiate it.

VIRGINIA.

Bath: West of Millboro, Steele, Sept. 3, 1906 (N, U); Wherry, June 11,
1930 (P).

Clematis albicoma Wherry, nomen novum

C. ovata of current manuals, not Pursh

So far as recorded this plant was first collected on Kates Mountain by Gus-
tav Guttenberg in 1877. The way it came to be identified with Pursh's C-
ovata has been discussed above. It differs from that, however, in the respects
enumerated in the key, and accordingly requires a new name, which is appro-
priately derived from its most unique character, the whitish hairs on the
achene-appendages. It occurs on various shale-barrens, and has been col-
lected from the following :

VIRGINIA.

Allegheny: 1§ miles west of Covington, Wherry, June 10, 1930 (N, P).
Bath: Hot Springs, Hunnewell, May 11 to July 3, 1911 (G)
Southwest of Hot Springs, Wherry, June 10, 1930 (P).

WEST VIRGINIA.

Greenbrier: Kates Mountain, many collectors and dates (G, N, P, U).
This is to be taken as the type locality of the species, and as type specimen
should be designated :

Guttenberg, July 31, 1877 (U).

12 Steele, Contr. U. S. Nat. Herb. 13: 364. 1911.

:

Reprinted from Bartonia, No. 12, 1930

The Eastern Short-styled Phloxes'

Edgar T. Wherry

ft

In last year's number of '* Bartonia ''^ the Phloxes of the
section Suhulatae were discussed as to their history, geog-
raphy, ecology, variation, and cultivation. The second section
to be treated in like manner is that termed by Peter^ the
Divaricatae. Its members differ from those of the former
section in that the stems are scarcely if at all woody, the leaves
are fewer, larger, and less persistent, and the inflorescence is
a more compound cyme of short-pedicelled flowers; in these
the sepals are united for 4 to ^ their length, and the stamens
and styles are much shorter than the corolla-tube. The west-
ern species included in the section by Peter are not treated
here, but four eastern ones are recognized. In the following
synopsis these are arranged in the order of increasing com-
pactness of inflorescence:

PHLOX, SECTION DIVARICATAE: KEY TO SPECIES

Sterile shoots becoming decumbent, often rooting at nodes ; leaves rather
broad, obtusish to acutish; inflorescence lax, glandular; calyx-lobes
short-awned; corolla-tube always glabrous, lobes sometimes notched,

and limb pale violet, ranging to purple or to white 4. P. divaricata

Sterile shoots erect or decumbent, not rooting at nodes; leaves mostly
narrow; inflorescence compact; corolla- tube sometimes pubescent,
lobes never notched, and limb purple, ranging to violet or to white.
Leaves linear to lanceolate, or some nearly ovate, acuminate,
sparingly if at all persistent; bracts spreading, scattered through
the moderately compact cyme; inflorescence-hairs fine or excep-
tionally coarse, sometimes gland-tipped, rarely lacking; calyx-
awns often long.

Nodes few; leaves pubescent or sometimes glabrous, the upper

spreading, passing rather abruptly into bracts 5. P. pilosa

Nodes numerous; leaves always glabrous, the upper ascending,

passing gradually into glandular bracts 6. P. floridana

Leaves oblong-elliptic or sometimes lanceolate, obtusish to acumi-
nate, many of them persistent; bracts ascending, closely sur-
rounding the decidedly compact cyme; inflorescence-hairs coarse,
never gland-tipped ; calyx-awns short 7. P. amoena

1 Presented in abstract, with lantern-slide illustrations, at the meeting
of the Philadelphia Botanical Club on December 19, 1929. Contribution
from the Botanical Laboratory of the University of Pennsylvania.

2 Bartonia 11: 5. 1929.

sin Engler & Prantl's Pflanzenfamilien 43a: 47. 1891.

(24)

V

■

IRREGULAR PAGINATION

Barton I A, No. 12

PHILADELPHIA BOTANICAL CLUB

25

Plate 1

Fig. 1. Phlox (livaricatu canadensis.
Preserve of Washington Biologists' Field Club, Cabin John, Maryland.

Fig. 2. Same s})e('ies, variety, and locality.
Forms with notched, entire, and apiculate corolla-lobes growing together.

I See page 34.]

*

j

4. Phlox divaricata Linn6. Blue Phlox. Plate 1.

History. — This species, like several others, was discovered
in early colonial days, being listed in 1700 by Plukenet^ as
^'Lychnidea Virginiana Blattariae accedens, Alsines aquaticae
foliis, floribus summo caule brachiatis, sive in ramulos divar-
icatis.** In 1739 John Bartram, the pioneer American bota-
nist, collected it along the Susquehanna River in Pennsyl-
vania, and sent it to Peter CoUinson, who catalogued it, ac-
cording to Dillwyn,^ as **Lychnidea foliis lanceolatis obtusis,
flore pallide caeruleo.'* The specimen on which Linne^
founded the species in 1753 bears on the sheet the initials
**H.U/'* signifying that it represented a plant being culti-
vated in the botanical garden at Upsala (Hortus Upsaliensis).
His description contains the erroneous statement that the
lower leaves are alternate, but is otherwise applicable. The
corolla-lobes in the type specimen are conspicuously notched,
although in the earliest known illustration of the species, that
published by Miller^ in 1760, the notches in the lobes are much
shallower, indicating that material had already reached
Europe from more than one source.

In 1796 Salisbury® applied the name Phlox vernalis to what
was probably the same species, but gave no description.
Thirty years later a supposedly distinct plant was described
by Sweet^ as P. canadensis, and this name has been consider-
ably used in horticultural writings, although most botanists
have recognized the differences from the Linnean plant — more
upright habit and lobes ** about half the length of P. divaricata
and much broader'* — to be insufficient to constitute a separate
species. Two other names, P. amplexicaulis Rafinesque and
P. glutinosa Buckley, have been regarded by some authors as
synonyms of P. divaricata, but are here discussed in connec-
tion with P. pilosa,

1 Mantissa 121. 1700.

2 Hortus Collinsonianus 39. 1843.

3 Species Plantarum (1): 152. 1753.

* As shown in a photograph kindly obtained in 1928 by Dr. William
K. Maxon of the U. S. National Herbarium.

5 Figs. Plants Miller's Gardeners' Diet. 2: pi. 205, fig. 1. 1760.

6 Prodromus stirpium, etc. 123. 1796.

7 British Flower Garden 3: pi. 221. 1827.

IJAirroMA, Xo. 12

Plate 1

PHILADELPHIA BOTANICAL CLUB

25

Fi(!. 1. I'lilo.r ili I (iricahi r'nnnh iisi.s.
Pitsi'ivc of \V:is!iiiijrt()ii P>i()l(»^lsts ' Field Club, ('.-ihin .lolm, M.-iivI.-iimI.

Fm. '2. S;iMU' sjK'cit's, \ Miii'tv, -nul lociility.
Forms witli uotclicd, ciitiri', .-lud ;i|ticnl.-itt' corollM-lobcs ^louiiio- tojjctlicr

I Set' p.-i^c ."U. I

' ^ ■

*■ i

-^^ i

,/

1

4. Phlox divaricata Linne. Blue Phlox. Plate 1.

History. — This species, like several others, was discovered
in early colonial days, being listed in 1700 by Plukenet^ as
'^Lychnidea Virginiana Blattariae accedens, ALsines aquaticae
foliis, floribus summo caule brachiatis, sive in ramulos divar-
icatis." In 1739 John Bartram, the pioneer American bota-
nist, collected it along the Susquehanna River in Pennsyl-
vania, and sent it to Peter Collinson, who catalogued it, ac-
cording to Dillwyn,''' as ''Lychnidea foliis lanceolatis obtusis,
flore pallide caeruleo." The specimen on which Linne*
founded the species in 1753 bears on the sheet the initials
*'H.U."* signifying that it represented a plant being culti-
vated in the botanical garden at Upsala (Hortus Upsaliensis).
His description contains the erroneous statement that the
lower leaves are alternate, but is otherwise applicable. The
corolla-lobes in the type specimen are conspicuously notched,
although in the earliest known illustration of the species, that
published by Miller' in 17(30, the notches in the lobes are much
shallower, indicating that material had already reached
Europe from more than one source.

In 1706 Salisbury*' applied the name Phlox vernal is to what
was probably the same species, but gave no description.
Thirty 3'ears later a supposedly distinct plant was described
bv Sweet" as P. canadensis, and this name has been consider-
abh' used in horticultural writings, although most botanists
have recognized the differences from the Linnean plant — more
upright habit and lobes ** about half the length of P. diraricata
and much broader" — to be insufficient to constitute a separate
species. Two other names, P. amplexicaulis Rafinesque and
P. glutinosa Buckley, have been regarded by some authors as
synonj'ms of P. divaricata, but are here discussed in connec-
tion with P. pilosa.

1 Mantissa 121. 1700.

2 Hortus Collinsonianiia 39, 1843.

3 Species Plantarum (1): 152. 1753.

* .A-s shown in a photograph kindly obtained in 1928 by Dr. William
R. Maxon of the U. S, National Herbarium,

•iFigs. Plants Miller's Gardeners' Diet. 2: pi, 205, fig, 1. 1760.
*• Prodromus stirpiuni, ete. 123. 1796.
"British Flower Garden 3: pi. 221. 1827.

INTENTIONAL SECOND EXPOSURE

26

PROCEEDINGS OP THE

PHILADELPHIA BOTANICAL CLUB

27

When explorations came to be made in the Mississippi basin,
the Phlox divaricata occurring there was found to differ some-
what from that farther east, and attempts were made to recog-
nize this by special nomenclature. In enumerating plants
observed in Arkansas territory, NuttalP included a ''P. glom-
erata," with a star to indicate that he regarded it as a new
species, but as he furnished no description the name lacks
validity. A specimen from his original collection, preserved
in the Gray Herbarium, is an immature individual of the com-
mon representative of P. divaricata in that region, which is
characterized by having entire corolla-lobes. The same thing
was collected by Lapham in Wisconsin, and named by Wood^
P. divaricata P ? laphami, with the description ''Lvs. ovate;
pet. obtuse, entire. . . . Intermediate between P. divaricata
and P. glaherrima, and may prove distinct from both. ' ' Gray^
modified Wood's designation to variety laphamii; and many
years later, Clute* held the difference in corolla-lobe outline
to justify the recognition of two distinct species, applying to
the western one the name P. laphamii. To what extent these
different interpretations correspond to the existing relation-
ships will be considered after the variations shown by P. di-
varicata have been described.

Whitish variants occasionally appear in all large colonies
of the Blue Phlox, and the names **P. divaricata alba," *'P.
canadensis alba,'' and even *'P. laphami alba" have long been
in use for such material by horticulturists, although none of
these combinations appear to have been validated by formal
publication. In 1920 FarwelP proposed P. d. forma alhi-
flora for a plant from Michigan, while a few years later
House® applied the same term to a New York occurrence.
FarwelF subsequently furnished data to show that a pallid
rather than a complete albino color-form was represented.
These and other forms will be discussed under Variation.

1 Trans. Amer. Phil. Soc. 5: 196. 1837.

2 Class-book of Botany 439. 1846.

3 Manual of Botany, ed. 2. 331. 1856.

4 Amer. Botanist 25: 101. 1918.

5 Eept. Mich. Acad. Sci. 31 : 369. 1920.

6 Ann. List Ferns & Fig. Plants N. Y. 581. 1924.

7 Amer. Midi. Nat. 11 : 68. 1928.

h

<

Geography. — Phlox divaricata is one of the most wide-
ranging of all the species, occurring in various physiographic
provinces, though rarely in the Coastal Plain and Piedmont,
from western Florida to eastern Texas, southern Minnesota,
and southernmost Quebec. This is shown graphically in fig-
ure 1 by circles representing the counties in which it is known
to have been collected. The terminal moraine of the final
(Wisconsin) glacial stage, marked by a dotted line, has evi-

FiG. 1. Distribution of Phlox divaricata.

dently been ineffective as a barrier; but the mountain front
(row of As) and the Fall-line (cross-hatched band) have
clearly restricted its migration, and it has crossed them only
along a few river valleys. Curving southward from the lower
end of Lake Michigan to the coast of Georgia runs a boundary
between an eastern and a western variety; the significance of
its course remains to be explained.

28

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

29

Alabama : Rather frequent in the upland regions, and rare
in the Coastal Plain, being recorded from Blount', Bul-
lock, Clarke, Etowah', Franklin, Jackson', Jefferson', Madi-
son', Marshall', Morgan', Tuscaloosa, and Wilcox' counties.
The boundary between the areas of the eastern and western
varieties crosses the northeast corner.

Arkansas: Occasional throughout, specimens having been
seen from Benton, Carroll', Garland, Hempstead, Madison',
Miller, Newton', and Washington counties. NuttalPs ''P.
glomerata" was ascribed to this state.

District of Columbia' : Found only on alluvial flats of the
Potomac River in the Piedmont province, toward the north-
west boundary.

Florida: Limited to the northwestern part of the state,
where it grows along the Apalachicola and Chipola rivers in
Gadsden, Jackson', and Liberty' counties.

Georgia: Rare, specimens having been seen only from the
Coosa valley in Floyd and the Ocmulgee valley in Bibb and
Houston counties.

Illinois: Abundant practically throughout, and recorded
from 36 counties: Calhoun, Champaign, Clinton', Cook, Du
Page, Hancock, Henderson, Jackson', Jeiferson, Johnson,
Kane, Kankakee, Knox, Lake', La Salle, Macon, Macoupin,
Marion, Marshall, McHenry, McLean, Menard, Ogle, Peoria,
Piatt, Pulaski', Richland, Rock Island, St. Clair, Stark, Taze-
well', Union', Vermilion, Wabash, Will, and Winnebago. The
boundary between the varieties runs near the eastern edge of
the state.

Indiana : Known in nearly every county, largely as the re-
sult of the collecting activity of Charles C. Deam; this does
not mean that it is more frequent here than in adjacent states,
but merely that they have been less thoroughly covered. The
list is: Adams, Allen, Blackford, Brown, Carroll, Cass, Clark,
Clay, Clinton, Crawford, Daviess, Dearborn, Decatur, De
Kalb, Delaware, Dubois, Elkhart, Fayette, Fountain, Frank-
lin, Fulton, Gibson, Grant, Greene, Hamilton, Hancock,
Harrison, Howard, Huntington, Jackson, Jay, Jefferson,

t Designates counties from which fresh material has been seen.

^y

Jennings, Johnson, Knox, Kosciusko, Lagrange, Lake, La-
porte, Lawrence, Madison, Marion, Marshall, Miami, Monroe,
Montgomery, Morgan, Newton, Noble, Ohio, Orange, Owen,
Parke, Perry', Porter, Posey, Putnam, Randolph, Ripley,
Rush, St. Joseph, Scott, Shelby, Spencer', Steuben, Sullivan,
Switzerland, Tippecanoe, Vanderburg, Vermilion, Vigo, Wa-
bash, Warrick, Washington, Wayne, Wells, and White. A
variant with reduced, acuminate lobes, the * * Crawf ordsville
form, ' ' occurs in Montgomery and Tippecanoe counties.

lowA: Scattered nearly throughout, specimens having been
seen from Decatur, Dubuque, Emmet, Fayette, Floyd, Fre-
mont, Ida, Johnson, Jones, Marshall, Pottawattamie, Po-
weshiek, Scott, Shelby, and Story counties.

Kansas: Known from 20 counties toward the eastern end:
Atchison, Bourbon, Brown, Butler, Chautauqua, Cherokee,
Cowley, Douglas, Geary, Jackson, Leavenworth, Lyon, Mont-
gomery, Morris, Nemaha, Riley, Shawnee, Wabaunsee, Wil-
son, and Wyandotte. The westernmost reported occurrence
of the species is that in Butler County.

Kentucky: Extends throughout, although as less collect-
ing has been done here than in the states adjoining on the
north, the county list is relatively small: Anderson, Boone',
Christian', Estill, Fayette, Franklin, Gallatin', Henry', Jef-
ferson, Jessamine, Kenton', Lyon, Mercer, Morgan, Rockcastle,
Rowan', Shelby, and Warren.

Louisiana : Recorded from 5 parishes along the Mississippi
and Red rivers : Caddo, Feliciana, Natchitoches, Rapides, and
St. Landry.

Maryland: Known only from the basins of the Potomac
and Susquehanna rivers, in Allegany, Anne Arundel, Cecil',
Frederick', Garrett, and Montgomery' counties.

Michigan: Throughout the southern half of the lower pe-
ninsula, there being records from 19 counties:* Allegan',
Benzie, Berrien, Cass, Eaton, Genesee, Grand Traverse,
Gratiot, Hillsdale, Ingham, Ionia, Jackson, Kalamazoo, Kent,
Oakland, Ottawa, Washtenaw, Wayne, and Wexford.

1 Prof. H. T. Darlington kindly furnished data on these from the
State College herbarium, where most of them are represented.

;^

V

30

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

31

Minnesota: In the State University herbarium there are
13 counties represented :^ Chippewa, Dakota, Goodhue, Henne-
pin, Houston, Le Sueur, Meeker, Nicollet, Olmsted, Ramsey,
Scott, Waseca, and Winona. The first is the northwestern-
most point the species is known to have reached.

Mississippi: Occasional in the northern and central parts
of the state, being recorded from Chickasaw, Clay, Hinds,
Lafayette, Oktibbeha, and Tishomingo counties.

Missouri : Wide-spread, specimens having been seen from 25
counties: Atchison, Bates, Butler, Carter, Cass, Christian,
Cole, Cooper, Dunklin, Gasconade, Greene, Iron, Jackson,
Jasper, Jefferson, Knox, Marion, Phelps, Pike, Ralls, St.
Charles, St. Francois, St. Louis', Shannon, and Taney.

Nebraska: Found only in the southeastern corner, the
county records being: Cass, Douglas, Lancaster, Nemaha,

Otoe, and Sarpy.

[New Jersey : In the herbarium of the Academy of Natu-
ral Sciences of Philadelphia there is a specimen labelled Cam-
den, but it is believed to represent cultivated material.]

New York : Common in the western and central portions,
extending to the banks of the Hudson and the western border
of the Adirondacks, in the counties: Broome, Cattaraugus,
Chemung, Erie, Genesee, Greene, Herkimer, Jefferson, Mon-
roe, Oneida, Onondaga, Ontario, St. Lawrence, Steuben,
Tompkins', Ulster, Wyoming, and Yates.

North Carolina: Evidently rare, specimens having been
seen only from the valleys of the Roanoke River in Halifax,
and the French Broad in Madison counties.

Ohio: Frequent throughout, there being records from 50
counties : Adams', Auglaize, Butler, Clark, Clermont, Clinton,
Columbiana, Coshocton', Crawford, Cuyahoga, Darke, Dela-
ware', Erie, Fairfield', Franklin', Gallia, Greene, Hamil-
ton', Hancock, Hardin, Harrison', Highland', Huron, Jack-
son', Knox, Licking, Lorain, Lucas, Medina, Mercer, Miami,
Monroe, Montgomery, Muskingum', Ottawa, Pike', Portage,
Preble, Richland, Ross', Sandusky, Seneca, Shelby, Stark, Tus-
carawas', Vinton, Warren, Washington, Williams, Wyandot.

1 Thanks are due to Prof. C. O. Eosendahl for sending specimens.

<V»

i

A

Oklahoma: In the Missouri Botanical Garden herbarium
there is a specimen from Chelsea, Rogers County.

Pennsylvania: Occurs in all parts of the state, though
rare northward, the county list being: Allegheny', Beaver',
Blair, Bradford, Butler, Cambria', Center, Chester, Clear-
field, Clinton, Dauphin', Delaware, Erie, Fayette, Hunting-
don, Lancaster', Lawrence, Luzerne, Lycoming', Mercer,
Mifflin', Union, Venango, Washington, Westmoreland, York.

South Carolina: In 1817 Elliott* reported this species
with a question from swamps of the Savannah River 40 miles
above Savannah, which would place it in Hampton County,
South Carolina. In his herbarium, preserved at the Charles-
ton Museum, a typical specimen of it is included, although it
is mounted along with a fragment of P. glaherrimay and his
doubt was evidently due to his confusing the two species.

Tennessee: As little collecting has been done here, there
are records only from Anderson, Blount, Davidson, Franklin,
Grainger', Hamilton', Hawkins', Knox', Madison, Marion',
Roane, and Shelby counties. The Madison and Shelby speci-
mens represent the western variety.

Texas: Herbaria include specimens from Liberty and San
Augustine counties, near the eastern edge.

Vermont: Reported from Sheldon, Franklin County.^

Virginia: Apparently rather rare, and known from but 9
counties: Amherst', Botetourt', Campbell, Chesterfield', Fair-
fax', Giles', Rockbridge', Russell, and Washington.

West Virginia : Scattered throughout, except in the higher
mountains, in the counties: Cabell, Doddridge, Fayette', Gil-
mer, Hardy, Harrison, Jefferson', Kanawha, Lewis, Marion,
Mercer, Monongalia, and Ohio.

Wisconsin:^ Rather common below latitude 45° 30', the
county list being: Barron, Brown, Calumet, Columbia, Dane,
Dunn, Green, Iowa, Jefferson, Manitowoc, Marathon, Mil-
waukee, Outagamie, Pepin, Pierce, Polk, Racine, Rock, Rusk,
Sauk, Shawano, Sheboygan, and Winnebago.

1 Sketch Botany S. C. & Ga. 1: 248. [1817.]
2Vt. Agr. Expt. Sta. Bull. 187: 237. 1915.

3 Data for this state were supplied by Mr. A. M. Fuller of the Mil-
waukee Public Museum.

32

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

33

«
«

Canada.^ Ontario: Frequent between Lake Huron and
the St. Lawrence, being known in 14 counties : Carleton, Elgin,
Essex, Frontenac, Grey', Hastings, Huron, Lambton, Lanark,
Lincoln, Peel, Russell, Welland, and York.

Quebec: The original *^ Phlox canadensis' ' was from Mon-
treal, Hochelaga County, this being the northeasternmost
point P. divaricata is known to have reached. There is also
a report from the Ottawa valley in Hull County.

Ecology. — Phlox divaricata is typically a rich woods plant,
growing on alluvial flats, on ravine slopes, and to some extent
on rock ledges, especially when these are calcareous. The soil
may be sandy or clayey, and is usually well supplied with
humus ; in reaction it is normally circumneutral. Though ap-
pearing in various successional stages, this Phlox is most
characteristic of climax associations. Beyond the area where
it is common — from the Appalachians to the Missouri, and
the Blue Ridge to the Great Lakes — it follows the river val-
leys, as is brought out in the map of its distribution. Its
rarity in the Atlantic coast states, where it occurs only along
a few streams that rise far up in the mountains, is particu-
larly striking. In most directions it has evidently reached
the limits to which the climate will permit it to migrate,
although there is no obvious reason why it may not ultimately
cross the Hudson and enter southern New England.

A member of the Spring flora. Phlox divaricata begins to
bloom in the south in March, and in the north in April, con-
tinuing through May into early June. The stigmas lie below
the anthers, but do not become receptive until much of the
pollen has been shed, and the delicate fragrance of the flowers
attracts many kinds of insects, which bring about cross-
pollination; 25 species of moths, butterflies, bees, etc., are
listed by Robertson^ as visiting it.

Variation. — Although on the whole more constant in its
characters than the members of the section Suhulatae, this
Phlox is variable in several respects. The lower leaves range
from narrowly to broadly elliptic, the upper from lanceolate

1 Records from the National Herbarium sent by Dr. M. O. Malta.

2 Flowers and Insects 151. 1928.

4

J

t

4

*l

>

to ovate, within single colonies. Their tips are normally ob-
tusish, but may become acutish toward the inflorescence.

The most striking variability appears in the corolla char-
acters. The common name, Blue Phlox, refers to the fact
that the color of the corolla-limb is bluer than in most other
species, although comparison with the plates in Ridgway^
shows it to be more correctly described as violet. The usual
range is from pale violet (59 d) to light amparo purple
(63 d). West of the Great Lakes and the Mississippi River
bluer hues are dominant, pale bluish violet (57 d) being occa-
sionally approached. Eastward there is a tendency to vary
toward light phlox purple (65 d), most marked in eastern
Kentucky and southern Ohio ; indeed Short,^ in his * * Florula
Lexingtonensis, * ' questioned the identity of the plant there
with P. divaricata because its flowers are * 'frequently rose-
coloured.*' In most colonies all gradations from these colors
to white appear, and occasionally a large patch is whitish
throughout; a particularly striking occurrence of this kind
on the Mackinaw River flats south of Pekin, Tazewell County,
Illinois, was called to my attention by Mr. V. H. Chase, of
Peoria. Eye striae may be present, a single broad stripe often
lying at the base of each lobe, its hue being the same as that
of the rest of the limb, but its tone somewhat deeper. Excep-
tionally these striae are broadened and reddened, coalescing
into a purple ring.

The corolla-tube ranges in length from 10 to 20 mm., and
though usually longer than the lobes, is sometimes exceeded
by them. The lobes are from 9 to 19 mm. long and 5 to 13
mm. wide. West of the varietal boundary shown on the map
(fig. 1) the lobe-ends are normally obtuse and entire, mucro-
nate or apieulate, but only exceptionally definitely notched.
East of it, on the other hand, a well-marked sinus 1 to 3 mm.
deep is commonly present, although in nearly every colony
a few individuals have entire or even apieulate lobes {cf.
plate 1, fig. 2). Rarely the lobes are much shorter than the
tube, and long-acuminate.

1 Color Standards and Nomenclature. 1912.

2 Transylv. Journ. Med. 1 : 415. 1823.

34

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

35

If the boundary line mentioned separated plants of con-
sistently different features, the presence of two distinct spe-
cies, which as noted under History has already been suggested,
might be considered. Since, however, mutants within the
colonies on one side of the line repeatedly imitate the plants
normally found on the other, only varietal distinctness can be
admitted. Correspondingly, no key can be drawn up for the
identification of isolated specimens, because exceptional indi-
viduals would be continually falling into the wrong place.
The plan used with the section Suhulatae is accordingly fol-
lowed here, the varieties being tabulated in the relative posi-
tions they occupy in nature, and the more noteworthy forms
being referred to by English descriptive terms.

VARIETIES AND FORMS
Corolla-lobes normally

ENTIRE; COLOR FRE-
QUENTLY VIOLET.

Phlox divaricata laphami Wood.
P. d. laphamii Gray. P. lap-
hamii Clute. Occurring west of
the varietal boundary described
in the text.

Forms :

White (albino). ''P. laphami
alba^' Hort. An occasional
mutation in many colonies;
sometimes dominant.

Notched. Lobes with a broad
sinus up to 1 mm. deep, imi-
tating the eastern variety.
Typified by specimen in Gray
Herbarium from Franksville,
Racine Co., Wise, collected by
S. C. Wadmond May 18, 1899.
Rare, very few specimens seen.

Mucronate or apiculate-lobed.
Typified by specimen from near
Starkville, Oktibbeha Co., Miss.,
collected by Fannye A. Cook
April 3, 1927, in herbarium of
E. T. W. Frequent, grading
into erose-margined.

Long-acuminate-lobed. Typified
by specimen from 5 miles north
of Huntsville, Madison Co., Ark.,
collected by J. E. Benedict, Jr.
April 16, 1928, in herbarium of
E. T. W. Very rare.

OF PHLOX DIVARICATA

Corolla-lobes normally
notched; color fre-
quently purplish.

Phlox divaricata canadensis

(Sweet) Wherry, status novua
[already in horticultural litera-
ture]. P. canadensis Sweet.
Occurring east of the boundary.

Forms :

White (albino). '^P. divaricata
alba*' or "P. canadensis alba'*
Hort. P. divaricata f. albiflora
Far well. An infrequent mutation.

Entire. Lobes with entire, ob-
tuse terminations, imitating the
western variety. Typified by
specimen from south bank of
James River east of Granite
station, Chesterfield Co., Va.,
collected by E. T. W. April 15,
1927. Fairly common.

Mucronate or apiculate-lobed.
Typified by specimen from f
mile southwest of Cabin John,
Montgomery Co., Md., collected
by E. T. W. May 1, 1930.
Much rarer than in the western
variety, and usually only erose.

Long-acuminate-lobed. Dominant
in a colony near Crawfordsville,
Montgomery Co., Ind., and
widely known as the ''Craw-
fordsville form. ' ' An exception-
al mutation in Lawrence Co., Pa.

./

■i-

Cultivation. — The earliest reference to the introduction of
Phlox divaricata into horticulture is a notation in Collinson's
catalog of his garden, recorded by Dillwyn:^ **A very pale
blue early Lychnidea in flower May 5, 1740, not in England
before ; now in most gardens ; from the Susquehanna River. * *
A letter from Collinson to Bartram, dated June 10, 1740, and
later published by Darlington,^ includes the remark that
** amongst the last things, there is a very pretty Lychnis, with
pale blue flowers, and sweet smell, ^* the same plant evidently
being referred to, for Bartram 's correspondence shows that
he confused the two genera, Lychnis and Lychnidea. In vari-
ous writings, Linne^ replaced the latter name by Phlox, and
in the 6th edition of the Gardeners' Dictionary, Miller* listed
the present species as "3. Phlox foliis lanceolatis obtusis,
floribus majoribus umbellatim dispositis. Begins flowering
in May; flowers large and of sky-blue colour.*'

The horticultural value of this Phlox has been referred to
by numerous subsequent writers, and many dealers now list
it, using various combinations of the names divaricata, cana-
densis, laphami or laphamii, alba, etc. The variety with
violet (''blue") entire-lobed corollas, the correct name for
which is P. divaricata laphami, is on the whole the more
showy plant, some splendid vigorous strains of it having been
developed. That with notched lobes and more purplish (lilac)
hue, P. d. canadensis, is usually relatively delicate, but also
has considerable merit from this standpoint.

Phlox divaricata has been used as one parent of a series of
hybrids which are offered by some nurseries under the name
** Phlox arendsii.'' In this case the tall, late-blooming
P. paniculata was the second parent, and the hybrids are in-
termediate in habit, flower color, and blooming time between
the two. Garden hybrids of P. divaricata with other species
are probably in the trade to some extent, but appear to have
received no distinctive names.

1 Hortus Collinsonianus 39. 1843.

2 Memorials of Bartram and Marshall 136. 1849.

3 Genera Plantarum 52. 1737; Hortus CliflFortianus 63.

4 Gardeners' Diet., ed. 6. 1752.

1737.

* K. »

36

PROCEEDINGS OF THE

]>ART()XIA, Xo. 12

Plate 2

5. Phlox pilosa Linn^. Downy Phlox. Plate 2.

History. — Of the three figures of Phlox published by
Plukenet^ in 1691, two represented members of the Suhulatae,
and have been discussed in the paper on that section. The
other, described as **Lychnidaea umbellifera Blattariae ac-
cedens Virginiana major, repens, Pseudo-melanthii foliis
pilosis, flore albo, pentapetaloide, fistuloso,'' belongs to the
Divaricatae. What is clearly the same species was listed a
few years later by Ray^ as **Lychnoides Marilandica, Jasmini
flore quinquepartito, calycibus lanuginosis, foliis angustis
acutis.'' When Linne^ came to characterize the species of
Phlox, no specimen of this was at hand, so he cited the above
references, and based the diagnosis of P. pilosa upon them.
The specimen from which Plukenet^s figure was evidently
drawn is preserved, however, in the Sloane herbarium at the
British Museum, and in 1928 it was kindly compared by Mr.
A. J. Wilmott with some eastern American material taken to
England by Dr. William R. Maxon. There proved to be
agreement with a recently collected Virginia specimen in leaf
characters, glandular calyx, and puberulent corolla-tube, thus
establishing the proper application of the Linnean name.

During his famous explorations in North America, Michaux*
encountered three of the Phloxes of the section Divaricatae.
He succeeded in recognizing one correctly as Linnets P. di-
varicata, but mistakenly applied the name P. pilosa to the
species here taken up last, proposing for the present one a
new name, P. aristata, with two varieties, a virens (purple-
flowered) and (3 canescens (rose- and white-flowered). His
usage was followed by various subsequent botanists, but was
recognized to be erroneous by Gray^ in 1856, and in recent
years the name aristata has fallen into disuse. Michaux's
specimens are preserved in Paris, and photographs of them
obtained for me in 1928 by Mr. Ivar Tidestrom, of the Bureau
of Plant Industry, confirm the correctness of Gray^s view.

1 Phytographia pi. 98, fig. 1. 1691.

2 Supplem. Hist. Plant. 490. 1704.

3 Species Plantarum (1): 152. 1753.

* Flora Boreali- Americana (1): 144. 1803.
5 Manual of Botany, ed. 2. 331. 1856.

< ^ r»>^

^

Fig. 1. I'hios pilosa virms.
Five \uUqs northwest of Cottoiivilh'. Marshall Coniitv, Alabama.

Fig. 2. P. pilosa ihtonsa.
Hillsborough Cotinty, Fh)ri«la.

Fig. ;J. P. floridana.
Leon Co., Florida (W. Kiirz, plioto).

36

PROCEEDINGS OP THE

liAinOMA. \(). r_*

I'lati; i!

5.

Phlox pilosa Linne. Downy Phlox. Plate 2.

History. — Of the three figures of Phlox published by
Plukenet^ in 1691, two represented members of the Suhulatae,
and have been discussed in the paper on that section. The
other, described as ''Lychnidaea umbellifera Blattariae ac-
cedens Virginiana major, repens, Pseudo-melanthii foliis
pilosis, flore albo, pentapetaloide, fistuloso," belongs to the
Divaricatae. What is clearly the same species was listed a
few years later by Ray^ as ''Lychnoides MarilancUca, Jasmini
flore quinquepartito, calycibus lanuginosis, foliis angustis
acutis.'' When Linne^ came to characterize the species of
Phlox, no specimen of this was at hand, so he cited the above
references, and based the diagnosis of P. pilosa upon them.
The specimen from which Plukenet's figure was evidently
drawn is preserved, however, in the Sloane herbarium at the
British Museum, and in 1928 it was kindly compared by Mr.
A. J. Wilmott with some eastern American material taken to
England by Dr. William R. Maxon. There proved to be
agreement with a recently collected Virginia specimen in leaf
characters, glandular calyx, and puberulent corolla-tube, thus
establishing the proper application of the Linnean name.

During his famous explorations in North America, Michaux*
encountered three of the Phloxes of the section Divaricatae.
He succeeded in recognizing one correctly as Linne 's P. di-
varicata, but mistakenly applied the name P. pilosa to the
species here taken up last, proposing for the present one a
new name, P. aristata, w4th two varieties, a virens (purple-
flowered) and 3 canescens (rose- and white-flowered). His
usage was followed by various subsequent botanists, but was
recognized to be erroneous by Gray^ in 1856, and in recent
years the name aristata has fallen into disuse. Michaux's
specimens are preserved in Paris, and photographs of them
obtained for me in 1928 by Mr. Ivar Tidestrom, of the Bureau
of Plant Industry, confirm the correctness of Gray's view.

1 Phytographia pi. 98, fig. 1. 1691.

2 Supplem. Hist. Plant. 490. 1704.

3 Species Plantarum (1): 152. 1753.

-* Flora Boreali- Americana (1): 144. 1803.
fi Manual of Botany, cd. 2. 331. 1856.

f

^ N

(

I }

f ' »

'0h

Fi<;. 1. I'lifn.r itili'.sii iiiiii.^.
V'wr milc^ iKiit liw ('>t <»f ( '<»t tnii\ i Mr. .M;ir<li:ill <'«miiii\. A I.iIi.-imi.i.

Kl'i. L'. /'. fiilnsfi <h foiisd.

Hill««lnti(iii^li (■(•iiiity. rinri»|;i.

I'k;. ;.. /'. flnriihum.

I II < '<>.. I'ImiJ.I;! I I . K 111/, plmt !.

.<•

l»

INTENTIONAL SECOND EXPOSURE

(

PHILADELPHIA BOTANICAL CLUB

37

i

In 1817 Rafinesque^ proposed for **a real Phlox akin to
P. pilosa/' observed by Robin in Louisiana, the name P. am-
plexicaulis. Most authors have ignored this or indicated its
identity to be unknown, but Brand^ assigned it, with a ques-
tion, to P. divaricata. The characterization, "foliis amplexi-
caulis lineari subulatis rigidis; panicula confertiflora, calyx
striatus pilosus, dentibus ovatis acuminatis, ' * sounds like a
composite of several species, but comes as close to describing
the southern Mississippi basin variety of P. pilosa as could be
expected. His name is accordingly here taken up for this
variety, although the feature to which it refers is not really
distinctive.

The next name to be considered is Phlox glutinosay applied
by Buckley^ in 1843 to an occurrence at Black *s Bend, Wil-
cox County, Alabama, differing from P. '^aristata'* {pilosa)
in having fewer and taller stems, broader leaves which are
mucronate instead of long-acuminate and are covered with
gland-tipped hairs, and bright red or scarlet flowers. Buck-
ley *s own specimen of this is at the Missouri Botanical Gar-
den, but the type is in the Gray Herbarium. Gray, who
seemed to take delight in criticizing Buckley, wrote on the
latter sheet ** Surely flowers not 'red or scarlet.' Dr. Tor-
rey's four specimens from Buckley plainly show this to be a
form of P. divaricata.'^ He added similar remarks to Tor-
rey's sheet, which is preserved at the New York Botanical
Garden, and reaffirmed his view^ in his writings ; but he over-
looked the long awns on the calyx-lobes and the pubescence
on the corolla-tube, which, together with the foliar characters,
point unmistakably to its being a hybrid of P. divaricata with
P. pilosa. In the hope of obtaining fresh material for study,
I have visited the type region on three different occasions, but
since Buckley 's day the woods have been extensively cut over,
and although P. pilosa is locally abundant, and P. divaricata
sometimes growls near it, the search for the hybrid has been
unsuccessful.

1 Florula Ludoviciana 36. 1817.

2 In Engler 's Pflanzenreich IV. 250
8 Amer. Journ. Sci. 45: 177. 1843.

72. 1907.

I^MaObMUbMU

38

PROCEEDINGS OF THE

Phlox cuspidata, applied to a Texan plant by Scheele^ in
1850, is sometimes regarded as synonymous with P. pilosa, but
the description fits better P. drummondii Hooker.

In revising the Polemoniaceae in 1870, Gray^ assigned the
glabrous-leaved phases of P. pilosa, such as are especially
common in the Gulf States, to a variety detonsa. This was
raised to species rank by Small ;^ but it grades too freely into
the wide-spread variety of P. pilosa to be so maintained.

One more name remains to be considered, and in order to
account for the interpretation of it adopted, the northwestern
representative of the species must be referred to at this point.
Over much of its range, P. pilosa has its inflorescence strongly
glandular, with only occasional glandless variants. On the
prairies of the upper Mississippi basin, however, the hairs on
the inflorescence are normally glandless, dense, and lustrous,
constituting a well-marked geographic variety. In describing
their P. argillacea, Clute and Ferriss* contrasted it with this
prairie Phlox, noting it to have the calyx and bracts glandular-
hispid, and emphasizing its * 'lighter green leaves, greater
height, less compact flower clusters, restricted habitat, and
above all . . . pale flowers and later and longer season of
bloom.'' Most of these, however, are the very respects in
which Linne's P. pilosa differs from the prairie variety.
Plants with pallid or near-white corollas are common in colo-
nies of P. pUosa,^ and indeed the Plukenet specimen, which is
taken as the type of the species, was one of these. When the
Illinois ^'P. argillacea'^ is grown beside such Virginian
material, the only way in which they can be distinguished is
by the greater vigor of the former, a difference of horticul-
tural but not taxonomic significance. It can only be con-
cluded that technically P. argillacea Clute and Ferriss is
synonymous with P. pilosa Linne.

1 Linnaea 23 : 139. 1850.

2Proc. Amer. Acad. Arts Sci. 8: 251. 1870.

3 Flora Southeastern U. S. 978. 1903.

4 Amer. Botanist 17: 74. 1911.

5 The combination P. pilosa forma alhiflora has been published by
Macmillan (Metasp. Minn. Valley 432. 1892) but applies to the albino
form of the prairie variety.

%

PHILADELPHU BOTANICAL CLUB

39

< u

\

V

*

0- '

i

\

k

.H

" " y

1

Geography. — The range of Phlox pilosa extends farther
west and northwest, but not so far northeast, as that of
P. divaricata, comprising all provinces except those at the
highest elevations, from Florida to central Texas, southern
Manitoba, and southwestern Connecticut. Figure 2 presents
the details as to the four varieties here considered.

Fio. 2. Distribution of Phlox pilosa.

The curiously patchy distribution shown in many parts of
this map can not as yet be fully accounted for, but the indi-
cations are that the species has migrated out from a dispersal-
center in the Ozark region. None of the physiographic
boundaries traced on the map appear to have restricted it, but
its absence from the Appalachians is evident.

k

40

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

41

Alabama: Common except in the higher mountains, there
being records from 24 counties: Autauga', Baldwin, Bibb',
Butler', Clarke, Cullman, Dallas, De Kalb', Fayette', Frank-
lin', Hale', Jackson', Lawrence, Lee, Madison', Marengo',
Marion', Marshall', Mobile', Montgomery, Perry', Tuscaloosa',
Washington, and Wilcox'. Both wide-spread (virens) and
Gulf Coast (detonsa) varieties are represented.

Arkansas: Probably occurs in most parts of the state,
although specimens have been seen from but the following 13
counties : Benton, Boone', Carroll', Faulkner, Garland, Greene,
Hempstead, Howard, Lawrence, Logan, Ouachita, Prairie, and
Washington'. A broad-leaved variety or possibly distinct
species also occurs locally; it is to be discussed in a future
paper dealing with the Phloxes of the Ozark and Texan

regions.

Connecticut: The northeasternmost known stations for

the species are in New Haven County.

Delaware: Thus far observed only in New Castle, the
northernmost county in the state.

District of Columbla.:' Has been collected from two sta-
tions toward the northwest side.

Florida: Both wide-spread and Gulf Coast varieties are
known, the county list being : Alachua, Columbia, Dixie, Her-
nando', Hillsborough', Jackson, Jefferson', Lake, Leon', Lib-
erty', Madison', Pasco, St. Johns, Suwannee', Volusia,
Wakulla, and Walton.

Georgia: Apparently rather rare, having been collected
only in Cobb', Dougherty, Floyd, Meriwether, Muscogee,
Peach, Spalding, Walker', and Whitfield' counties.

Illinois : Common and abundant, there being records from
36 counties: Champaign, Christian, Clay, Cook', Du Page,
Fulton, Hancock, Henderson, Henry, Jackson, Jefferson, Jo
Daviess, Kane, Kankakee, Lake', Livingston, Macon, Ma-
coupin, Madison, Marion, Marshall, McHenry, McLean,
Menard, Ogle, Peoria, Piatt, Richland, Rock Island, St. Clair,
Stark, Stephenson, Tazewell, Wabash, Will, and Winnebago.
The wide-spread and prairie (fulgida) varieties are about
equally developed.

f*::^

y

1

Indiana : Less frequent than P. divaricata, even Mr. Deam 's
thorough collecting having turned it up in but 31 counties:
Allen, Benton, Cass, Clark, Elkhart, Floyd, Fulton, Harrison,
Jasper, Kosciusko, Lagrange, Lake, Laporte, Marshall, Mont-
gomery, Newton, Noble, Perry', Porter', Posey, Pulaski, St.
Joseph, Spencer', Starke, Steuben, Tippecanoe, Tipton, Vigo,
Warren, Wells, and White. The southern Mississippi basin
variety extends into Perry and Spencer counties.

lowA: The prairie variety is known in 12 widely scattered
counties: Carroll, Crawford, Decatur, Dickinson, Dubuque,
Emmet, Fayette, Jones, Pottawattamie, Poweshiek, Story, and
Winneshiek. The wide-spread variety is reported from John-
son County.

Kansas: Recorded from Atchison, Brown, Chautauqua,
Cherokee, Cowley, Douglas, Johnson, Labette, Linn, Mont-
gomery, Riley, and Shawnee counties. Both wide-spread and
prairie varieties are represented.

Kentucky : Apparently rare, being known from but 6 coun-
ties: Bath, Christian', Fayette, Henderson, Jefferson, and
Morgan. In addition, Short distributed specimens labelled
*' barrens of Kentucky," which may have come from Barren
or Logan County. The southern Mississippi basin variety is
present locally.

Louisiana: Collected from 9 parishes: Acadia', Caddo',
Calcasieu, Feliciana, Grant, Natchitoches, Orleans, Rapides,
and St. Tammany. Two varieties are included.

Maryland : The observation of this species in Maryland in
early colonial days was mentioned under History. In subse-
quent times it has been largely destroyed by agricultural de-
velopment, but specimens are preserved from Baltimore and
Montgomery' counties.

Michigan : Scattered in the southern part, the county list
being: Cass, Ingham, Ionia, Jackson, Kent, Livingston, Oak-
land, St. Joseph, Van Buren, and Washtenaw.

Minnesota: Occurs throughout except northward, speci-
mens having been seen from 34 counties: Becker, Brown,
Carver, Cass, Chippewa, Dakota, Douglas, Faribault, Fill-
more, Goodhue, Hennepin, Houston, Hubbard, Jackson, Kana-

42

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

43

bee, Kandiyohi, Lincoln, MeLeod, Martin, Meeker, Mille Lacs,
Morrison, Nicollet, Olmsted, Otter Tail, Pope, Ramsey, Rice,
Rock, Scott, Stearns, Wabasha, Waseca, and Winona. Only
the prairie variety is present.

Mississippi : The commonest Phlox in the state, there being
19 county records : Choctaw, Clay, Forrest, Franklin, Grenada,
Hancock, Harrison, Holmes, Itawamba', Jackson, Jasper,
Lafayette', Marshall', Noxubee, Oktibbeha', Pontotoc', Scott,
Tippah, and Tishomingo. Both wide-spread and Gulf Coast
varieties are included.

Missouri : Specimens of the varieties covered in this article
have been seen from 34 counties: Atchison, Bates, Benton,
Butler', Carter, Cass, Christian, Cole, Crawford, Greene,
Iron, Jackson, Jasper, Jefferson, Laclede, McDonald, Madison,
Nodaway, Phelps, Pike, Platte, Putnam, Ralls, St. Francois,
St. Louis', Scott, Shannon, Stone, Taney, Vernon, Washing-
ton, Wayne, Webster, and Wright.

Nebraska: The prairie variety occurs in the eastern part
of the state, in Cass, Dixon, Douglas, Lancaster, Richardson,
and Saunders counties.

New Jersey: Occasional north of the Pine-barrens, there
being 6 county records : Burlington, Essex, Morris, Somerset,
Union, and Warren'.

New York : The only specimen seen from the state is one in
the Gray Herbarium from Niagara Falls, Niagara County.

North Carolina : Apparently rare, and reported from but
4 counties : Forsyth, Halifax, New Hanover', and Polk.

North Dakota : The prairie variety is recorded from Cass,
Grand Forks, and Richland counties, along the eastern border.

Ohio: Known in 17 counties, chiefly northern and south-
central : Athens, Cuyahoga, Defiance, Erie, Fairfield, Frank-
lin, Fulton, Hancock, Henry, Hocking, Lawrence, Lucas,
Madison, Ottawa, Summit, Wood, and Wyandot.

Oklahoma: Scattered throughout the eastern part, speci-
mens being preserved from 9 counties: Creek, Delaware',
Johnston, Le Flore, Logan, Mayes', Murray, Muskogee, and
Oklahoma'. All represent the wide-spread variety.

^ I '• »

Pennsylvania: Restricted to the western edge and the
southeast corner, in 9 counties : Beaver, Bucks, Chester, Dela-
ware, Franklin, Lancaster', Lehigh, Montgomery', and North-
ampton'.

South Carolina: Rare, specimens having been seen only
from Berkeley, Clarendon, Florence, Richland, Sumter, and
Williamsburg counties.

South Dakota: The prairie variety occurs in a few east-
ern counties : Brookings, Clay, Lincoln, and Roberts.

Tennessee: Scattered throughout, specimens being pre-
served from Chester, Davidson, Fayette, Franklin, Hamilton,
Knox, and Madison counties. The southern Mississippi basin
variety is represented in the second.

Texas: Wide-spread east of longitude 98° 30', there being
records for 24 counties: Bexar, Cherokee, Comal, Comanche,
Cooke, Dallas, Erath, Gregg, Grimes, Hardin, Harris', Lamar,
Liberty, Montague, Montgomery, Parker, Polk, Rusk, San
Augustine, Smith, Tarrant, Titus, Walker, and Wood. Be-
sides the varieties here discussed, another occurs westward.
Its distribution and relationships will be treated in a subse-
quent paper on phloxes of the south-central states.

Virginia: Although the type locality of the species was in
this state, it is not common there, being definitely known from
but 6 counties: Augusta, Botetourt, Chesterfield', Fairfax',
Frederick, and Spotsylvania.

Wisconsin : Common throughout, the county records being :
Barron, Bayfield, Brown, Buffalo, Burnett, Clark, Columbia,
Crawford, Dane, Dunn, Grant, Green, Jackson, Jefferson,
Juneau, Kenosha, Lafayette, Milwaukee, Monroe, Pierce,
Portage, Racine, Richland, Rock, St. Croix, Sauk, Shawano,
Sheboygan, Trempealeau, Walworth, Washburn, Washington,
Waukesha, Waupaca, Waushara, Winnebago, and Wood. The
wide-spread variety barely enters the southeastern corner.

Canada. Manitoba : The prairie variety extends to Winni-
peg, in Iberville County.

Ontario : Limited, so far as known, to Essex and Lambton
counties, at the southern end.

•"^

44

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

45

Ecology. — Phlox pilosa grows in many sorts of habitats, —
fields, rocky slopes, open woods, and even exceptionally
swamps. It seems to prefer subacid soils. Occasional as a
pioneer, it increases in abundance rapidly as later successional
stages develop, but thins out again as climax forest conditions
are approached. It is more xerophytic than P. divarieata,
and extends westward a short distance into the semi-arid
parts of Texas. Since the ice of the last glacial stage re-
treated it has migrated northward at least 350 miles (550
km.), corresponding to an average rate of a mile per hundred
years.

The distribution map, fig. 2, suggests the species as a whole
to have had a dispersal center in the south-central part of the
United States, perhaps in the Ozark region, from which the
several varieties spread out in different directions. On the
north and west, climatic barriers have no doubt been reached,
but such can hardly be the case toward the northeast, and if
man does not prevent, the plant may gradually extend its
range there.

In the south Phlox pilosa begins to bloom about the first of
April, further north in early or even late May. The duration
of flowering varies widely from one colony to another, being
at times two or three weeks, or again as many months. As in
most Phloxes, the stigmas ripen after the anthers. The
flowers of some of its races are even more fragrant than those
of P. divarieata, the scent resembling that of Azalea viscosa
or Dianthus spp. Robertson observed twenty insect visitors,
chiefly butterflies and long-tongued bees.

Variation. — The variability of Phlox pilosa is so marked
that the tendency on the part of some students to split it into
several species is easily accounted for. Field study through-
out its range shows, however, that all gradations between the
extremes exist, so that only a separation into varieties is
justifiable.

Most frequently the plants are pubescent throughout, but
locally there may be a tendency toward loss of hairs. This
appears first on the lower leaves, then on the upper ones,
and finally on the stem, the inflorescence usually (though not

^:>

>4f Jl N

;\

% 4

invariably) retaining its pubescence intact. More or less
glabrous-leaved plants may occur in any part of the range, as
isolated individuals in the midst of numerous normal ones,
but in certain regions they increase in frequency. Thus, in
both Illinois and Indiana colonies are known in which pubes-
cent and glabrate plants are roughly equal in number. The
climax of this phenomenon appears, however, along the Gulf
Coast from Mississippi to Florida, where many colonies are
made up entirely of glabrous-leaved individuals, leading to
the recognition there of a distinct variety. Exceptionally the
relative hairiness of the upper and lower parts of the plant
may be reversed, — the leaves be pubescent but the inflo-
rescence glabrous.

Over the greater part of the range the hairs on the inflo-
rescence are conspicuously gland-tipped, while those on the
lower part of the plant are glandless. Occasionally, however,
glands extend down the stem, and sometimes even over the
leaves. This feature reaches its maximum development in
the Ozark region, where in some colonies the plants are viscid-
glandular practically throughout. The opposite tendency,
toward disappearance of glandularity, is more wide-spread.
In the midst of many colonies of plants with glandular inflo-
rescence occasional mutants occur in which the hairs lack
glands entirely. Colonies made up wholly of glandless plants
are not known in the eastern states, but in the Mississippi
basin they are frequent, and northward become dominant.
The inflorescence-hairs on such plants vary considerably,
tending to be coarser in the south.

The normal leaf-outline of Phlox pilosa is lanceolate-
acuminate, but deviations are not infrequent. Broadening of
at least the upper leaves to an ovate shape may occur in any
colony. Narrowing to linear outline is most marked along
the Gulf Coast. The calyx-lobes vary from subulate to
broadly linear, without evident correlation with leaf-outline.

Phlox-purple is the typical corolla-color in this species, but
it is by no means constant. Pallid color-forms are occasional
in all colonies, and dominant in some, w^hile complete albinos
are not infrequent. A light violet or lavender hue similar

46

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

47

to that of P. divarieata appears locally, and in certain colonies
seen in the uplands of Alabama purple, violet, and white
forms were about equally abundant. A pale eye with more
or less prominent superimposed striae is often developed.
Sometimes there is a single central broad stripe on each lobe,
of similar hue to the outer portions ; more often there is a pair
of lateral ones, either redder or bluer, and occasionally both
occur together. These striae may broaden and coalesce into
a deep-colored ring, which is especially striking in otherwise

pallid corollas.

In corolla-dimensions this species is rather less variable than
P. divarieata, and no geographic segregation in this feature
can now be recognized. Usually a whole colony is character-
ized by having the corollas all relatively small, or all large.
The lobes range from cuneate to orbicular-obovate, and from
terminally obtuse to mucronate or apiculate. The corolla-
tube is often pubescent, but may be glabrous on individual
plants or throughout whole colonies.

On the opposite page the well-marked varieties^ and forms
which it seems possible to recognize in the midst of this vast
series of variations, are tabulated according to the ** geo-
graphic'' plan already used. The wide-spread variety is
regarded as ancestral to the others not only because these
occupy more restricted areas, but also because mutations
which appear within colonies of the wide-spread one imitate
or approach all of the other varieties.

Cultivation.— Phlox pilosa is little known in cultivation,

probably largely because in rich garden soils it has proved to

be short-lived. The vigorous race from the southern end of

Lake Michigan, **P. argillacea, ' ' seems, however, well adapted

to garden conditions, and has received favorable comment in

horticultural circles. Other varieties and forms deserve more

attention from this standpoint.

1 Only varieties known to grow east of the Mississippi Eiver are here
included. Additional ones occur further west, one with foliaceous calyx-
lobes (P. aspera E. Nelson) in Texas, and others with often very broad
leaves in Arkansas, Louisiana, and Missouri. These will be discussed m
a later article.

/;

^j*

^i.ii^

4i^

?

VARIETIES AND FORMS OF PHLOX PILOSA

Intlorescence-hairs glandless,

riNE; CALYX-LOBES BEOADISH.

Phlox pilosa fulgida Wherry, var.
nov. Named from the lustrous
inflorescence-hairs. Typified by
specimen in U. S. National Her-
barium from Fort Snelling,
Hennepin Co., Minn., collected
by E. A. Mearns June 12, 1891.
Dominant on the prairies of the
upper Mississippi basin.

Form:

White (albino). P. pilosa forma
alhiflora MacMillan. Rare.

Usual features: leaves pubescent with glandless hairs, inplorks-
cence-hairs glandular, calyx -lobes narrow and lonq-

AWNED, and COROLLA-TUBE PUBESCENT.

Phlox pilosa vlrens (Michaux) Wherry, comb. nov. P. aristata virens
Michx. The widespread variety: Tex. to Fla., northward to Kans.,
SE. Wise, and SW. Conn., avoiding the highest elevations.

Forms :

Glabroua-leaved. Typified by specimen from li miles southwest of
Knox, Starke Co., Ind., collected by C. C. Deam May 30, 1929, in her-
barium of E. T. W. A variation, most frequent in 111. and Ind.

Glandless (as to inflorescence-hairs). Typified by specimen from li
miles north of Zieglersville, Montgomery Co., Pa., collected by E. T. W.
May 26, 1930. An occasional mutation throughout.

Glabrous- tubed. Typified by specimen from 1^ miles east of Hunts-
ville, Madison Co., Ala., collected by E. T. W. April 29, 1929. A
rare mutation northward, but often dominant in southern colonies.

Light-colored (pink, pale violet, white, etc). P. pilosa Linn6 as to
type citation from Plukenet; P. aristata canescens Michx. P. argil-
lacea Clute and Ferriss. Common and locally dominant.

Leaves consistently glabrous ;

inflorescence much as in

widespread variety.

Phlox pilosa detonsa Gray. P.
detonsa Small. Common near
the Gulf coast, Fla. to Miss, (re-
ports farther W. refer to prec).

Form:

Glabrous-sepalled. Typified by
specimen from 3 miles east of
Hopewell station, Hillsborough
Co., Fla., collected by E. T. W.
April 18, 1930. Known in Fla.
only.

INFLORESCENCE-HAIRS GLANDLESS,

COARSE; CALYX-LOBES OFTEN

RATHER BROAD.

Phlox pilosa amplexicaulis (Raf-
inesque) Wherry, status novus.
P. amplexicaulis Raf. Rare, E.
Tex. to La. and Tenn. to S. Ind.

Form :

Glabrous-sepalled. Typified by
specimen in U. S. National Her-
barium from near Covington,
St. Tammany Parish, La., col-
lected by Bro. G. Arsene April
12, 1920. Known in La. only.

The following hybrids have been
P. divarieata canadensis x P. pilosa
P. divarieata canadensis x P. pilosa
P. divarieata laphamix P. pilosa vi

observed :

amplexicaulis. Ind.f
virens. Ala./
rens. (P. glutinosa Buckley.)

48

PROCEEDINGS OF THE

6. Phlox floridana Bentham. Florida Phlox.i Plate 2, fig. 3.
History. — This species was first collected by Chapman, and
named by Bentham^ in 1845. Shuttleworth distributed speci-
mens of it under the name ''P. rigida/' first published by
Brand." In the Kew Index* '*P. nuttalli'' or * * nuttalliana "
Hort. and '*P. Carolina'' Sweet^ are referred to P. floridana,
but judging from Sweet's plate really belong to P. maculata.
Geography. — Specimens have been seen only from counties
indicated by dots on the accompanying map, figure 3, showing
the species to be endemic in a rather small area.

Alabama : Reported from
3i<j; several localities by Mohr,^
but his specimens so labelled
all represent forms of P. pi-
losa. In 1925, however, I col-
lected it at Ozark, Dale' Co.
Florida: The county list
is: Alachua, Gadsden, Gil-
christ, Hamilton, Hernando',
Jefferson, Lafayette, Leon',
Liberty, Suwannee, Taylor,
Walton, and Washington.
Georgia: In Thomas Co.

Fig. 3. Distribution of
Phlox floridana.

Ecology. — The Florida Phlox grows chiefly in open oak-pine
woods on dry sand-clay soil of subacid reaction. Whether its
restricted range is due to lack of hardiness or to some defect
in its means of dispersal has not been ascertained. It blooms
about a month later than the related P. pilosa in the same
regions, — during May and June.

Variation. — Apparently more uniform than che other mem-
bers of the section.

Cultivation. — Reports of Phlox floridana being in cultiva-
tion have proved to be based on mistaken identification.

1 For data on this species I am indebted to Professor Herman Kurz
of the State College for Women at Tallahassee, Florida.

2 In De Candolle's Prodromus 9: 304. 1845.

3 In Engler 's Pflanzenreich IV. 250 : 69. 1907.

4 Index Kewensis (3): 500. 1894.

5 British Flower Garden 3 : pi. 190. 1826.

6 Plant Life of Ala. 685. 1901.

4 >

Barton I A, No. 12

Plate )}

PHILADELPHIA BOTANICAL CLUB

49

Fig. 1. Phlox amocna waiter i.
Six miles north of Jeniison, Chilton County, Alabama.

f.^>

>i

Fig. 2. Phlox amocna valtcri.
In cultivation; originally from North Carolina.

7. Phlox amoena Sims. Hairy Phlox. Plate 3.
History. — This species is recorded to have been discovered
by Fraser in 1786, but it was not named until 24 years later.
Meanwhile Walter^ had listed it with a question, and Michaux^
without question, as P. pUosa Linne. In 1810 Sims^ pub-
lished a colored plate of it and ventured to give the name now
accepted, though considering that **This species of Phlox is
too nearly allied to the one figured in the preceding plate,''
which was the real P. pilosa., Pursh* endeavored to reach a
compromise by proposing the combination P. pUosa variety
amoena. Elliott,^ on the other hand, returned to Michaux's
usage, while Bentham^ overlooked Sims's name entirely. In
the second edition of his Manual, Gray' termed the present
plant P. pilosa var. ? walteri, and retained this designation
in the following two editions. Chapman^ was satisfied that
the two are distinct species, and made for the present one the
new combination P. walteri; and Gray® then came to the same
view, identifying it at first with P. procumhens Lehmann,^°
but soon afterward^^ recognizing this as a hybrid, and adopt-
ing Sims's name, as required by the rule of priority. In the
last cited work, Gray also noted that Nuttall had proposed
the name P. involucrata, but had never adequately pub-
lished it.

The only other date of significance in the history of P.
amoena is 1903, when SmalP^ segregated from it a presum-
ably distinct species, under the name P. lighthipei. This was
reduced to varietal status by Brand" (mis-spelled lightipei),
and reasons for accepting his view will be presented in the
discussion of variation in the species.

1 Flora Caroliniana 96. 1788.

2 Flora Boreali- Americana (1): 144. 1803.
8 Botanical Magazine 32: pi. 1308. 1810.
♦Flora America Sept. (1): 150. 1814.

5 Sketch Botany S. C. & Ga. (1) : 247. [1817.]

6 In De Candolle^s Prodromus 9: 305. 1845.

7 Manual of Botany, ed. 2. 331. 1856.

8 Flora Southern States 339. 1860.

9 Manual of Botany, ed. 5. 372. 1868.

10 Index Sem. Hamburg 1828: 17; Linnaea 5: 383. 1830.

11 Proc. Amer. Acad. Arts Sci. 8: 251. 1870.

12 Flora Southeastern U. S. 978. 1903.

13 In Engler's Pflanzenreich IV. 250: 70. 1907.

liAUTOXIA, Xo. 12

Plate :\

PHILADELPHIA BOTANICAL CLUB

49

■k '■*.

Fi(i. 1, Phlo.r <n)K>rn(i ualtcri.
Six iiiik's north of .loinisctii, Chilton County, Alnbnnin.

1

r

4

r

Fl(i. L'. I'lilo.)' amoi nil iniHrri.
In cult iv;i1 ion : ori«;iu:illy t'i(uu N(»rtli Cjirolina.

7. Phlox amoena Sims. Hairy Phlox. Plate 3.
Histonj.— This species is recorded to have been discovered
by Fraser in 1786, but it was not named until 24 years later.
Meanwhile Walter^ had listed it with a question, and Michaux^
without question, as P. pilosa Linne. In 1810 Sims"* pub-
lished a colored plate of it and ventured to give the name now
accepted, though considering that ''This species of Phlox is
too nearly allied to the one figured in the preceding plate,"
which was the real P. pilosa. Pursh^ endeavored to reach a
compromise by proposing the combination P. piloaa variety
amoena. Elliott,^ on the other hand, returned to Michaux's
usage, while Bentham^ overlooked Sims's name entirely. In
the second edition of his Manual, Gray' termed the present
plant P. pilosa var. ? walteri, and retained this designation
in the following two editions. Chapman" was satisfied that
the two are distinct species, and made for the present one the
new combination P. walteri; and Gray-^ then came to the same
view, identifying it at first with P. jyrocumhens Lehmann,^*'
but soon afterward" recognizing this as a hybrid, and adopt-
ing Sims's name, as required by the rule of priority. In the
last cited work, Gray also noted that Xuttall had proposed
the name P. involucrata, but had never adequately pub-
lished it.

The only other date of significance in the history of P.
amoena is 1903, when SmalP^ segregated from it a presum-
ably distinct species, under the name P. lighthipei. This was
reduced to varietal status by Brand^'^ (mis-spelled Ughtipei),
and reasons for accepting his view will be presented in the
discussion of variation in the species.

1 Flora Caroliniana 96. 1788.

2 Flora Boreali-Americana (1): 144. 1803.

3 Botanical Magazine 32: pi. 1308. 1810.

4 Flora America Sept. (1): 150. 1814.

r. Sketch Botany S. C. & Ga. (1) : 247. [1817.]
« In De Candolle's Prodromus 9: 305. 1845.

7 Manual of Botanv, ed. 2. 331. 1856.

8 Flora Southern States 339. 1860.

9 Manual of Botany, ed. 5. 372. 1868.

10 Index Sem. Hamburg 1828: 17; Linnaea_^o: 383. 18JU.

11 Proc. Amer. Acad. Art« Sci. 8: 251. 1870.

12 Flora Southeastern U. S. 978. 1903.

1-5 In Engler's Pflanzenreich IV. 250: 70. 1907.

INTENTIONAL SECOND EXPOSURE

50

PROCEEDINGS OF THE

Geography. — This species has a rather restricted range —
from Florida to eastern Mississippi, southern Kentucky, and
north central North Carolina. The records of the two recog-
nizable varieties, Phlox amoena walteri and P. a. lighthipei, are
indicated in figure 4 by solid and open circles, respectively.
The distribution of these suggests the species to have spread
from a dispersal center in the northwestern corner of Georgia
or adjacent Alabama. It is probably still expanding its range,
there being no indication that the present boundaries are
climatic.

0«llKhthip«i

FiQ. 4. Distribution of Phlox amoena.

r

Alabama : Abundant and wide-spread, being recorded from
33 counties: Autauga, Baldwin, Barbour, Bibb', Blount',
Calhoun', Cherokee', Chilton', Clarke, Cleburne', Colbert',
Cullman', De Kalb', Elmore', Escambia, Etowah', Fayette',
Franklin', Jefferson', Lawrence, Lee, Limestone', Marion',
Marshall', Mobile, Montgomery', Morgan', Perry', Randolph',
St. Clair', Shelby', Walker', and Winston'.

[Arkansas: Ascribed to this state by Mohr,^ but evidently
through misunderstanding.]

Florida: Occasional in the northern part, specimens —
chiefly of the southeastern variety — having been seen from
Clay, Duval, Gadsden, and St. Johns counties.

1 Plant Life of Ala. 686. 1901.

PHILADELPHIA BOTANICAL CLUB

51

Georgia : Distributed much as in Alabama, the county list
being: Bartow', Bibb', Bulloch, Camden, Catoosa', Charlton,
Chatham, Clarke, Coffee, Dade, De Kalb', Emanuel, Floyd',
Fulton, Glynn, Gwinnett, Habersham', Hall, Haral§on, Jas-
per, Madison', Peach, Richmond, Stephens', Sumter, Telfair',
Thomas, Walker', Wheeler', and Wilcox. The southeastern

variety is present locally.

Kentucky: Enters along the southern border, specimens
having been seen from Logan, McCreary', and Warren coun-
ties. Short's records from ''barrens" may represent others.

Mississippi : Rare and probably limited to the eastern edge
of the state. The records published by Lowe^ proved, on ex-
amination of the specimens in the State University lierbarium,
to represent mis-identified P. pilosa. The localities given on
two sheets seen in other herbaria can not be certainly placed,
but are thought to lie in Alcorn and Wayne counties.

[Missouri : Mohr extended the range of P. amoena to this
state, but must have mistaken some other species for it.]

North Carolina: Common in the Blue Ridge and inner
Piedmont, but apparently absent from the Coastal Plain. The
county list is : Buncombe', Burke, Caldwell, Catawba, Chero-
kee', Forsyth', Hayw^ood, Henderson', Iredell, Lincoln, Mc-
Dowell', Madison', and Polk'.

South Carolina : Occurs throughout the southwestern half,
there being 10 county records: Aiken', Anderson, Berkeley,
Cherokee', Greenville', Jasper, Lexington, Oconee', Pickens',
and Spartanburg. The type locality of the species was on
the Santee Canal in Berkeley.

Tennessee: Frequent except toward the west end, the
county list being: Blount, Bradley', Coffee', Cumberland,
Davidson, Franklin, Hamilton, Knox, Marion', Monroe, Mor-
gan, Polk', and Sumner.

[Virginia : In the second edition of his Manual of Botany,
Gray2 ascribed ''P. pilosa var. ? walteri'' to Virginia, and
this has been copied from one compilation to another ever
since. No substantiating evidence is, however, at hand.]

1 Plants of Miss. 234. 1921.

2 Manual of Botany, ed. 2. 331. 1856.

52

PROCEEDINGS OF THE

PHILADELPHIA BOTANICAL CLUB

53

[West Virginia : Phlox amoena was reported by Millspaugh^
in Fayette County ; no specimens seem to have been preserved,
however, and trips to that region have yielded only P. divari-
cata. Mr. L. W. Nuttall, who found the plant in question
advises me that its identity was never checked by comparison
with herbarium material, and that he regarded it as an in-
troduction from the west. It therefore seems likely to have
been some other species.]

Ecology. — The most frequent habitat of the present species
is a thin woods in rather sterile soil, although it sometimes
extends into deep woods where the soil is richer, and again
pushes out into swamp thickets or even boggy meadows. In
reaction preference it is to be classed as subacid, and only
rarely thrives in circumneutral soils. Its partially persistent
stems and rather small hairy leaves mark it as somewhat
xerophytic. From the successional standpoint it is inter-
mediate, being exceptional either as a pioneer plant or an
occupant of climax forests.

Its blooming period extends from early April through May
in the lowlands, and from May into June at higher elevations.
The flowers have little scent, evidently attracting insects by
their brilliant color. Fertile seeds do not seem to be pro-
duced very freely, however, and this species rarely becomes
abundant. Its failure to occupy a wider range is perhaps
in part connected with this lack of reproductive vigor.

Variation. — Phlox amoena is less variable than most other
species, and except toward the southeastern end of its range
is rather uniform in aspect. The leaves, which are normally
elliptic-oblong and obtusish, show a tendency to become
lanceolate and acuminate in many colonies, and in northeast-
ern Florida and southeastern Georgia this tendency is locally
dominant; the bracts being also narrowed and the stems
elongated, the plants then take on a striking resemblance to
those of P. pilosa or P. floridana. It was material of this kind
which, as noted under History, has been considered a distinct
species, P. lighthipei; but as many intermediates with typical
P. amoena exist, only varietal separation seems justified.

1 Living Flora of West Virginia 335. 1913.

vl t

The only other noteworthy variation occurs in the corolla-
color. This is normaUy phlox-purple (65 b), but a brilliant
hue approaching the true purple (65) of Ridgway is also
frequent. The bluer amparo purple (63 b) and the redder
mallow purple (67 b) are exceptional. In a few colonies
light-colored plants become prominent, their corollas being
pink, light violet (59 d), near-white, or even pure white.
Intense eye striae, usually rhodamine purple (67), are often
present, each lobe bearing near its base either a single broad
central stripe, two narrow lateral ones, or both in combina-
tion, yielding a striking stellate pattern.

VARIETIES AND FORMS OF PHLOX AMOENA

Leaves mostly elliptic-oblono and obtusish ; bracts broad.

Phlox amoena waiter! (Gray) Wherry, comb, nov P. pilosa Michau^,

not Linn^; P. amoena Sims; P. pilosa amoena Pursh; P. pilosa wal-

teri Gray P. walteri Chapman, in part; P. procumhens Gt&j, not

tlm^^-: P. involucrata Nuttall ex Gray. The widespread variety.i

Form:
Light-colored (corolla pink, Ught violet, white, etc.). Typified ^y
sDecimen from 9 miles southeast of Beechgrove, Coffee Co., Tenn.,
collected by E. T. W. May 7, 1929. Occasional in many colonies.
Leaves mostly lanceolate and acuminate; bracts narrow.
Phlox amoena lighthipei (Small) Brand [spelling corrected] ; P. wal-
ttri ChapmanTin part; P. lighthipei Small. In Eastern Fla. and Ga.

Cultivation.— Being adapted to growth only in sterile, acid
soils. Phlox amoena fails to thrive in ordinary gardens, and
is not in the trade. Many dealers offer under this name, how-
ever the hybrid more correctly known as P. procumhens
Lehmann; also listed as ^*P. verna'^ Hort. As noted under
History, Gray for a time confused this plant with P. amoena,
but soon corrected the error ; in doing so, however, he made
another, in that he suggested the parents of the hybrid to be
P suhulata and P. amoena. Actually the spatulate leaves,
strongly glandular pubescence, and long style show the second
parent to have been P. stolonifera instead. Horticulturists
who desire to label their plants correctly will do well to re-
member that ^' Phlox amoena'' Hort. is not the P. amoena of
botanists.

IP. rugelii Brand, in Engler^s Pfi^nzenreich IV. 250: 73 1907
= P. divaricata canadensis x P. amoena walteri. Ala.,/ Tenn.,/ and N. C.

i

IRREGULAR PAGINATION

Reprinted from Journal of The Wa.hin(;ton A. ademv of Sciences
Vol. 21, No. 4, IVbrimry l'.», iy31

BOTANY.— A neiv spiral-orchid from the sonthern states.' Edg.'^r T.
Wherry, University of Peiinsyl\-;inia.
While studying the soil-reaction relations of nati^^-e plants in the
sol, I have'repeatedly observed in boggy pinelands a spiral-ordj.d
Hadies-tresses) not corresponding to any species nicluded in bniall s
Flora' It is closely related to the Slender Spiral-orchid [Ibulnan
araciie (Bigel House)], and search for morphological differences be-
?ween them has not been particularly successful. They are, however
more or le"s distinct in flower color, sepal length, habitat, range, and
Zoming period, and show no evident intergradatiou. The southern
plant is accordingly here described as a new species.

Lateral sepals little exceeding the bond ^-^}^f^^^^^^l^:Ti:^
moderately acid grassy fields, b. t. to lex. and nortn^a ^^,^^.^^^^^ ^^^^.^^

Ibidium floridanum Wherry, sp. nov.

FIG. 1.

7 .^7/ dmili^ sed floribus vernalibus et labii medio intense flavo.

L=UTlv:lfroots severa^^^^^^^^^

autumn and withering the foU^mg s«^m n^^^^ ^' ^^^^.

elliptic, 1 to 4 cm long and 5 *» 20 3. «'««'o«^^™ {^ .i^glc ranked, often
4 to 7 remote scale-like leaves; mceme .5 to 10 cm long B ,j ^^

strongly spiralled; flowers usually oP«"'"^,".-^^'lXr creamy white with the
early as mid-December o[„=;f »^\«4%'^J^-Y' ^ '^bUngr^^^ (Raf .)

SSlSsS' K P-if ti"^ -ther markedly

beyond the beld in the lip; callosities stubby, 1 mm. long.

. Contribution from the Botanical Laboratory of the University of Pennsylvania.

Received November 26, 1930. .^^

t Flora of the Southeastern I nitcd States. 319. IMS.

49

50 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 21, NO. 4

Type in U. S. National Herbarium, no. 1,466,427, collected by Edgar T.
Wherry April 14, 1930, near Loretto, Duval County, Florida. Named from
the fact that it is widespread and abundant in this state, specimens having
been seen from the following counties: Broward, Duval, Flagler, Gadsden,
Hillsborough, Jackson, Lake, Lee, Marion, Orange, Pinellas, St. Johns, and
Walton.

Noteworthy specimens are as follows:

FLORIDA :

Tampa Bay, Burrows, 1834; the earhest known collection (New York
Botanical Garden)

Fig. 1. Ibidium floridanum
Type locality. Natural size.

Fort Myers. Standley, December 14, 1919; an unusually early blooming
date (United States National Herbarium)

Fort Lauderdale to Miami, Small & Carter, February, 1911; the southern-
most known station (N. Y. B. G.)

GEORGIA:

Milledgeville, Boykin, 1836 (N. Y. B. G.).
Wrightsboro, Chapman (N. Y. B. G.)
SOUTH CAROLINA: Andcrsou, Davis, April 9, 1919; the northernmost known

station (U. S. N. H.)
ALABAMA: Mobile, Mohr, May, 1868 (U. S. N. H.)
MISSISSIPPI: Koshtaw, Tracy, May 20, 1898. (N. Y. B. G.)
LOUISIANA: Alexandria, Hale, April. (N. Y. B. G.)

TEXAS: Houston, Hall, April 1, 1872; the westernmost known station (N. Y.
B. G.)

FEB. 19, 1931

WHERUY: A NEW SPIHAI.-OKCHin

51

The southernmost occurrences of /. gracile represented among specimens

soTt^carolixa: Aiken, Ravend September 1869 (US^^^ HO
ALABAMA: Auburn, Polked f^ ^^{«--vJf y ^-n, 1^00 (bb. N. H.)
ARKANSAS: Tcxarkaua, Heller, August, 1898 (N. Y B. U.j
The ranges of the two species thus barely overlap.