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VOLUME xin

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PHILADELPHIA
1937

^v

CONTENTS

Morris Arboretum

FORWARD

The present volume of the Contributions of the Department of
Botany and of the Morris Arboretum of the University of Pennsylvania
contains a collection of the papers published by the members of the
Botanical department in several periodicals between 1935 and 1936.

In this collection the subject of Botany is ^iven a broad definition
reflectinji the several lines of interest developed in the group.

Rodney H. True

Chairman, Department of Botany
Director, Morris Arboretum.

V
i

I

••>

i

I

I

.1

i

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4

4
i

ANDERSON, LEWIS E.: Mitochondria in the life cycles of certain
higher plants. American Journal of Botany, 23:490, (1936).

BOESHORE, IRWIN and WILLIAM D. GRAY: An Upper
Cretaceous wood : Torreya antiqua. American Journal of Botany,
23:524, (1936).

CARLSON, J. GORDON: Effects of several fixatives on staining
reactions in Zea mays, especially with reference to the Feulgen
reaction. Cytologia, 7:104, (1936).

CHILDS, THOMAS W.: See H. H. York.

EDWARDS, JOHN K. : Cytological studies of toxicity in meristem
cells of roots of 7.ea mays. I. The effects of the neutral salts.
American Journal of Botany, 23:483, (1936).

FOGG, JOHN M., JR.: Lophotucarpus spongiosus in Salem County,
New Jersey. Bartonia, 17:21, (1936).

GRAY, WILLIAM D.: Myxomycetes of Clark County, Indiana.
Proceedings of the Indiana Academy of Science, 45:69, (1936).

Notes on plasmodial behavior of Stemonitis

fusca Roth. Proceedings of the Indiana Academy of Science,
45:74, (1936).

See I. Boeshore.

See E. T. Wherry.

HUTCHINSON, W. G. : A method for staining rust mycelium in
woody tissues. Phytopathology, 26:293, (1936).

JUMP, JOHN AUSTIN : Wound responses of Ficus australis. Bul-
letin of the Torrey Botanical Club, 63:477, (1936).

McQUILKIN, WILLIAM EVERETT: Root development of pitch
pine, with some comparative observations on shortleaf pine. Journal
of Agricultural Research, 51:983, (1935).

McVAUGH, ROGERS: Studies in the taxonomy and distribution
of the Eastern North American species of Lobelia. Rhodora,
38:241,276,305,346, (1936).

MOTHER MARY CHRYSOSTOM: The influence
benzene derivatives on the roots of Lupinus a/bus.
Journal of Botany, 23:461, (1936).

SEIFRIZ, WILLIAM: Breakdown of fruit and vegetable tissue
due to an electric current. Plant Physiology, 11:195, (1936).

Vegetation Zones in the Caucasus. Geo-
graphical Review. 26:S9, (1Q36).

of several
American

July, 1936]

ANDERSON — MITOCHONDRIA

491

!^IITOCHONDRIA IN THE LIFE CYCLES OF CERTAIN

HIGHER PLANTS '
Lewis E. Anderson

TiiK BKiiAVioR of cytoplasm and cytoplasmic inclu-
sions in the life cycles of plants has received very little
attention, and up to the present the genetical data
which indicate that the cytoplasm may serve as the
bearer of certain heritable characters have been sup-
ported bv only mea-er cytological evidence Chloro-
phvU deficiencies which do not follow regular Men-
delian ratios, and which may accordingly be inferred
to l)e inherited in a manner other than through the
chromosomes, have been found in approximately forty
species of plants. It is therefore desirable that ad-
ditional knowledge be secured of the behavior of any
possible bearer of the factors concerned.

It is perhaps significant that such a large propor-
tion of these uon-Mendelian characters should be
concerned with plastids, and the question arises as to
the possibility of a direct transmission ot these cell
organs or of their i)rim()rdia from one generation to
the next. In certain algae fully developed chloro-
i)li<ts pass directly into the egg and thus into the
daughter plant; but in seed plants the chloroi)lasts
as such do not persist through all phases of the lite
cycle, although they may be represented in certain
stages by their primordia. The purpose of this m-
ves'tigation is to trace the behavior of these bodies
through the entire life cycle, especially during the
development of reproductive structures and the pro-
cesses leading to fertilization. It is hoped that such
a study may throw some light on the material basis
of maternal inheritance.

Cytologists, in general, agree that plastids arise
from some visible i)rimordium in the cytoplasm, the
transition of which has been figured many times.
There is much confusion, however, in the terms used
to designate these primordia, and most of the dis-
agreement as to what constitute i)lastid primordia is
])r()bablv due to this confusion of terms. All cyto-
l)lasmic' inclusions (plastid primonha and other bod-
ies) preserved by certain "basic" fixatives exhibit
similar staining qualities; and thus it is not possible
at present to distinguish between these primordia and
other similar appearing bodies— if, indeed, any such
distinction actually exists. Until we are able to iden-
tify definitelv the bodies which give rise to plastids
and those which do not, it seems best to use some
inclusive term tn embrace them all. For convenience,
therefore, the term " mitochondria " will be used for

1 Rocoivod for publication May 16, 1936.

Contribiition from tho Botanical Laboratory an<l tho
Morris Arboretum of the University of Pennsylvania.

This investigation was suggested by ami earned on
under the direction of Prof. Conway Zirkle. to whom the
writer wishes to express his sincere thanks and ai)i)ieoi-
Mtion Prof. Kdgar T. Wherry and Prof. John M. to«ig.
'ir ))nve rendered helj.ful assistance in the pivi.aration
of the manuscript. The work was done while ine hum .
held a Morris Arboretum fellowship.

all those small c%toplasmic inclusions preserved by
bichromates on the basic side of pH 4.2-5.2 and de-
stroyed by more acid fixatives and by mixtures ot
l,ichromates and acetates. This term thus includes
the primordia of plastitls, whereas the term plastid
itself will include only those bodies containing starch

or chlorophyll. _ ,

The literature bearing on the genetics of chloropli>ll
deficiencies has been extensively reviewed by de Haan
(11)33) and East (11)34). It is necessary here to re-
view, briefiy, onlv phases of the cytology of the work
which concerns this investigation. The cytology ot
variegated plants has been investigated by a tew
workers. Miles (IDlo) examined albino seedlings ot
Zca Maifs and found no evidence of plastids in pure
white plants, llamlolph (11)22) made rather exten-
sive studies on several types of chlorophyll deficien-
cies in the same species and reported that all types
contain minute proplastids. In contrast to Miles
findings, white seedlings were found to contain plas-
tids wdiich were retarded in their development. Ran-
dolph found onlv one type of plastid primordia in
meristematic tissue and no differences ^v^^e found
between green and colorless plastids. Zirkle (ID-J)
found that mitochondria in the root tips of Zea Mays
developed into plastids, all stages of development
beincr figure.l. In addition, he found two types of
mitodiondria, one of which develops into plastids,
the other, a smaller type, apparently undergoing no
change. In the stem growing point the primordia
are usually minute plastids, although occasionally
they are mitochondria. In variegated plants of Zea
exhibiting maternal inheritance, Zirkle distinguished
two kinds of plastids in lighter and darker stripes of
the leaf with no intermediate forms on the border-
line of the two regions. No evidence was presented,
however, that these two types of i)lastids are (le-
rived from two distinct classes of primordia. Tsinen
(11)23), working with Abutilon, maintained that
variegation is due to changes in the ])lastid primordia
or th'e plastids themselves. In Hosta, Yasui (1030)
could not distinguish between i)rim()rdia of green,
yellow, and white plastids, nor did she find inter-
mediate forms in mature cells.

The presence of plastids in the egg has been long
known (Schimper, 1885), and more recently Guil-
liermond (1924) described i)roplastids or mitochon-
dria in the egg of Lilhnn. Mitochondria have been
figured in the pollen grains of several ])lants, among
which are Cucurbita Pepo (Guilliermond, 1012) and
Hcllcborus (Wagner, 1927), as well as in the spores
of Equhetum (Lewitsky, 1925). In the pollen tul)e
of Liipiniis Iiitcus, Hiihland and Wetzel (1924) ob-
served structures resembling mitochondria which orig-
iiiMtPfl from plastids in the ])ollen grain. The transfer
of any of these cytoplasmic bodies from the pollen

'■ A

. I

47

tube to the egg, however, has not been demonstrated
cytologically.

Matehials and methods. — The plants investigated
were Antirrliinum majiis L., Pliiladelphus coronarius
L., and Hyacinthus orientalis L. Flowers of Hyacin-
thus were obtained from bulbs i)lanted in the green-
house and forced during the months of December
and January, at which time they flowered freely.
Root tips, stem growing points, anthers, and ovaries
in various stages of development were fixed at in-
tervals. Specimens of Antirrhinum were likewise
ol)taiiied from i)lants grown in the greenhouse from
seed, those of Philadelphiis from plants grov/ing at the
Morris Arboretum. In order to obtain the various
fertilization stages, the i)lants were hand-i)ollinated
and the rate of pollen-tube growth was determined
by smearing the styles on a slide at various intervals
after i)ollinatioii. The specimens were then stained
in crystal violet for 30 seconds and washed in water,
the excess stain being removed with filter jiaper.
The pollen tubes stain a brilliant violet while the
stylar tissue stains only faintly. Pollen tubes could
thus be traced to the microi)yles, at which time the
ovaries were fixed at two-minute intervals until fer-
tilization had occurred. Pollen tubes were fixed
directly in the style as well as on glucose agar where
they had been germinated. In the latter case small
blocks of agar upon which they were growing were
cut out, plunged into the fixative, imbedded in para-
Hm, and sectioned in the usual manner.

In order to preserve mitochondria, it was necessary
to use bichromate fixatives on the basic side of the
1)H range 4.2-5.2, a number of which were employed.
By far the most satisfactory fluid, however, was
Zirkle's (1934) modification of Erliki's potassium
bichromate-co])i)er suli)hate mixture in the following
proportions: Iv.Cr.O., 1.25 g.; (Xn^)..CrA' 1-^ 8-5
CuSO,, 1.00 g.;"ll Jo,' 200.00 g. Enough pyridine was
adtleil to this mixture to make ^/4 of 1 per cent.
After the resulting precii)itate had settled (which
required about 20 minutes), si)ecimens were placed
in the fluid for 48 hours. They were then washed in
water for an h(nir, dehydrated in l)utyl alcohol (Zirkle,
1930) and embedded in paraffin. Sections were gen-
erally cut 8 microns in thickness. The only stain
employed was Heidenhain's iron-alum haematoxylin.

All figures were drawn at the level of the table with
the aid of an Abbe camera lucida under a Si)encer
objective, 1.8 mm., N. A. 1.30, with either an 8X
or a 20 X ocular. They have been reduced ai>
]iroximately 2y2 times and show a magnification of
about 975 diameters unless otherwise designated.

All the specimens investigated were preserved in
bichromate fixatives on the })asic side of the pH rangc^
4.2-5.2. The cells accordingly show the typical
*' basic image," including both mitochondria and im-
mature plastids which would have been destroyed by
more acid fixatives. Unfortunately, the fixatives em-
]iloyed dissolve chromatin; and as a result the re.st-
ing nucleus is ])reserved as n solid mass of nuclear
Ixiiipu

(fig. 1), while in dividing cells the coagulated nuclear
lymi)h forms a spherical, dark-staining mass (fig. 7).
A number of artifacts are thus found in dividing
cells. No attemi)t was made to follow the behavior
of mitochonilria during nuclear division, as this would
have hatl little direct bearing upon the subject under
investigation.

MiTOClIONDHIA IX Al'ICAL MERISTEMS. — As mito-

chondria arise through the division of preexisting
mitochondria (Guillienuond, 1912; Friedrichs, 1922;
et al.) and as they are transmitted through the divid-
ing cells of the meristcm, it is i)ossible to trace the
course of their develoi)ment from the stem growing
l)oint of the young sporophyte to the micro- and
megaspores and finally to the male and female game-
tophyte. Thus the i)ollen tube and the embryo sac
derive their mitochondria from the same primordia,
as both of these organs are developed from the same
ai)ical meristem. The mitochondria in the male and
female gametes, therefore, can be traced to a com-

mon origin.

.^Ul 1 UlillMlil4

tiiU

iii. t< t it >

nucleolus

In the meristematic cells of the root the mitochon-
dria assume various forms, comprising both straight
and curved rods, and granules or chains of granules
(fig. 1). In Hyacinthus they are clustered about the
nucleus and around the perii)hery of the cell, al-
though some are also scattered throughout the cyto-
plasm. ^Mitochondria which fix as hollow spheres are
not uncommon and their transition into plastids can
be followed by tracing the intermediate forms from
the ai)ex of the root to the region of differentiation,
where they become mature plastids (fig. 2). This
transition is similar to that described by Zirkle (1929)
for root tips of Zea .l/r/j/.v.

Longitudinal sections of stem growing points of
Hyacinthus reveal both rod-shaped and granular
mitochondria as well as small plastids, the latter in-
creasing in number toward the region of elongation.
All stages in the transition of mitochondria into plas-
tids may be observed in the vicinity of the growing
tip. Even cells with mature i)lastids, however, always
contain some undifYercntiated mitochondria. Both
mitochondria and plastids are exceedingly abundant
in the procambial strand and vascular bundles. Em-
bryonic leaves also show the characteristic stages in
this transition, and oMer leaves contain in addition
mature plastids. The cells of the stem growing point
of Philadelphus contain many minute plastids which
are fi.xetl as small hollow spheres, together with a few
granular mitochondria and an occasional curved slen-
der rod (fig. 3). There is no sharp dividing line
between the jilastids and the granular mitochondria,
as there are many intermediate forms, particularly in
cells shortly removed from the growing point (fig. 4).

Floral branches of the difTerent plants studied show
inclusions characteristic of their respective shoot
meristems. Mitochondria and ])lastids in the meris-
tematic jirotuberances originating at the tip of the
young flower Inid are identical with tho.'^e in the
more mature floral iian- which arise from such ]iro-

1

+ 11 f ^nfo •■*onc"

O V» Cr paT t , uCii J

PAGINATION BRGINS

492

AMERICAN JOURNAL OF BOTANY

[Vol. 23,

all derived from the growing point of the same floral
branch, then receive the same types of cytoplasmic
inclusions. Thus it is evident that there is an equal
distribution of mitochondria to microspores and mega-
spores; and even if there should be more than one
kind of mitochondria, as Meyer (1920), Bowen (1927,
1929), Huhland and Wetzel (1924), and others have
reported, there is no evitlence of any segregation of
ditfeient types in the two kinds of spores.

MiT(H;HONI)RIA in microspores and MALE GAME-

Toi'nvji:.— The microsjjore mother cells of Philadel-
yhus iiudANtin'himim are extremely small, the nucleus
occupying the greater i)art of each cell (lig. 5). This
is particularly evident after meiosis is initiated, for
at that i)oint the nucleus increases in size perceptibly

(fig. 7). The cytoplasm forms a relatively thin layer
about the nucleus and the ])erii)hery of the cell, with
vacuoles forming the greater i)art of the cytoplasmic
volume. JMastid primordia are numerous and are
preserved by the iixatives as large spherical dark-
staining bodies, although there are in addition some
smaller sjjheres and rods (lig. 5). IVIitochondria in
the i)rocess of transformation into jilastids surround
some of the nuclei, while minute plastitls already
transformeil are not uncommon. The plastids fix as
hollow s])heres with a minute starch grain in the
center. Transitional stages become more abundant as
meiosis i)roceeds.

In Hi/ncinthifs the mother cells are much larger
than in the two above-mentioned genera, although
the nuclei are also corresj)ondingly larger and occupy
most of the cell (fig. 6). The cytoplasm is strikingly
vacuolate and is densely filled with mitochondria
which stain intensely, they fix generally as solid
sjjheres and short thick rods, although there are
occasional slender threads. In mother cells there are
ai)parently no transitional stages between mitochon-
dria and plastids. The cytoi)lasm becomes increas-
ingly vacuolate as division proceeds (fig. 7). IVIito-
chondria are transmitted through the reductional
divisions directly to the microsi)ores, and each
microsjiore apparently receives an approximately
ecjual share of the cytoplasmic inclusions. There is a
very striking increase in tlie number of plastid pri-
mordia in the microspore over those appearing in the
sporocyte (cf. fig. 7, 18). Whether this increase is
due to an actual nuiltiplication of these bodies or
whether they represent nutochondria formerly ]we-
served below the limit of visibility is not certain.
The latter theory, however, seems less prol)able, since
at no stage were mitochondria observed to grade off
in size to the border line of visibility.

Mitochondria are exceedingly abundant in the
mature microspore or pollen grain just before it is
shed and are jireserved as short, thick, solid rods
and spheres which have increased somewhat in size
(fig. IS). The cytoplasm is very vacuolate, leading
to the clumping of the mitochondria into definite
aggregations or bundles, n normal condition in rela-
tively scanty cytoplasm (fig. 21). In the mature
pollen grains at the tinip nf chpddino' both the larger

solid rod-shaped mitochondria and the larger' spheres
begin to api)ear hollow (fig. 21). At this time the
transition into plastids occurs, and later, at the time
of i)ollination, young plastids are numerous. The
latter can be demonstrated in thin sections in which
most of the mitochondria have been completely de-
stained, for in such sections the plastids are very
consi)icuous (fig. 25). They are hollow spheres or
rods each with a minute starch grain in the center.

Mitochondria in the microsjiores of Philadelphus
and Antlrrhinmn are not nearly so abundant as in
those of Ilyaclnthus, and they fix as extremely small
spheres and thin straight and curved rods. Most of
the inclusions are i)lasti(ls. Possibly this difference
in the number of the cytoplasmic inclusions in Phila-
(felphiis and Antirrhinum on the one hand and
Hyacinthus on the other is due to variations in their
fixing i)roi)erties. It is also possible that substances
leached out of the cells of the specimens fixed may
have dissolved some mitochondria.

Pollen-tube growth is readily followed in Hya-
cinthus, owing to the shortness of the style and to
the relatively large amounts of cytoplasm in the
pollen tubes. Living i)ollen tubes were grown directly
on glucose-agar, and both active cytoi)lasmic stream-
ing and the passage of the nuclei down the tube could
l)e observed. The streaming cytojilasm is filled with
granules and with distinct rod-shaped bodies. The
cytoplasm of living pollen grains exhibits these same
inclusions, and as the germination tube makes its
appearance they can be seen flowing directly from
the pollen grain into the elongating tube (fig. 19).
The nature of these inclusions visible in the living
material is not altogether certain, but they strongly
suggest mitochondria and plastids. Both mature
pollen grains and i)ollen tubes react i)ositively to the
iodine test for starch, which is evidence of the exist-
ence of plastids.

In the i)ollen tube the mitochondria are fixed either
as small spherical bodies or as rods, which may l)e
either short and thick or thread-like (fig. 15, 19).
Both the transitional forms of mitochondria and the
plastids observed in i)olIen grains flow into the pollen
tube with the streaming cytojilasm (fig. 19, 20). The
cytojilasmic inclusions are more concentrated at the
tip of the tube where they siuTound the tube nucleus.
Each male nucleus is likewise accompanied bv a dense
mass of mitochondria. In Hyacinthus the male nuclei
are not enclosed in individual cells but lie free in the
jwllen tube. As they migrate down the tube, they
are surrounded by a characteristic mass of inclusions
(fig. 15) which accomi)any them until thev are dis-
charged into the embryo sac. In the pollen tubes
of both Antirrhinum and Philadelphus mitochondria
are abimdant, l)ut they are not easily seen because
of the small size of the tubes and the difTiculty of
fixation. Plastids were not observed in either of these
genera. The mitochondria usually occur as tiny
si)heres and very slender rods, either straight or
curved, forms which were characteristic of the micro-
spore iiiulijui cdl* and growing points. The male

!

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Fig. 1-12. — Fig. 1. Cells from root tip of //war//j///j/s— Ficr 9 n^iu f^ *i • /• ,

.howin. „,il„cho„,l,iu, „la..id.,. and Ira '.ilion'fl « ts e.,^onthe two fL ■,' ^T."' "^T"*'"" '" "«!"'""■'"'
of I'hIlm/rlphus.-FiK. 4. Oil from olonnlinL. >•,.. 71 If IT . r ^7, f,^; *-' "'°'" ""' '"™' K™vin!-' Point
PhihuUML-Fi^. 6 l>olk!„ , [o or COM // I^ r,;°f/^T "-" "< '•'"''"'' ll'hm.-Fii,. 5. I'ollon n.o.hor cell, of

of I,,,.rin,lu,l with n.i.o;.ho„.lr ^Z^^o^i' ^.IJ^'::":^'T^,:' II !..„„„.. F,.. 10. M..a.pore

494

AMERICAN JOURNAL OF BOTANY

[Vol. 23,

nuclei are spherical and extremely small, in sharp
contrast to those of Hyacinthus, but they are simi-
larly surromided by mitochondria which are present
as the i)ollen tube enters the embrj'o sac.

MlTOCllONURIA IN MEGAS?0RES AND FEMALE GAME-

TOPiiYTE. — The cells of the meristematic protuber-
ances which give rise to the ovules are in general
similar to the i)remeiotic cells of the anthers, which
have already been described. In Ilijacinthus the
nuclei are surrountled by both spherical and rod-
>hai)ed mitochondria and the cytoi)lasm contains sev-
eral ])rominent vacuoles. There are no visible plas-
tids or transitional forms, and all the cells of the
l^rimordia of ovules are alike until the megaspore
mother cells are differentiated. At this time the
nucellus has become prominent and the inner and
outer integuments have formed. These structures
contain an abundance of mitochondria, which collect
around the nucleus and in the cytoplasmic strands
between the vacuoles and just within the cell wall.
The megasi)ore mother cell develoi)s from one of these
cells (fig. S). It enlarges rapidly and becomes more
vacuolate as meiosis is initiated. The cyto])lasmic
inclusions of the mother cells of Antirrhinum are
similar to those of Hi/ncinthus except for their smaller
size and predominantly granular form.

Meiosis occurs rapidly in both genera. The nuclear
details are diilicult to follow, however, when the
material is preserved in basic fixatives, although the
e(iual distribution of mitochondria to the four mega-
spores can easily be seen. In both Ilyncinthus and
Antirrhinum the megasi)ore nearest the micropylar
end of the ovule develops into the embryo sac, the
others disintegrating in the normal fashion. All four
megaspores jwssess mitochondria which maintain their
characteristic groui)ing around the nuclei. The func-
tioning megaspore enlarges and gradually encroaches
upon the other megaspores, which finally disintegrate.
These disintegrating cells retain their mitochondria
for a time, but eventually their staining properties
become such as to obscure any inclusions that might
be contained. Continued enlargement of the inner
megasjiore results in the others being i)ushed together
until finally they are either obliterated or reduced to
a cap at the end of the functioning megaspore. As
the latter enlarges, its cytojilasm becomes increasingly
vacuolate until it is limited to a thin layer about the
periphery of the cell with a few strands traversing
the vacuoles to connect with the centrally located
nucleus (fig. 0). Mitochondria are limited almost
entirely to the cytoi)lasm surrounding the nucleus,
where they form a dense mass (fig. 10). Solid, spher-
ical mitochondria predominate, but there are also
numerous short or elongate rods and threads. Un-
developed plastids are present usually as hollow
spheres and rod-shaped mitochondria which have be-
come vacuolate at one or both ends (fig. 11).

The development of the megaspore in Antirrhinum
differs in certain minor details from that of Hi/acln-
Liiuis. Aiiu iiuLjm.-5 ui lilt' eiiialgcii lllf^^^.>]>ult• in iiKe-

wise surrounded by a dense mass of mitochondria

4yu

which differ from those in Hijacinthus o»ly in size.
In both genera the grouping of mitochondria around
the nucleus is pronounced, and only scattered inclu-
sions occur throughout the rest of the cytoplasm (fig.
10). Occasional transitional forms are i)resent and
not infrequently there are small plastids.

The megaspore enlarges perceptil)ly before the first
division of the nucleus, after which the daughter
nuclei migrate, one to each end of the embryo sac,
and the cytoj^lasm becomes progressiveh' more vacuo-
late (fig. 12). Each daughter nucleus is also sur-
rounded by the characteristic grouping of cytoplasmic
inclusions similar to the i)arent nucleus and each
receives aiii)roximately an equal number. The four
nuclei jiroduced by the next division are all sur-
rounded by the characteristic clumi)s of mitochondria
(fig. 16). A thirtl division produces eight nuclei,
four at each end of the embryo sac. Each of the
nuclei is surrounded by a mass of cytoplasmic inclu-
sions which consist of mitochondria, transitional
forms, and plastitls. During the migration of the
polar nuclei to the center of the embryo sac each
nucleus retains its individual sjihere of surrounding
inclusicms. In Antirrhinum, the ])olar nuclei fuse as
soon as they come in contact with each other and a
very prominent grouping of mitochondria is formed
in the center of the embryo sac (fig. 22). ^litochon-
dria are jireserved generally as solid spheres which
are comjiaratively small in Antirrhinum but very
abundant. Rods and threads are also present but
they are not so numerous as the granules. Tran-
sitional forms as well as plastids occur not infre-
quently. In H unci nth us the two polar nuclei usually
do not fuse imtil after the pollen tube has grown
ajijiroximately half way down the style. They are
both surrounded l)y plastid primordia, however, which
are larger and more consi)icuous than those in
AntirrJiinum.

The mature embryo sac in both Hijcicinthjts and
Antirrhinum contains eight nuclei l)efore the two
polar nuclei fuse. The antijHKlal nuclei of Antirrhi-
num are small and are occasionally included in three
somewhat inconspicuous cells, each definitely de-
limited by a membrane (fig. 28). Usually, however,
the nuclei lie free in the embryo sac. All three nuclei
are surroimded by mitociiondria. In Hijacinthus, on
the other hand, the antipodals are exceptionally large
(fig. 14), with their cytojilasm very vacuolate. They
are difficult to fix satisfactorily and are often so
shrunken and distorted as to be scarcely recognizable.
When projierly fixed, however, the nucleus of each
cell is surrounded by numerous mitochondria and
some i)lastids. The polar nuclei, likewise surrounded
by these inclusions, usually lie adjacent to the anti-
podals and are often in contact with them. The
greater part of the volume of the embryo sac in both
Antirrhinum and Hijacinthus is occui)ied by large
vacuoles (fig. 22), the cytoplasm being limited to a
thin layer around the periphery and to the egg
aj)paraius, aniipodais, and endosperm nuclei. The
mitochondria and plastids, of course, are limited to

1

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Go „,„ml,n« „o on pa,„ of //„„.,„//,„. sh,„vi„s n,i.„ohon,l,ia an,l plas.kl. flowing i L tho pollon „bo - Fk 2o'
21. Matiuo i,olloii gram o( Ilyaanlhm. Mitochondria appoar clun.iied owing to var„oles "'

495

496

AMKKICAX JOURNAL OF BOTANY

[Vol. 23,

July, 1936]

ANDKUSON — MITOCHONDRIA

497

I

the cj-toplasm and are most ahuiidant in the regions
adjacent to the nuclei, although scattered inclusions
are present throughout the cytoj^lasm of the entire
embryo sac. The cells of the nucellus contain few
vacuoles and stain heavily. These cells contain more
mitochondria than any other cells of the ovule; in
fact, in Autirrhinutn, the nucellus is jiractically filled
with small granular mitochontlria (lig. 22). The cells
of the integuments also contain mitochondria, but
they are less abundant than in the nucellus.

In tixed material of lK)th Antirrhinum and Hya-
cinthns the cells of the egg apparatus as well as the
synergids are bounded by distinct membranes (fig.
13, 17, and 22), and consecjuently their cytoplasm is
definitely separated from the rest of the female
gametophyte. The two s\nergids lie one on each
side of the egg, which extends somewhat more deejily
into the embryo sac. The nucleus of each synergid
is located at the center of the cell and is comi)letely
surrounded l)y mitochondrii of the same nature as
those in the microspore. The cytoplasm is denser
in the region toward the mi('ro]nle and becomes more
vacuolate at the other end, usually with a single
hirge vacuole filling most of the l^roader extremity of
the egg (fig. 13, 22). In Hijacinthns the synergids
are somewhat larger than the egg, with mitochondria
very al)un(lant in the nudeir region but less numer-
ous toward the microi^ylar end (fig. 13). The syner-
gids are smaller in Antirrhinum and in closer contact
with the egg cell. The nuclei are very small and
inconsi)icuous, and both jilastid primordia, which are
l)reserved as rods and solid or hollow si)heres, and
small plastids are i)resent (tig. 22).

The nucleus of the egg of HjiacintJius is large and
conspicuous and lies in the end away from the micro-
pyle (fig. 14). The surrounding cytoplasm is rather
dense, becoming more vacuolate toward the micro-
])ylar end (fig. 17 1. Mitociiondria are likewise most
numerous around the nucleus, forming the character-
istic den.'je mass already described in the pollen tube
and other structures. They are ]ireserved as solid
spheres, thick rods, or threads, and any of these
forms may serve as the primordia of plastids. These
are not uncommon in the eggs of both Antirrhinum
and Hyacinth us and develoj) in the same general
manner in both species (fig. 22). The mitochondria
in the egg of Antirrhinum are almost as numerous
as they are in Ilyacinthus, but they are much smaller
and aiipear mostly as spheres and short rods. A
similar transition into ])lastids occurs in this species,
although the hollow sjiheres are much more difficult
to distinguish and the resulting plastids are corre-
spondingly smaller (fig. 27).

Mitochondria in fkrtilization. — Actual fertiliza-
tion was observed only in Antirrhinum, which, be-
cause of the large number of ovules in a single ovary,
is relatively easy to find. As the pollen tul)e enters
the embryo sac, the tube nucleus is surrounded by
mitochondria. The tube ])a-ses between the synergids
and lies along ihe .-idt' of the egg where, in ^ume
cases, it may remain for some time before it ruptures.

The m;de nuclei pass down the tube, errch nucleus
being surrounded by mitochondria. The pollen tube
after entering the emVjryo sac becomes much swollen
and moves toward the region of the egg between
the female nucleus and the micropyle (fig. 23). A
protuberance ajipears on the unruptured pollen tul)e
and presses against the egg. The mode of rupture of
the tube was not determined, but it apparently occurs
rapidly and with considerable force as described by
several previous investigators. No perforation such
as that reported by Schafi'ner (1897) and others was
observetl. The male nucleus which is to fuse with
the iiolar nuclei is api)arently discharged first, and
migrates through the cytoiilasm of the embryo sac
(fig. 23), finally fusing with the polar nuclei to form
the endosperm nucleus (fig. 24). In the course of
this migration through the cytoplasm it is surrounded
by mitochondria (fig. 23), although it was not pos-
sible to determine with certainty whether these sur-
rounding inclusions were discharged along with it
from the jwUen tube or whether they were originally
jiresent in the cytoplasm of the embryo sac. Prob-
al)ly, however, some cytoi)lasm escapes from the
jiolien tul)e at the time of rupture, for this breaking
is ai)parently due to the high osmotic pressure in the
tube, and in view of the large number of mitochon-
dria in the cytoplasm it is hardly conceivable that at
least a portion of them can fail to pass into the
embryo sac along with this male nucleus.

The male nucleus which is to unite with the egg
nucleus enters the egg immediately after it is dis-
charged from the pollen tube. In a preparation such
as that shown in figure 23, there seems to be an
actual connection between the pollen tube and the
egg. If such is the case, this opening would afford
ample opiiortunity for the jiassage of jilastid pri-
mordia from the tube to the egg. Furthermore, since
mitochondria are so numerous around the male
nucleus during its migration down the pollen tube,
it is difficult to see how, under such conditions, the
nucleus could enter the egg without at least part of
these inclusions accomi^anying it. After the male
nucleus has entered the egg, it is surrounded by
mitochondria which are similar to those accompany-
ing it in the i)ollen tube (fig. 23). To be sure, the
actual ])assage of mitochondria from the pollen tube
into the egg was not observed. Accordingly, it can-
not be stated definitely that these mitochondria,
which surround the male nucleus in the egg cell, were
discharged along with it when the pollen tube rup-
tured, for no mitochondria or other inclusions present
in the egg after fertilization could be definitely iden-
tified as having lieen passed into it from the tube
along with the male nucleus. However, since the
male nucleus is surrounded by mitochondria just be-
fore it is discharged and the cytoplasm of the entire
pollen tube is filled with inclusions, it would seem
impossil)le for the pollen tube to discharge its nuclei
without some of these inclusions escaping into the

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by no special cell membrane or wall (fig. 22), the

resulting endosperm will almost certainly contain
some mitochondria which were released from the
pollen tiilie. Since the opening through which the
male nucleus passes from the tube into the egg is
sufficiently large to i)ermit the entrance of inclusions
along with it (fig. 23), there seems no reason to doul)t
that they do enter. As the male nucleus migrates
through the cytojilasm of the egg to the egg nucleus,
it is surrounded by both mitochondria and plastids.
After fusion, the fertilized egg nucleus is likewise
surrounded by many small granular mitochondria
and some scatteretl hollow si)lieres and small i)lastids
(fig. 24). The ruiHured ])ollen tube still contains a
few mitochondria, although they are not nearly so
numerous as they were before the male nuclei were
discharged.

Mitochondria in the embryo. — After fertilization,
both the dii)loid nucleus of the egg and the triploid
endosjierm nucleus are surrounded by mitochondria,
I)lastids, and transitional forms between the two (fig.
24). The endosjierm nucleus divides immediately
and a wall is laid down dividing the two daughter
cells, both of which contain mitochondria and plas-
tids. Further tlevelopment of the endos])erm is rai)id,
walls being formed after each division and each
nucleus in turn being surrounded by mitochondria.
The endosjierm cells are large and highly vacuolate,
the cytojilasm being limited to the nuclear region
and to a thin layer just within the cell wall, in which
there are likewise scattered inclusions (fig. 2()). The
endos])erm contains mature plastids, as can be seen
in figure 2(]. The fertilized egg does not divide until
the develojiment of the endosjierm is well advancejl,
but remains a single, large, vacuolate cell, with mito-
chondria scattereil through the cytoplasm and char-
acteristically clustered about the nucleus (fig. 2()).
After the en(losi)erm forms a many-celled structure,
the nucleus of the fertilized egg divides (fig. 27),
forming a two-nucleate structure at the end of the
endosperm, after which a wall is laid down. Subse-
f|uent divisions result in the formation of an elongated
row of cells constituting the proembryo. all of which
])ossess mitochondria derived from the fertilized egg.
Further difTerentiation results in the develoiiment of
the susjiensor an<l of the embryo jirojier, wliicli con-
tain cytoplasmic inclusions in no way difTerent from
those of the fertilized egg. The terminal cells of the
embryo continue their division, forming the cotyledons
and growing jioints, into which are transmitted the
identical inclusions which were present in the fertil-
ized egg.

Discus.s^ioN. — Geneticists, in general, have agreed
that the inheritance of certain chlorophyll abnormal-
ities which do not conform with regular Mendelian
ratios are conditioned by the cytoplasm or its inclu-
sions. Hence, certain chloroj^hyll deficiencies which
are transmitted only through the female jilant have
been explained by the assumi)tion that the eng does
not receive any male cytoi)lasm during fertilization.
This e\j)lanation was first suggested by Cnrrens in
1909, uht'ii ije a.-'suuied liiaf tiie transmission of varie-

gation in Mirabilis, the classical example of maternal
inheritance, was throuuh the cytoplasm, since the
variegation was not carried by the male nucleus.
He assumed further that this character is transmitted
only through the egg cell, for he believed that no
cytojilasm enters the egg with the si)erm nucleus.
The develoi)nient of the '" defective " jilastids which
cause variegation was believed by Correns (192S) to
be conditioned by diseased cytoplasm, a theory which
has the recent siipi)ort of East (1934) in a critical
review of the subject. Other investigators ((Jregorv,
1915; Winge, 1919; Chittenden, 1925; et al.) postu-
lated that the jjlastids themselves or their i)rimordia
are the bearers of the deficiency. Baur (1909), on
the other hand, explained the non-Mendelian variega-
tion in Pelorqonium zonule, which is inherited through
both ])arents, by assuming that i)lasti(ls were brought
into the egg cell along with the male nucleus, since
this character is transmitted by both i)arents. There
are other genetical data indicating that male cyto-
plasm enters the egg in fertilization (see East, 1934).

The investigations of the writer do not supiiort
the view that no male cytojilasm passes into the egg
during fertilization. The jiersistence of i)lasti(Ls or
their iirimordia during all i)hases of the life cycle
can easily be demonstrated. They exist in the apical
meristems and in all the organs derived from the
stem growing i)oints. They assume identical forms
in both the male and female gametophytes, where
they are exceedingly numerous. They are especially
concentrated in the cytoijlasm of the pollen tul)e,
where they surround both male nuclei, and, as the
tube is ruptured, they are exjielled into the cytoi)lasm
of the embryo sac. The resulting endosjierm, there-
fore, contains ])lastids contributed l)y both i)arents.
These relations have not been recognized heretofore,
because nearly all ])revious investigators of this prob-
lem have been limited to fixed cytological prejiara-
tions where the cytojilasm has l)een jireserved in such
a manner that all its inclusions were dissolved.

The egg itself, on the other hand, is delimited by a
distinct membrane. Whether or not the fertilized egs
contains jilastids contributed by the male plant will
depend upon the ability of this meml)rane to divest
the male nucleus of its surrounding cytoi^lasmic in-
clusions. The exact mode of entrance of the male
nucleus could not be determined. Heimans (1928)
reported that the male gamete enters the egg cell
through a small openini: in the membrane, which may
well be, in view of the observations made by the
writer. After the male nucleus has entered the egg.
the oi)ening through which it has passed remains for
a time in the membrane (fig. 23). This pore is large
enough to iiermit mitodiondria to enter along with
the male nucleus, and a- shown in figure 23, they are
jiresent both inside the <"zg and in the adjacent pollen
tube. In view of these ob.servations it is diflficult to
conceive of the male muleus entering the egg without
a certain numlier of mitochondria as well as some
cytojilastu acc()mi)anyin- i^. As the mitochondria of
tiie pollen tul)e do not dilter in present staining re-

I I

July, 1936]

ANDERSON — MITOCHONDRIA

499

If

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Fiji. 22-27 .—Fifi. 22. MicT()i)ylar end of maturo oml>ryo sac of Anlirrhunim, with surrounding nucoUus; (a) synorjiul,
(h) ffiir. (r) fused i)olar nurloi.— Fig. 23. Kinhryo sac of Antinhhnnn just aftor tho i)()ll('n lul)e has ruptured. ()no
nialo nucleus (/>) has ontcrrd tho ogg leaving an opening in tho niomhrano. Tho other male nucleus is migrating
throiigh the cytoi>lasm of the embryo sac to the fused jiolar nuclei (c). Other .<<tructures .shown are (a) pollen
tul)e."(r) synergid. {d) egg nucleus, (/) antipodals.— Fig. 24. Enilnyo sac of Anlirrhhuim immediately after fertiliza-
(i()„ _Fijr." 25. Mature itollen grain of 11 ua<-inlhu^ with mitochondria destained t(} show i)laslids and transitional
f-j,.nis.— Fig. 26. Kmhryo sue after f(>rtifization. showing endosi)erm (6), and zygote (a) which has not yet divided.
X 412.— Fig. 27. Young emlnyo showing first di\ ision of fertilized egg nucleus. The endosperm is not shown. X 1050.

i

49S

4'JU

actions and in form from tho.se of the egg, they can-
not be distinguished from the latter once they have
entered the egg.

If the cytoplasm with some of its inchision.s passes
into the egg along with the male nucleus, Correns'
original exi)lanation of maternal inheritance will have
to be abandoned, and the caii.se of the failure of the
pollen to transmit these plastid deficiencies will have
to lie sought elsewhere. In some forms (Zea, Oryza,
Hosta, etc.) only two tyjies of plants reach maturity,
those which are variegated and tho.se normally green.
The lighter jilants die. The variegated plants iiro-
duce a certain amount of normal jiollen— i.e., ])ollen
without any defective plastitls. When the flower is
pollinated with a mixture of normal and defective
pollen, selective fertilization could exjilain the failure
of the male parent to tran.smit the defect.

It is evident that only a few mitochondria enter
the egg with the male nucleus. They are scarce,
therefore, in comparison with the large number already
pre.sent in the egg cell. In the subsequent divisions
of the fertilized egg which form the ])roembryo, the.<e
few defective ]irimordia would have little chance of
being distril)uted to just tho.se cells which form the
embryo proper, and even if they reached the embryo
they might not be in those ceils which give rise to
the shoot meri.stem. Thus the defective plastid pri-
mordia might well iiarticijiate in the act of fertiliza-
tion and yet have little influence ujion the progeny.

An alternate exjilanation can be found in the
assumption that the cytojilasmic inclusions of the
pollen tube do not function in the egg cytojilasm.
Since any plastid ])rimordia which may be contributed
by the male cytoi)lM.><m during fertilization are indis-
tingui.shable from tho.-e alreiuly jireseut in the ("zn,
the plastid ]irimordia from the i)ollen tube cannot be
traced as such in the sub.se(|uent develoi)ment of the
zygote. Consequently, there is no way of determin-
ing whether mitochondria derived from the male
parent develoj) normally after they come in contact
with the cyto))la.sm of the egg or whether they are
incapable of develojiing into jilastids after they enter
the egg. Large numbers of mitochondria are trans-
formed into ])lastids in the fertilized egg and during
its inunediate divisions ffinr. 27), but none of these
can be recognized as havim: been derived from the
jiollen tube. Any mitochondria from the male cyto-
pla.sm which enter the egg may disintec^rate, and thus
the primordia of defective iilastids from the male
plant would not ])e evident in the offsjirinir. There
is evidence of a jiarallel occurrence in certain of the
green algae. In Zj/cinoHn, for examjile, the chloro-
l)lasts contributed by the '' male " gamete in con-
jugation degenerate in the zygosjmre, only the chloro-
plasts of the female gamete remaining functional
(Kurssanow, 101 1 ) . A somewhat similar development
has l)een reported for tspirogym (TWindle, 1911).
AMiether the same sort of thing occurs in the higher
])lants cfnmot lie determined until some method of
distinguishing between the cvtoplasmic inclusions of
the pollen tube and of the egg has been devised.

There is, of course, the possibility that the develop-
ment of normal jilastids is inhibited by either diseased
or defective cytoplasm, and that the factors which
determine aberrant chlorojihyll are not transmitted in
any visible structure. Such an a.ssumjition does not
markedly affect the nature of the problem, however,
since the distribution of the cytoplasm and of the
cytopla.smic inclusions is essentially the same.

SUMMARY

The plants investigated were Antirrhinum mnjus
L., Philadelphus coronarius L., and Hyacinthus orien-
tal is L.

It was ])o.ssible to demonstrate mitochondria and
plastids in structures in which they had not heretofore
been rejiorted by using fixatives on the ba.sic side of
pll 4.2-5.2. The more acid fixatives employetl by
most i)revious investigators dissolve them.

The term '* mitochondria " is u.sed to include i)la.s-
tid i^rimordia, since all the.se groujis of inclusions
exhibit similar fixing and staining ])roperties and are
indistingui.shable from one another with jiresent
methods of investigation.

All living i)lant cells studied were found to contain
the i^rimordia of plastids, the develojiment of which
can be traced in the stem growing jioint and in the
leaf cells. Subsequent organs developed from growing
points contain these iiriniordia.

Plastid ])rim{)rdia are i)re.sent in microsjwre mother
cells from which they are distributeil approximately
equally to the microsjjores, where they are particu-
larly abundant. Transition of the.se primordia into
plastids could be followed in both pollen grains and
l)ollen tubes. The.se cytojila.smic inclusions are carried
from the microspores into the imllen tube by the
streaming cytoplasm. 1'hey are particularly numerous
about the male nuclei and per.si.st at least until the
pollen tube ruptures in the embryo snc.

Megaspore mother cells contain mitochondria iden-
tical in appearance with those of the microspore
mother cells, and they are likewi.se di.stributed equally
to the four megaspores. The functional megasjiore
posse.s.ses mitochondria which are strikingly abundant
a-ound its nucleus. The nucleus of each daughter
cell is in turn surrounded by mitochondria. The egg
cell contains both plastids and mitochondria, as do
all the other cells of the embryo sac. The polar
miclei are surrounded b> the.se inclusions before and
after they fu.se.

Fertilization was observed in Antirrhinum. The
pollen tube does not rup'ure until some time after it
has entered the embryo sic; instead, it remains for
a time in contact witli the egg. A i)rotrusion is
formed toward the egg. ind at that jmint the tube
luptures.

The male nucleus whii fuses with the polar nuclei
is surrounded by mito londria, and the resulting
endosjierm consequentl> (-(utains cytoj^lasmic inclu-
sions from the male plani. The second male nucleus.
.«"nrronndpd In- mi^^w.Jw,. .i. rm+o- . +1,^ — , . i r

■ ■ , -..., 1. iin_ «-^^ a lilt 1 U.-'f .■^

With the egg nucleus. The membrane surrounding

500

AMERICAN JOURNAL OF BOTANY

[Vol. 23,

Roprintod from the Amkhkan Joukxal of Botany, Vol. 23, No. 8, 524-528, October, 1936

printed in U. S. A.

li

! I

the ejiff, nftor it h.-is ])OPn ])niK'ture(l by the male
imclei'is, contains an oponins larso onoush for mito-
chondria to pass in from the jwllen tube.

Plastids and their i)rinu)r(lia are present in all the
cells of the deve!oi)ins eml)ryo where the transitional
stajjes between mitochondria and ])lasti(ls are most
alnmdant. The^e inclusions are fomid in the cyto-
])lasm of all cells formed before the growinji points
api)ear, as well as in the srowins points themselves.

It is shown that if mitochondria from the male
cytoplasm enter the e«rp: fell, any defective cytoi)lasm

or cytoplasmic inclusions present in the pollen tube
wouid be transmitted to the egg.

The exi)lanations of maternal inherit;uice which
assume that no cytoplasm is transferred from the
male plant to the female plant in fertilization are
thus inade(iuate. Several alternate possibilities are
suggested.

DfKE rxIVKKSITY.

Durham, Nokth (^^HOLINA

LITERATURE CITED

Baur, E. 1909. Das Wesen und die lMblichk(>i'sver-
hidtnisse (1(M- " Vari(>t:ilt's alboniurjjinatae hort '' von
Pdiin/oiiiiint ZitniiU . Z.'itschr. Ind. Abst. Vererb. 1:
330-351.

BowKN, K. H. 1927. A v.n^liniinarv r(M>ort on the
structural elements of ihe cytoplasm in plant cells,
liiol. Bull. 53: 179-196.

1929. Studies on the structure of plant proto-

l>lasm. II. The ])lastidome and pseudochondriomr
Zeit.schr. Zellf. Mikro. Anat. 9: 1-65.
CiiiTTKNDKX, R. J. 1925. Studies in variejiation. II.
Illlflnniina and P< htif/oniuni : with notes (m certam
chinuM-ieal arraujiemcaiUs which involve .^^tenlity.
Jour. (J{>netics 16: 43-61.
C'ouRKNs. (\ 1909. Vererbunnsversuche mit blass ({relb)
* fininen un<l bmilblattrif:en-Sip])en bei MImhilis
hihipa Vrlirn pUiiliirm, und LmKirla (unnia.
ZrWschv. Ind. Abst. Vererb. 1: 291-329. ^
. 192S. Vhvv niehlmendelnde Ver(>rbun;i. Z.'it-
schr. Ind. Abst. Vererb. (Suppl.) 1: 131-138.
K.^sT. K. M. 1934. The nucleus-plasma problem. Am<T.

Nat. 63: 289-303; 402-139.
FijiKDRicus. (;. 1922. Die Kntsl(>hunti der Chromato-
I.hon-n aus Chon.lriosomen Ixn Ildo'h't nni,i<liiis>s.
Jahrb. Wiss. Bot.Ol: 430-458.
(;kk<;ory. U. r. 1915. On variesation hi PrimuUi

sinrnsis. Jour. (Jenetics 4: 305-321.
GuiLLTKHMoxD, A. 1912. Rechcrches cytolojiiques sur le
mode (if formal iim de ramidcm .^ur les plas'es
ve<:claUN. Archiv. Anat. Micnwop. 14: ;:09-428.

"" 1924. Reeherehes sur rJvolu!i(m du chondri-

ome pendant le dcv(>loppement du sic embrytmnare
Pt des cellules mires des grains de poll(>n dins l;'s
Liliacces ct mr la sijinification des formation erjias-
toi)lasmi(|ues. Ann. Sci. Xat. Bob X. 6: 1-52.
Haan, H. I)K. 1933. Inheritance of chlorophyll <le-

ficiencies. Bil)liojjr. (Jeneticu 10: 357-116.
Hkimxxs. J. 1928. Chromosomen und Bcfnichtun-z j)ei
Liliitm Miutnuou. Rec. Trav. Bot. Ncerl. 25A:

138-167. ,^ ., ,

KuRssANow. L. 1911. Cber B(«fruchtuiif:_, Reifun- und
Keimmi}^ l)ei Zijgiuma. Flora 104: 65-84.

Lewitsky. O. a. 1925. Die Chondriosomen in der
Gonofiene.se bei Eqnisd urn jHilnstrc L. Planta 1:

301-316.
Meyer, A. 1920. Morpholojiische und Phys:olosische

Analy.se d(>r Zelle der Pflanzen und Tiere. Jena.
MiLE.s. F. C. 1915. A frenetic and cytolojrical study of
certain types of albini.sm in maize. Jour. Genetics
4: 193-214.
Raxdolph, L. F. 1922. Cytolojiy of chlorophyll types

of maize. Bot. Gaz. 73: 337-375.

RuHLAXi). W.. AND K. Wetzel. 1924. Der Xachweis

von rhloroi)lasten in den {lenerativen Zellen von

Pollen.schlaiichen. Ber. Deutsch. Bot. Ges. 42: 3-14.

SciiAFFXER. J. H. 1897. Contribution to the lifejiistory

of Sntfilldiiii vnrinbilis. Bot. Gaz. 23: 252-273.
ScHiMi'ER. A. F. W. 1885. rntersuchun«>;en iiber die
(Miloroi)hyllk.)rper und die ihn(>n homologen
Gel)ilde.
TRr)Ni)i.E, A. 1911. t'ber die Rcductionsteilung in den
Zyjrolen von Siiirot/i/ra und iiber die Bedeutuug der
Synapsis. Zeitschr. Bot. 3: 593-619.
TsiXKX, S. J. 1923. Reeherehes histolo^iques et cvto-
lo^dtiues sur la Panachuiv dans 1(> jienre Abiilil<nt.
Bull. Soc. Sci. Nancy IV. 2: 2.5-26.
W.\(;ner. N. 1927. Sur la formation " de novo" des
chcmih-iosomes dans l:i cytoplasmi« des cellules-
mcr(>s des grains de jmllen chez les aniiiosperms.
Biol. Generalis3: 329-346.
WixcE, O. 1919. On th(> non-Mendelian inheritance in
varie^jated i>lants. Gompt. Rend. Trav. Lab. Garls-
berj:34: 1-20.
Y.xsui. K. 1930. Studies cm the maternal inh(>ritanco of
l)las id characters in Ilostn jnixmica Aschers et
(;rael). f. alhomnnjinala Mak. and its derivatives.
Cytolonia 1 : 192-215.
ZiHKLFX C. 1929. Development of normal and diverg-
ent plastid types in Zva Mays. Bot. Gaz. 88: 186-

203. , , ,
1930. The u.-^e of n-butyl alcohol in dehydrat-

in<z woody tis.Mie for paraflin embeddinti. Science

71: 103-104.

1934. Amines in cytolofiical fixing fluids. Pro-

toplasma 20: 473^82.

AN UPPER CRETACEOUS WOOD: TORREYA ANTIQUA

Irwin Boeshore and William D. Gray

iTf^

Oct., 19361

ftOESrtORE AND GRAY — TORREYA ANTIQUA

525

h

I

AN UPPER CRETACEOUS WOOD: TORREYA ANTIQUA^

Irwia Boeshore and William D. Gray

The fossil wood here described was collected from
the Upper Cretaceous of Bhick Creek Age near Fay-
etteville, North Carohna, by H. A. Rankin. The
original 'specimen was a fragment of secondary wood
measuring one foot long, two inches wide, and one
inch thick. It is in a fair state of preservation but
with non-uniform infiltration of mineral substances,
showing regions of a white hyaline substance and
other regions of a soft, brown-colored substance (fig.
1). The wood is without fusiform rays, tracheae,
wood i)arenchyma, or resin i)assages of any kind; it
consists wholly of pitted tracheitis with consi)icuous
spiral thickenings. The latter feature and the ab-
sence of tracheids in wood rays are the chief diag-
nostic characters which ally it promptly with the
genus Toncija {Tiimion of Rafinesque).

Since spiral thickenings also occur on the tracheid
walls of the secondary wood of Pseudotsuga, Taxiis,
Larix americam, and Finns taeda, additional evidence
as to the identity of our specimen is presented. The
presence of well developed spirals on the tracheid
walls of the entire growth ring in Torreya and Taxus
is a constant and tvi)ical feature, while in Pseudo-
tsuga, Larix, and Pinus the spirals occur in more or
less isolated regions and may be imperfectly de-
veloped. Furthermore, the resinous containers are
absent from Torrciin and Taxus, but present in
Pseudotsuqa, Larix. ;.nd Pinus. The well developed
spirals in the fossil wood and the absence therefrom
of resinous containers exclude it from the last three
genera. The main choice for its position then lies
between Torrem and Taxus. The structure of our
specimen conforms more to the wood structure of
Torreya in the large, ( hiefly squarish, tracheids with
,1 ' . . , :^ 1 ;„ .»_« «orie*' ind »n tlip oval or

rulllt*! open .^piiiii.^ iiJ --t: b\.li^-, aii I -

1 Received for publication May 15, 1936.

oblong ray cells in tangential view, than to that of
Taxus with its small, chiefly rounded, tracheids with
rather close spirals in 2 (rarely 3) series, and the
narrowly oblong ray cells in tangential view.

A consideration of the distribution of the present
day representatives of Torreya reveals that the four
living species comprising the genus are confined to
more or less localized areas which are widey sep-
arated from each other. The two American species
are separated by the breadth of the entire continent;
T. taxifolia, the Florida Stinking Cedar, being found
on the ridges which extend into the Apalachicola
l^iver swamps, while T. californica, the California
Nutmeg, is found in the Santa Cruz and Sierra
Nevada mountains. The two other living species of
Torreya are Asiatic; T. nucifera being found in the
mountains of Japan, and T. grandis occurring in the
mountains of northern China. In addition to the
four si)ecies just listed, a fifth, T. Fargesii, has also
been described; this species, however, is little known
and is considered by some to be merely a variety of
T. grandis.

Historical. — There have been many reports of
the occurrence of fossil representatives of Torreya;
the s])ecies generally being described on the basis
of leafy branches or seeds. Such rejiorts are from
widely separated locations and inchcate that the dis-
tribution of the genus in i^ast geological times was
much wider than the distribution of the living s])ecies.
Berry (lOOS) pointed out that Torreya is not strictly
si)eaking a member of the existing flora but rather a
relic of a prehistoric one; furthermore, he predicted
that only a few centuries will elapse until the genus
will be extinct. This i)redicti(m was more or less
refuted by Sellards (1014), who stated that T. taxi-

Florida trees now ranks seventh in abundance.

The following reports of findings of fossil Torreyas
will serve to give some idea of the former distribution
of the genus:

Heer (1875) named two species, T. Dicksoniana
and T. parvifolia, from the Kome beds (Cretaceous)
of Greenland; in 1883, he named another species,
T. boreal is, from the Tertiary of the same country.
All three species were named on the basis of leaf
characters, although in the case of T. Dicksoniana,
cones were also found. Sa porta and Marion (1876)
described an undoubted species of Torreya from the
Pliocene of France which they named T. bilinica.
Saporta (1879) frequently mentioned the genus Tor-
reya as being a constituent of the Tertiary flora;
he cited no species from the Oligocene or the Mio-
cene but mentioned the occurrence of T. nucifera
and T. nucifera var. brevifolia in the Pliocene of
France. Dawson (1882) described T. densifolia from
the Upper Cretaceous of Protection Island, and T.
Dich-sonioides from the Middle and Ui)per Cretaceous
of the Pine River region, British Columbia; the
latter species was described as being near to T.
Dlchsoniafia Heer. Lesquereux (1883) described T.
oblanceolata from the Cretaceous black shale near
Golden, Colorado (see Knowlton, 1898). Fontaine
(1889) added two more si)ecies, T. virginica and T.
falcata, from the Potomac formation (Younger Meso-
zoic) of Virginia. He described T. falcata as being
quite similar to T. parvifolia, reported from Green-
land by Heer. Zittel (1890) mentioned Torreya as
being one of the companion genera of Gingko dur-
ing the U])i)er Cretaceous l)ut added no new species
or localities to the list. Zeiller (1900), discussing the
flora of the Tertiary, mentioned the finding of T.
nucifera in the Pliocene in the vicinity of Lyon,
France; however, it is iiossil)le that he was referring
to the same sj^ecimen which Sai)orta had previously
listed. This citation is particularly significant in
view of the fact that T. nucifera, as we now know it,
is a living species which occurs only in the moun-
tains of Jai)an. Menzel (1900) insisted that Sequoia
Langsdorfii Ettingshau.'^en was identical with Torreya
bilinica Saporta and Marion. It is possible that the
jiarticular specimen which Menzel examined was really
a Torreya, but most of the numerous records of Sequoia
Langsdorfii really re])resent Sequoias. Oliver (1903)
mentioned no particular species but stated that Tor-
reya is recognized as far back as the Lower Cre-
tace(ms. Berry (li)08, 1914) described T. caroliniana
from the Mid-Cretaceous of North Carolina; in 1919
he collected this same s])ecies from the Up])er Cre-
taceous near MacBride's Ford, Georgia. Krausel
(1918) de.scribed T. miocenica from the Upper Mio-
cene of Silesia; this sj)ecies was described on the
basis of seed characters, and cellular details of the
seed coat w'ere described and figured. Chaney (1925)
listed two unnamed species of Torreya from the
Lower Eocene of the John Day Basin, Oregon.
Hollick and Martin (1930) described two species

T.

along the Yukon River were made, and T. suspecta,
a rather doubtful species of which four collections
were cited. Dorf (1933), in discussing the Oligocene
flora of California, cited Torreya as being one of the
typical redwood forest types of that period.

In all of the above citations wood structure is not
mentioned in a single instance; it appears worth-
while, therefore, to give a detailed description of the
wood structure with the hope that it may add some-
thing of value to the knowledge of fossil Torreyas.

Preparation of material. — Polished transverse,
radial, and tangential sections of the fossilized wood
were fastened to ordinary glass microscope slides
with a glue consisting of four grams of j^owdered
rosin to one gram of lanolin, then ground to micro-
scoi)ic thinness and i)()lished. They were then dis-
solved from the slides with xylol and placed in a
mixture consisting of equal parts of xylol, ethyl
alcohol, and butyl alcohol. This mixture, which is a
modification of that advocated by Zirkle (1934) as a
cleaning solution, i)roved to be quite useful in rapidly
clearing the sections of ghie, particles of polishing
comiKJund, etc. Sections were then washed with
xylol and mounted in balsam for study.

Torreya antiqua sp. nov.

Traxsversk. — Growth rings are inconsj)icuous and
variable in width (fig. 4, 5). The spring wood con-
stitutes most of the ring; tracheids are squarish (fig.
14), uniform in size, mostly in regular rows, occa-
sionally in irregular rows, diameter of lumina 16-49
microns, the majority being about 40 microns; walls
of tracheids 7-11 microns in thickness. Summer
wood of 1-5 rows of tracheids with radial diameter
of lumina 7-14 microns; change to spring wood
abrui)t. Wood rays 1 cell in width; distant 2-12
rows of tracheids, length of cells about 175 microns.

Radial. — Rays without tracheids; ray cells straight
and uniform in width dig. 13); terminal walls gen-
erally straight and at right angles to the long axis,
thin; upper and lower walls rather thin; lateral
walls with small roimd jiits, 2-4 per tracheid (fig.
11). Bordered pits in 1 :~eries abundant on radial
walls of tracheids (fig. Hi; j)its rarely in two series.

Taxgkxtial. — Rays from 1-21 cells in height,
mostly uniseriate but with paired cells in places (fig.
10); ray cells with rather thick lateral walls and
small, oval or rounded I'lmina. Tracheids long (ap-
proximately 2.4 mm.); lew tracheids with bordered
pits on tangential walls g. 12). Spirals of tracheids
in 2-4 series in the s])riiij wood (fig. 2, 8) and i)lace(l

iiUiii

M 1.

1 <K

MUlii

1 1

4 1.

gracillima, of which citations of sixteen collections

at apj)roximateIy 45
the tracheids.

Biseriate rays are al>»
dorfii but any close c<
Sequoia is barred bec.i
on the tracheid walls a,
sages in the latter genu

Wood structure of T.
rings (fig. 3, 6). The

irles with the long axis of

ej)orted for Sequoia Langs-

lection of T. antiqua with

' of the absence of spirals

the presence of resin pas-

■xifolia and T. californica.
0( iCo ii,(v< definite gnjuih
cheids are squarish in out-

4^11

II

i

Oct.. 1936]

BOI'^SHOIJE AND (iHAY — TOUKEVA ANTIQUA

527

showing s,„n, 'hjokomn.. of n.^^^^^^^ ,.i„^ „, „,„„„„, ,,„„.,_ fi,. 5.

]

line, in regular rows, with occasional irregular rows.
Ihe diameter of the hunina in the si)ring wood varies
from S-35 micions in ta.vifolia and from 18-42
microns in calif arnica. The walls of the tracheitis
average al)out (5 microns in thickness in both si)ecies.
'Ihe sunmier wood of taxifol a, of 2-4 layers of trach-
eids, i)asKes abrui)tly into the s])ring wood, while
in caitjornica theie are ()-7 layers of tracheitis and
the trans't'on to si)ring wood is gradual. The walls
of the tracheitis in the summer wood average S
mic.ons in lH)th sjiecies. The wood rays are distant
1-MO rows of tracheitis in ta.vijolia and 1-lS rows in
califoniica. Length of ray cells 70-215 microns in
taxi folia; 115-285 microns in califoniica.

Radial. — The wood rays are withtmt tracheitis in
both sj-ecies. The terminal walls of the ray cells are
both straight and curved in tax'fuVa antl generally
straight but .sometimes placetl oblitjuely in cnl'foni'ca;
termdial walls are thin in both si)ecies. The ui)i)er
and lower walls of the ray cells are also thin in l)()th
s];ec:es. The lateral walls have 2-5 small i)its ])er
tracheitl in tax'folia and l-<) jiits })er tracheid in
califoniica. The bortlered ])its of tracheitis u-ually
occur in only one series but occasionally may l;e
found in two series in both tax'foVa and califoniica;
they are circular in outline and rather close in taxi-
fol'a but are ellii)tical antl somewhat distant in
califoniica.

Taxcjential, — The wootl rays are 1-21 cells in
he:ght in taxifol'a, anil 1-17 cells in cal'fornica (fig.
7, 9). In both s])ecies the cells are oval t)r oblt)ng.
Tlie tracheitis average about 2.8 or 2.4 nun. in length
in taxifol'a; in cal'fornica they Vary from 1.6 to 3.8
mm. in length. In l)oth species the bordered pit^
are chiefly on the radial walls of tracheitis, but are
occasionally ff)untl on tangential w:dls. The s])irals
of the tracheitis are 1-2 (mostly 2) series in taxifol'a
whereas in cal-fornica they occur in 2-5 series. In
taxifolia the s])iral< are i)laced at .slightly less than
right angles to the longitudinal axis of the tracheitis;
in cal'fornica thev roughlv ai)proximate an angle
of 45\

Comparison of T. antiqua w'th T . tax'folia
and T. calf arnica

TiJAxsvERSE. — Growth rings are more conspicuous
in both ta.vifolia and califoniica than in antiqua, the
sununer wt)od of the latter often consisting of only
t)ne rt)W t)f tracheitis (fig. 5). In all three sjiecies
the tracheitis are stjuarish in outline; the average
diameter of the lumina of antiqua is greater than
the diameter of the lumina of the other two species.
The tracheid walls of the ft).-<sil are thicker than
those of the living s])ecies, but this may simi)ly be
due to the fo.ssilization jiroce.ss. In the number of
layers of tracheitis comprising the sununer wootl,
antiqua, with 1-5 n)ws of tracheitis and an abrupt
change from s])ring to sununer wt)f)d, more nearly
ai)i)roximates taxifolia than californica, since the
latter sjjccies })o.s.<esses <)-7 layers t)f tracheids in the
sununer wood antl the tran.'^ition irom sprmg to sum-

mer wood is more or lo— gratlual. In the character-
istic distance between v.t)cd rays, antiqua is closer
to cal fornica, where tin r; ys are tlistant 1-18 rows
of t.acheitls, than to tdxifolia, where they are 1-30
rows tlistant. The av( igt ratlial length of the ray
cells of antiiiua is less dim the lengths in either ta.v'i-
fol'.a ciV californica.

Radial. — All three sixcits have wood rays devoid
oi tiaciie.tls. 'ihe teinhual walls of the ray cells are
generally stiaight in ant. qua as comparetl with the
stia.giit t)r curved te.-minal walls t)f taxifolia antl
the siiaight or ol)lit|ue walls of californica.. Pits
between lay cells ansl tratheitls vary from 2-4 i)er
trache.tl in ant qua, 2-5 per tracheitl in taxifolia, and
]-() jier tracheid in cal fornica. Bortlered pits on
tracheitl walls are circular in antiqua and taxifolia
and ellijitical and somewhat tlistant in californica.

TANtiENTiAL. — Wootl rays vary from 1-21 cells in
height in antiqua and tax'folia, while in californica
they vary from 1-17 cells in height. Biseriate rays
occur nuich more fret]uently in antiqua than in either
taxifal a oc cat fornica. Spiral thickenings occur with
2-4 (usually A) .-^erie- in antiqua as comjjared with
1-2 series in taxifolia and 2-5 series in californica.
The ang.e of in.-ertit)n of spiral thickenings is alnnit
the .same (approximately 45 ) in antiqua antl cali-
fornica, while in taxifolia it more nearly approaches
a right angle. The .-^ijiral thickenings are witler in
antiqua than are tht)se of the other two species;
however, this may also be tlue to the fo.ssilization
l)rocess.

T. antiqua has .st)me characteristics in common with
T. taxifolia as well as with T. californica; in still
other characteristics it "lifters from the other two
sj)ecies. A tiuestion of nomenclature arises, .since
Berry (1908, 1914) named Tuniian carolinianum from
the same general regit)n from which this species was
collected but mentioned no wootl structure in tle.scrib-
ing liis sp,ec;es. As no leaves or seeds were ctjllectetl
with the specimen here tlcscribed, it seems appro-
])riate tt) a.-^sign a ditferent sjiecific name from that
of Berry's specimen until further data are obtained

AcKNowLEDtjMENTs.— The autliors are indebted to
Dr. Rodney II. True, Chairman of the Department
of Botany, for .securing grinding apparatus used in
the i)reparation of material: to Dr. Etlgar T. Wherry,
University of Rennsylvania, for the specimen of fos.sil
Torrcija here tl.-ecribeil. They also wish to thank
Dr. Herman Kurz, Flt)ri<la State College for Women,
antl Dr. W. L. JejisoiL University of California, for
s])ecimens of T. taxifol i<i mtl T. californica. This in-
vestigation was aitled in part by a Faculty Re.-<earch
grant frt)m the Univer-ity of Pennsylvania to the
.-senior author.

SIM MARY

The distribution of li\ ing forms of Torrcija is dis-
cus.'^ed briefly. The ft)ur living s])ecies are confined
to localizetl, witlely-sep.irnted areas. The literature
concernetl with distribution of fo.ssil repre.'^enta fives
is reviewetl, anti all the tx itlence ciearlv intiicates that

M

Oct.. 19.U. 1

BOKSJIOKK AND (iKAV- -TOUHKVA ANTKilA

5-J7

hMl. 1. Alan. .<..!< ^u^^ ." ' ' ., ., -|-,,„., ,,,,,. ,,.,.,i..n el T. In.r>inha: n.,lc chiHi^n. Iron. >i.nny: to

' ..clioM of V. anlKiua showinir a li\ c-lMycica liny; ol suiimu'i wood. — I iji. a.

Tl■a^•"^^ <'i"''f' sanction ol

' ' ■ .Fiii. 7. Tanirmtial scchon ot 7. /r/j-/-

f

oclion of r. ra/./o,...; not,- ^ra.lual tiansilion fro.n .pnn^ lo sunnn..; woo,i.-ri«. .. . a.ui.n. >... -''"'•- ^

o m howin.^ <pral ihi.' -„in- of Ira.-lMi.l^.-Kiu. S. TanK^nhMl hm-Iiou ol / . ..//r,.m showinu spiral .l,H-krn,iu:
Zj^l^u^^lw I 'Ln that of liu. 7 an-l 0).-Fi.. U. Taim..nhal ..Hion ol 7. .. ,.f...... >howm. .,. a

u^k n--F... 10 ■ T ,...nlial mH.ou of V. .,,.1.,..,; unU- ,.;.nv.l n-11-^ ... .uo ol ihr w<mhI ...y. -1' ... 11. Ha.l al

huM n.n..^^ ... . . 1-1 _ ^^^^ ^^^^ ^^ ^^^^^ ^^ ^^^^^11 ^ .^^ l„,,w.-rn .av nlN and t.acl.c.ls n.v «■.... -I'ltf. 12.

;:;::;:' ;;:.rs:.H,n,; of 7- ;„/....; ..o.; l>onl.iva p.U on .anu.n.ial a. .hi a> .a.i.al wans ,., ,radi.„ls.-lM. I.>.

Tanncnlial si-ction of 7' <n,li(j

4
tl

lifadual. In the cliaractcr-
oMJ rays, (uitujun is closer

r. >s arc d. slant 1-lS rows
ifi>l:(i, where they arc l-oO
i( radial lenjith of the ray
11 the Icimih- in e.ther taxi-

Radial section of T. uhIhjuu showing nature ol woo

J j-ay^. — Fig. 14. Tranisverso sec

lion of spring wood of T. «/t//V/aa.

line, in rcjiiilar rows, with occasional irregular rows.
'1 he diameter of the lun.ina in the .s])rin<i' wood vanes
from s-o') micions in t<Lv'ijt)}iti and from lS-41.'
microns in caijoniica. d he walls of the tracdicids
avei'a'ic ahout <) microns in thickness in both spet-ies.
'Ihe siunmer wooil of td.c'iol a, of 2-4 la>-eis of tra(di-
eids, ])as-es :d»ni])tl\- into the sprinji wood, wliile
in (■(ihforn.cii theie are <>-7 layers of traidieids and
the transjJMi to s])riim' wood is gradual. The walls
of the tiacheids m the summer wood average N
iiLcons in hoth sp,ecies. The wooil rays are distant
l-MU row- of tra(dieids in taxijolUi and 1-lS rows in
ralifornicii. Leiinth of ray cells 70-21.') microns in
taxifoL'd : lla-^S') microns in cnlifornlca.

Hadiai.. — "Ihe wood rays -die witlio.it tracheids in
both sjiecies. Tlio terminal walls of the ray cells are
l;olh straiiiht and cur\-el in tax fold and jicnerally
straijiht Imt .sometimes jilacod ohlicniely in atijont'ca:
torniJial walls are thin in hoth sj.ocies. The upper
and lower walls of the ray cells are also thin in hoth
sjiecies. Th.e lateral walls have 2-') sihmII pits j.er
tra(dieid in inxjolUi and l-t» i)its j.er tracheid in
(■(illfttniicti. The honlered ])its of lr:i(die;ds u-.idly
occur in only one sor.e- l»ut occasionally may l;e
found in two series in hoth tnx'j(tl'(i and mlllant'cd :
they are circular in outline and rather close in tdx!-
fol'd hut are elliptical ami scjinewh.it distant in
r(;lif(t))uca.

Ta.\«;k\tial. — The wood rays are 1-21 cells in
heulit in t(iX'i(fl'a, and 1-17 cells in (-(djitrn'irn (liii.
7. !h. In lioth species the cells are oval or ()l)Ion'^
'idle tracheids averaiic ahout 2.."-» or 2.4 mm. in leii'ith
in t(tx.joi(i: in rdl'fofnirn they varv from l.ti to o.S
mm. in Ieny:'h. in hoth specic^s the ho'-deved jiits
are (di:efl\- on the radial walls of traidieids, hut are
occasionally found on fan'zential walls. The sjiirals
of the traidieids are 1-2 (mostly 2) ser'es in tdX'fol'a
whereas in r<il''j(>ni'c<i they occur in 2-7) .series. In
tdxifolin the sj)ira|s are placed at slightly Ie.<s than
riiiht an«ile> to the loniritudinal axis of the tratdieids;
in ((if htmica they rouixhiy api)roxiniate an aiijile
of 47) .

( 'oiii iKinsoii nj T.fnibf/ud irth 7'. tax fohd
oiul T. lud joniicd

Ti{A\svi:i{sK. — firowth rinjrs are more conspicuous
in hoth t(ixif(»l'(i and cidijitni'icd than in antifiua, the
summer wood of the latter often consistinj; of only
(»ne n»w of tra<dieids (liir. ')). In all three species
the tra(dieids are scpiarish in outline; the average
diameter of the lumina of nfitif/iui is greater than
the diameter of the lumina of the other two s])ecies.
The tra(dieid walls of the fossil are thicker than
those of tile liviiijr species, luit this may sinii)Iy he
due to the fossilization process. In the numher of
layers of tracheids comprisin«r the summer wood,
anti(/un, with 1-7) rows of tra(dieids .and an ahrui)t
ehanire from s|)rin<j: to summer wood, more nearly
ai)proximates tnxifoliti than (nhfonncn, since the
hitter snecies i,ns-('v^(v< il-7 I.i\ei- of tracheids in the
.-.immer wood and the tr.ii.sifion from sprinii to sum- is re\iewed. and all the i<!ence clearly indicates that

nicr wood is more or h
Istic distance helween
lo cal foniicd, wlicic fli
of 1.a(die;ds, than lo I,
rows distant, ddie a\c
cells of (nitujiid is less il
ji)l.(i or cdllfornicd.

Kadial. — All three -] r,'i( > have wood rays devoid
of tiacue. ds. 'ihe leim nil walls of the ray cells are
j;ene-aily sl.aijihl in iii'(jid as compared with the
.-i.a.<iiil or c.irved le.in.nil walls of tdxifol'd and
ihe s,ia.<ilit or ohrunic w ills of cdlifdnilcd.. Pits
I.etween ia.\- cells and iraiheids \ar.\- from 2-4 jier
ti'aciie.d in diii (/ua, '2-') pe- tracdieid in tdx'ijolht, and
1-1) per tiacdieid in nil joniicd. Bordered j.its on
tiatdieal walls are circular in (Uitiqud and tdxifolid
and ellii-tical and somewhat distant in CdiijoDi'iCd.

Ta.\(;i:ntial. — Wood rays vary from 1-21 cells in
he.jiht in dntif/ud and taxfolid, wdiile in cdlifonilcd
lliey vary from 1-17 cells in heiy;ht. Bi.seriate rays
oi'cui" much more fre(|uenily in dNt.'({dd than in either
idxijind (),• cdl joniicd. Sjiiral thi(d<:eniiiii;s occur with
2-4 (usually 4| .series in (iiiti<jHd as compared with
1-2 series in tdx'fot'a and 2--7) .series in cdlifonilcd.
'Ihe aiiff.e of insertion of .-j.iral thickeninjis is ahout
the .same (ai)proximatel\ 4.") ) in antirjud and cdll-
(ornicd, wdiile in tdxiiol'ii it more nearly approacdies
a rijilit aiisile. Tlie spiral thickeninjis are wider in
nnt'(/ud than are those oi the otlier two si)ecies;
however, this may al-o he due to the fossilization
proce.ss.

T. <uiti(ji(d lias .some cli.iracteristics in common with
T. tdxIfolid as well as with '/'. <(jlifoni:ca ; in still
other (diaracteristics it differs from tiie other two
sj.ecies. A (|uestion of iionieiKdature arises, since
Berry ( lUOS, ]iU4) named Tianion cdroimidnmn from
the same jreneral reirion from whi(di this species was
collected hut mentioned no wood structure in descrih-
inir his sp.ecies. As no Iea\es or .seeds were collected
with the specimen here de.sorihed, it .seems apjiro-
j.riate to assijrn a different specific n.anie from that
of lierry's specimen until further data are ohtained.

A('K\o\vm:i><;mknts.- '1 he authors are indebted to
Dr. Rodney II. 'I'rue, (ha rman of tiie l)ei)artinent
of Botan.\', for sccuriim irinidini; apj.aratus used in
the preparation of material: to Dr. IM{rar 'l\ Wherry,
i'niversity of Bennsylvaiiia, for tiie s])ecimen of fossil
Torre !id here d.secribed. 'I'liey also wisii to thank
Dr. Herman Kurz, Florida State Colleire for Women,
and Dr. W. L. .lepson. T'niversity of California, for
sjK'ciniens of T. tdxijolm iinl T. cdlifornird. This in-
vestijiation was aided in part by a Faculty Ke.seandi
irrant from the rniyer-ity of IVnnsylvania to tlie
senior author.

<5t rMAUY

The distribution of li\ tip forms of Torrvija is dis-
cussed briefly. The fn ' living s])ecies are eonfined
to localized, widely->cp,!' iteil areas. The literature
concerned with di>tiil <»n of fossil rei)resentati\es

rj5

or>

528

AMERICAN JOURNAL Of BOTANIT

[Vol. 2i,

in past geological times, certainly during the Cre-
taceous and Tertiary, the genus had a much wider
distribution than it has at present. Fossil Torreyas
heretofore reported were described on the basis of
leaf or seed characters.

Torreya antiqua is described on the basis of second-
ary wood structure. The structures of T. taxifolia
and T. cnl'fornica are reviewed and are compared
with the structure of T. antiqua. T. antiqua is simi-
l;ir in some characters to T. taxifolia, in others to
T. californica, and in still other features differs from

both. T. antiqua shows a greater abundance of bi-
seriate rays than the other two species, the lumina
of the tracheids are larger, the spiral thickenings are
wider, the radial length of the ray cells is shorter,
the amount of summer wood is considerably less,
and the growth rings are not so clearly defined as in
T. taxifolia or T. californica.

Department of Botany,
i'^niversity of pennsylvania,
Philadelphia, Pennsylvania

LITERATURE CITED

Beruy, E. W. 1908. A niid-cretaceous species of Torreya.

Amor. Jour. Sci.. 25: 382-386.
1914 The upper cretaceous and eocene floras

of South Carolina and Georgia. U.S. Geol. Surv.

Prof. Paper 84: 107.

1919. I'ppor cretaceous floras of the eastern

uulf ro'Mon in T( nnossee, Mississippi, Alabama, and
Goor-ia. U.S. Gool. Surv. Prof. Paper 112: 70

Cheney R. W. 1925. A comparative study of the
HridW C^rock flora and the modern redwood forest.
C^arnesie Inst. Washinston Publ. No. 349, Pt. 1: 13.

Dawson J W. 1882. On the cretaceous and tertiary
floras of l^ritish Columbia and the North West
Territory. Trans. Roy. Soc. Canada, vol. I, sec. IV:

15-34. » ... . r^ •

DoRF, E. 1933. Pliocene floras of California. Carnegie

Inst. Wa.shinjiton Publ. 412: 96.
FoNT.AiNE, W. M. 1889. The Potomac or younger meso-

zoic flora. U.S. Geol. Surv. Mon. 15: 234-235^
Heer, O. 1875. Flora fossilis arctica, III, Pt. 2: 70-72.

Zurich. ^- rr • U

1883 P^lora fossilis arctica, VII: 56^7. Zurich.

Hollick, a. and G. C. Maktin. 1930. The upper cre-
taceous floras of Alaska. U.S. Geol. Surv. Prof.
Paper 159: 55-56.

Knowlton, F. G. 1898. A catalogue of the cretaceous

and tertiary plants of North America. U.S. Geol.

Surv. Bull. No. 152: 234.
Krausel, R. 1918. Nachtrage zur Tertiarflora Schlesi-

ens. I. Jahrb. d. Preiiss. Geol. Land.: 342-345.
Lesquereux, L. 1883. Contributions to the fossil floras

of the territories. III. The cretaceous and tertiary

floras. U.S. Geol. Surv. of the Terr.: 30-31.
Mbnzel, p. 1900. Die Gymnospermen der nordbohm-

i.schen Braunkohlenformation. Abh. Nat. Ges. Isis:

104-106.
Oliver, F. W.1903. The ovules of the older gymno-

sperms. Annals Bot. 17: 451^76.
Saporta, G. 1879. Le monde des plantes avant I'appari-

tion do rhomme. Paris, p. 196, 253, 306, 332.
AND A. Marion. 1876. Recherches veget. foss.

de Meximieux. Nouv. Archiv. du Mus. d'hist. nat.

Lyon, t. 1. livr. 4: 131-355.
Sellards, E. H. 1914. Florida state geological survey.

Sixth annual report. Tallahassee, p. 212, 215, 354,

400, 412. .

Zeiller, p. 1900. Elements de paleobotanique. Pans.

p. 258-259.

ZiRKLE, C. 1934. Butyl alcohol and cytological tech-
nique. Science 80: 481-482.

Zittel, K. a. 1890. Handbuch der Palaeontolo^ie.
Miinchen und Leipzig, p. 258, 298.

1

H

m

'^nr

104

Cytologia 7

1936

Effects of several fixatives on staining reactions in Zea mays

105

Effects of Several Fixatives on Staining Reactions in Zea mays.

Especially with Reference to the Feulgen Reaction

By
J. Gordon Carlson

Biological Laboratory, Bryn Mawr College

(With 2 Plates)

Received June jg, 1934

Introduction

For the past few years cytologists have been interested in the
Feulgen nucleal reaction as a staining method "specific for
chromatin". This specificity is valid, however, only if we are willing
to identify chromatin with thymonucleic acid. Investigations by both
biologists and chemists have tended to confirm this identity, though
there is still disagreement as to just what proportion of the chromatic
material of the nucleus is chromatin. There is some evidence to
indicate that plant chromosomes are composed of both plastin and
the condensed chromatin of the reticulum (Lenoir '22; Van Camp
'24; Martens '25; Zirkle '28, '31).

Since chromatin reacts with the Feulgen stain because of the
presence of an aldehyde freed in the hydrolysis of thymonucleic acid,
Feulgen and Rossenbeck ('24) avoided fixatives containing form-
aldehyde. They fixed their sectioned material with a solution of
corrosive sublimate and acetic acid and their smear preparations by
drying. Bauer ('32), however, using the Feulgen reaction after a
variety of fixatives, found that formaldehyde in the fixing fluid did
not alter the selectivity of the fuchsin-sulfurous acid stain, even
though this stain reacts with the aldehydes. He concluded, there-
fore, that, if the formaldehyde were only adsorbed on the fixed
tissue, it would be removed in the subsequent washing and dehydra-
tion, while, on the other hand, if it were chemically united with the
proteins of the cell, its properties would be altered to such an extent
that it would no longer react with the fuchsin-sulfurous acid.

Another line of approach to the problem of stain selectivity is
represented by the recent work of Zirkle ('28-'33), who has in-
vestigated the effects of different fixatives on the mordanting of
plastin, chromatin, and mitochondria in the plant cell. He concluded
that the staining capacity of these cell organs was conditioned by both
the components of the fixative and the pH at which the fixation
occurred. Certain fixatives that mordanted the chromatin fixed the

il

I

I

plastin in such a manner that it did not retain the dye. Other com-
binations mordanted the plastin but not the chromatin. Still others
mordanted both substances. The only stain he used was Heidenhain's
iron alum-hematoxylin.

The generally admitted fact that the Feulgen reaction has a
specificity for chromatin not possessed by such a stain as hematoxylin,
which can be made specific for any of the several cell parts, depending
on the fixative employed, directs attention to the problem of the
effects of fixation on stainability. It is a well known fact that many
of our commonly used fixatives differ in their capacities to mordant
certain cell components. Very little work, however, has been done
on the specific mordanting properties of fixatives with respect to dif-
ferent stains.

Four methods of staining are described in this paper. They
are the Feulgen reaction, Heidenhain's iron alum-hematoxylin, crystal
violet-iodine, and safranin-iodine. The crystal violet-iodine method
of Newton ('27) has been recommended by Sax ('31) as a useful
chromosome stain after fixation with Navashin's solution. Safranin
was also investigated, since it is commonly used with crystal violet
in Flemming's tri-color stain, where these two dyes, when combined,
stain different cell organs. Inasmuch as the hydrolysis necessary
for the Feulgen reaction is effected by immersing the sectioned tissue
for several minutes in N HCl at 60°C., it seemed advisable to compare
the reactions of each of the other stains both with and without this
treatment.

Material and Methods

Root tips of Zea mays from seeds sprouted in sphagnum moss
were fixed from 40 to 48 hours.

The following fixatives were selected because of the varied and
representative types of fixation they offer.

1. Bouin: picric acid (sat. aq. sol.) 75 parts, formalin 25 parts,
acetic acid 5 parts.

2. Navashin: (A) formalin 50 parts, acetic acid 12 parts, water
38 parts; (B) chromic acid (10% aq. sol.) 15 parts, water 85 parts.
"A" and "B" were mixed in equal parts just before using.

3. Copper Bichromate: chromic trioxide 1.25 gms., ammonium
sulphate .25-50 gms., water 100 c.c, slight excess of copper
hydroxide.

4. Copper-Chrome-Propionate : chromic trioxide 2 gms., pro-
pionic acid 1 c.c, water 100 c.c, excess of copper hydroxide.

5. Nickel-Chrome-Propionate : chromic trioxide 2 gms., pro-
pionic acid 2 c.c, water 100 c.c, excess of nickel hydroxide.

IRREGULAR PAGINATION

.

106

J. G. Carlson

Cytologia 7

6. Formol-Sulf uric Acid : formalin 4 parts, sulfuric acid 1
part, water 95 parts.

7. Formic Acid-Acetaldehyde : (A) formic acid 6 parts, acet-
aldehyde 19 parts, water 75 parts; (B) chromic acid (107^ aq. sol.)
20 parts, water 80 parts. 2 parts of "A" were mixed with 1 part of
"B" just before using.

8. Modified Erlicki: potassium bichromate 1.25 gms., am-
monium bichromate 1.25 gms., copper sulfate 1 gm., water 200 c.c.
Two drops of pyridine were added to 15 c.c. of this mixture one-half
hour before using.

9. Chromic Sulf ate-Formaldehyde : chromic sulfate 5 gms.,
formalin 4 c.c, water 96 c.c, slight excess of copper hydroxide.

After fixation the tissues were washed for 1 hour in tap water,
and then dehydrated, cleared, infiltrated, and embedded by the butyl
alcohol method. The root tips were sectioned longitudinally 8 micra
thick.

The stains were used as follows.

Feulgen Reaction. Hydrolysis was effected by treating the
the mounted sections with N HCl at 60°C. for 20 minutes, a length
of time found by experiment to give an intense color reaction after
most fixatives. This treatment was both preceded and followed by
immersion for 2 minutes in N HCl at room temperature. The sections
were then placed in fuchsin-sulfurous acid, where they were allowed
to remain for 3 hours — ^the time recommended by Feulgen and Rossen-
beck C24) for plant tissues.

Heidenhain's Iron Alum-Hematoxylin. Sections were mordanted
for 2 hours in 4% iron alum, rinsed in water, and placed in .5%
hematoxylin the same length of time. For differentiation 2% iron
alum was used. One series of these sections received a preliminary
treatment with N HCl at 60°C. for 20 minutes.

Crystal Violet-Iodine. Sections were transferred from water to
1% aqueous solution of crystal violet for 5 minutes, then rinsed
successively in 35% and 50 7^ alcohols, immersed in 80 %> alcohol
containing 1% potassium iodide and 1% iodine for 30 seconds, and
differentiated in absolute alcohol followed by clove oil. One set of
slides was given a preliminary treatment with N HCl at 60°C. for
20 minutes.

Safranin-Iodine. Material was treated similarly to that stained
with crystal violet-iodine, except that a 27^ solution of the stain
was used and the time of staining was increased to 10 minutes.

Sections from two root tips on separate slides were stained by
each of the above methods, not only as a check against the abnormal

1936

Effects of several fixatives on staining reactions in Zea mays

107

I

1

\

reactions of pathological tissues, but also as a means of comparing
the results of different degrees of destaining.

Staining Reactions

The main results of this study are given in the accompanying
table. Since the staining experiments were run in duplicate, two
sets of data are given for each procedure. Differences in the results
obtained from the two series are due to slight variations in the length
of time of the destaining. The depth of color as indicated by the
double and single crosses is absolute for the Feulgen reaction series,
but relative for the other stains. This is an arbitrary but necessary
method of interpretation, because the Feulgen technique demands
constant, set times for the several treatments, a procedure that is
not practicable with the other dyes, where the subjective element
necessarily enters into the differentiation. All the observations
tabulated refer exclusively to the zone of cell division of the root
tip, as it was found that the cells of the root cap and zone of elonga-
tion occasionally react differently from those of this region — a
problem beyond the scope of the present investigation.

Reticular granules and reticular threads are distinguished in
the resting nucleus only for the sake of descriptive clearness, with
no implication that there is a more fundamental difference between
the two than the relative degrees of condensation of the chromatic
network. This matter is considered in more detail in the discussion.

Though it is a question to what extent the Feulgen *'Nucleal-
reaction" is a stain in the strict sense of the word, it is referred to
as a stain in this paper merely as a matter of convenience.

A. The Fetdgen reaction

This biochemical staining method gives essentially the same end
results regardless of the fixative used. At all mitotic and inter-
mitotic phases the chromatin alone reacts. The only observable effect
of fixation is the intensity of the stain. Chromatin fixed in Bouin,
nickel-chrome-propionate, formol-sulfuric acid, and chromic sulfate-
formaldehyde is lightly stained, while after the other fixatives
it stains a deep violet. Bauer ('32) , working with animal tissues and
a number of different fixatives, has investigated the relation of the
fixative and the length of the hydrolyzing process to the density of
the Feulgen color reaction. He found that for each fixative there
was an optimum period of hydrolysis, and that a longer or shorter
time failed to give a color of maximum intensity. His findings sug-
gest that a time of optimum hydrolysis could be found, specific for

108

J. G. Carlson

Cytologia 7

1936

Effects of several fixatives on staining reactions in Zea mays

109

Bouin

Copper-
chromate

Cxipper-
Chrome-

pionate

IMickel-
Chrome-

Pro-
pionate

Formol-

Sulfuric

Acid

Acid
Acetal-
dehyde

Modified
Erlicki

0 0)

ll

U OD

0

1

3

Chromosomes

Reticular granules

Reticular threads

Nucleolus

Mitochondria

Cytoplasm

Cuticle

+ +
+ +
+ +

H++
++H
++++

WW

++H
++++
++H

++++

1

++++

+ +
+ ' + '

4- +

+ +
+ +
+ +

H++
H++
H++

++++

H++

+ +
+ +
+ +

+ +

Iron Alum-
Hematoxylin

Chromosomes
Reticular granules
Reticular threads
Nucleolus
Mitochondria
Cytoplasm

H++
++++
+ +

+ +

++++
++++
+ ++

+ +

++++
++++
WW
++++
H++
+ +

++H
H++
++ +
++++

+ +
++++
+ +

++ +
+ +

+ +

++++
++++
+ +
+ +

+ +

+ +
++4+
++H
+ +

++H
+ +
H++
H++
+ +

Iron Alum-

Hematoxylin

preceded by HCl

Chromosomes
Reticular granules
Reticuler threads
Nucleolus
Mitochondria
Cytoplasm

+ +

+ +
¥t¥t

+ +

++++
+ ++

+ +

++++
+ ++
+ ++

+ +
+ +

H++
++H
+ +
++++

+ +

+ +
+ +

+ +
+ ++
+ +

++++
++++

++4+
++++
+ +
++++
+ ++
+ +

++++
++H
+ +
H++
+ ++
+ +

Crystal Violet-
Iodine

Chromosomes
Reticular granules
Reticular threads
Nucleolus
Mitochondria
Cytoplasm

++H
+ +

0

- +

H++
++++
+ +
+ +

++++
++++

++++

^^ ^^

- +

++++

^ —

Crystal Violet-
Iodine preceded
by HCl

Chromosomes
Reticular granules
Reticular threads
Nucleolus
Mitochondria
Cytoplasm

++++

++H

n n

WW

+ -

++++

++++
++++
++ +
++++

_3_3

+ +

•H-H
+ +

_-3_3

1

c

• mm

1

Chromosomes
Reticular granules
Reticular threads
Nucleolus
Mitochondria
Cytoplasm

++++

++++
+ H

0 «
- +

4+++
H++
- +
+ +

++++
H++
+ +

++++

7,

— —

+ +

+ +
++++

_3_3

Safranin-Iodine
preceded by HCl

Chromosomes
Reticular granules
Reticular threads
Nucleolus
Mitochondria
Cytoplasm

H++

n r

++++
= ++++

++++

+ +
++++

+ -

+ -

+ +

+ +
-++

+ ++

++ Deeply Stained. + Lightly gtained. -Unstained. 1 Stained peripherally. 2 Greater affinity for
stain in mitotic and epidermal cells. 3 Erratic, the stain evident in a few cells only.

each of the above fixatives, which would give a color of greater in-
tensity. This seems probable in view of the fact that hydrolysis for
less than 20 minutes resulted in a very slight color reaction after
certain of these fixatives but not after others.

A very constant and distinctive feature of this stain is the ap-
pearance of the intermitotic nucleus (pi. 8, fig. 2). The stained
chromatin forms a dense network of fine threads with granules of
different sizes. The only noticable variation occurs when the speci-
men is fixed in nickel-chrome-propionate. In this case the reticular
granules are absent and the reticular threads are thicker and less
sharply defined. The chromosomes are stained uniformly after all
of the fixatives except two, nickel-chrome-propionate and formol-
sulfuric acid. When these fixing fluids are used, the peripheries of
the chromosomes are colored more deeply than the cores.

Nucleoli are unstained at all stages. After those fixatives that
effect the formation of the perinucleolar vacuole, however, the
surface of the nucleolus appears to be faintly stained. The material
fixed with nickel-chrome-propionate shows this very clearly. Since
this fixative is also one that appears to dissolve the chromatin some-
what, it is possible to explain this reaction by assuming that a very
small amount of the dissolved chromatin accumulated at the nucleolar
boundary during fixation. Only the nucleoli of specimens fixed in
modified Erlicki and chromic sulfate-formaldehyde fill the space in-
side the reticular area ; and these show no trace of the dark borders.
It is also possible that a very thin layer of chromatin surrounds the
nucleolus and that this is exaggerated by the shrinkage of the
nucleolus that occurs after certain fixatives.

The cuticle of the root tip is colored by the Feulgen technique
after most of the fixatives. It has been known for many years that
lignin, suberin, and cutin contain aldehydes, as they give a positive
reaction to the Schiff test. Margolena ('32) reported that they
react with fuchsin-sulfurous acid either with or without preliminary
hydrolysis. Though aldehydes are present in the cuticle of the living
root tip, it is a question to what extent fixation and hydrochloric acid
treatment affect the amount present. Margolena makes no statement
as to the effect of hydrolysis on the intensity of the Feulgen stain.
There is a fixation effect, however ; for the intensity of the stain in
the cuticle varies after different fixatives as does that of the
chromatin. When the intensity of the chromatin stain is great, the
color of the cuticle is relatively dark (after Navashin, copper
bichromate, copper-chrome-propionate, and formic acid-acet-
aldehyde) ; but when the chromatin stain is less intense, the cuticle
is very light (after nickel-chrome-propionate and chromic sulfate-

110

J. G. Carlson

Cytolcgia 7

formaldehyde) or even unstained (after Bouin and formol-sulfuric
acid). The only exception to this is provided by the material fixed
in modified Erlicki, in which the chromatin is deeply stained and the
cuticle lightly stained. It is quite possible that fixatives containing
oxidizing agents destroy some or all of the aldehydes originally
present; for there can be no doubt that there is a definite fixation
effect. To what extent, however, hydrolysis may be effective in
creating new aldehydes to replace those oxidized must for the present
remain an open question.

B. Heidenhain's iron alum'hematoxylin stain

1. Bouin Fixation. Nucleoli, chromosomes, and reticular
granules are stained black, while reticular threads and cytoplasm are
stained light. Following hydrolysis the chromosomes and reticular
granules lose some of their staining capacity, though they still take
up more stain than the reticular threads and cytoplasm. One slide
shows deeply stained prophase chromatin.

2. Navashin Fixation. Chromosomes, mitotic nucleoli, and
reticular granules are black, while intermitotic nucleoli are deeply
stained only in the outer one or two cell layers. Reticular threads
and cytoplasm are light-stained. After treatment with hydrochloric
acid the nucleoli stain more deeply in relation to the chromatin. This
effect is erratic, however; for a majority, though not all nucleoli
show it (pi. 8, fig. 1).

3. Copper Bichromate Fixation. All the chromatin is stained
very heavily. Mitochondria are somewhat lighter, but darker than
the lightly stained cytoplasm. The only visible effect of hydrolysis
is to cause the mitochondria to stain less dark, so that they are
barely distinguishable from the cytoplasm surrounding them.

4. Copper-Chrome-Propionate Fixation. Chromosomes, reti-
cular granules and nucleoli are black, while reticular threads and
cytoplasm are light. Treatment with hydrochloric acid seems to have
no effect on the staining reaction (pi. 8, fig. 6).

5. Nickel-Chrome-Propionate Fixation. Nucleolus and reticular
granules are stained black; chromosomes, reticular threads, and
cytoplasm are light. Hydrochloric acid treatment, even for an hour
and a half, produces no visible effect.

6. Formol-Sulfuric Acid Fixation. The appearance of the
stained sections varies greatly with the amount of destaining. To
a certain point both nucleoli and chromosomes remain black; beyond
this point the former lose color very rapidly, the latter slowly. All
the nucleoli will not lose their color, however, before some of the

X936

Effects of several fixatives on staining reactions in Zea mays

111

1

chromosome color fades. A satisfactorily differentiated slide is one
in which most of the chromosomes are dark, with the reticulum,
cytoplasm, and most of the nucleoli light. Treatment with hydro-
chloric acid causes the nucleoli to stain erratically and the chromo-
somes to lose the stain more readily, so that these two structures
come to exhibit the same relative depth of stain.

7. Formic Acid-Acetaldehyde Fixation. This fixation resembles
that of the formol-sulfuric acid, in that the nucleoli lose their black
color very rapidly after a certain point in the destaining is passed.
The chromosomes and reticular granules, however, retain their black
color sufficiently far beyond this point that there is no overlapping
(pi. 8, fig. 5). Reticular threads and cytoplasm stain lightly.
Hydrolysis does not affect the stainability of the cell parts.

8. Modified Erlicki Fixation. Chromosomes, reticular granules,
nucleoli, and mitochondria are black, reticular threads and cytoplasm
light. Treatment with hydrochloric acid produces no effect pi. 8,
fig. 4). This is especially interesting in view of the fact that acid
fixing solutions usually dissolve mitochondria. Apparently, once
fixed, mitochondria are not destroyed by even a hot solution of normal
hydrochloric acid.

9. Chromic Sulfate-Formaldehyde Fixation. Chromosomes,
reticular granules, nucleoli, and mitochondria stain black, while
reticular threads and cytoplasm stain lightly (pi. 8, fig. 3). Hydro-
chloric acid treatment produces no effect on staining.

C. Crystal violet'iodine stain

1. Bouin Fixation. Nucleoli stain dark at all stages. The
chromosomes, the only other stained structures, show a somewhat
lighter color (pi. 9, fig. 7). The effect of hydrolysis is to render
the chromosomes unstainable.

2. Navashin Fixation. Sax ('31) has suggested the use of
this fixative followed by the crystal violet-iodine stain as a good
technique for chromatin. This is confirmed by the present study.
Only the chromosomes and the reticulum are stained (pi. 9, fig. 8),
though there is a very slight tendency for the nucleoli in the epidermal
layer and in dividing cells to show some color. Hydrolysis increases
the number of epidermal and mitotic nucleoli staining, and decreases
the stainability of the reticular threads (pi. 9, fig. 9).

3. Copper Bichromate Fixation. Chromosomes and reticular
granules are stained dark. Nucleoli are light, but have dark, sharp-
ly defined borders. The reticular threads stain lightly. The hydro-
chloric acid treatment results in a strikingly changed appearance of

112

J. G. Carlson

'ytologia 7

the nucleoli. Instead of light centers and dark peripheries, they have
dark centers, and gradually become lighter toward the edges where
they blend with the unstained portions of the nuclei. Reticular
threads do not stain at all, the reticular granules only occasionally;
and, though the chromosomes stain less deeply, they still show a color
as dark as that of the nucleoli.

4. Copper-Chrome-Propionate Fixation. Reticular elements
and chromosomes are heavily stained. Nucleoli are unstained. The
cytoplasm is slightly colored. The effect of hydrolysis is to render
the nucleoli deeply staining, while the cytoplasm fails to show any
color (pi. 9, fig. 10). This technique gives a good reticular stain,
in which respect it resembles the Feulgen reaction.

5. Nickel-Chrome-Propionate Fixation. The nucleoli are the
only cell parts to stain. Treatment with hydrochloric acid lessens
the staining capacity of the nucleoli, so that some stain lightly and

some not at all.

6. Formol-Sulfuric Acid Fixation. However rapidly these sec-
tions are passed through the alcohols and clove oil into xylene, none
of the structures is sufficiently mordanted to retain any of the
stain. Following hydrolysis a very small percentage of the nucleoli
show a faint color.

7. Formic Acid-Acetaldehyde Fixation. The cell structures are
so little mordanted by this fixative that even the most rapid dehydra-
tion leaves only a trace of color in the chromosomes. The reticular
granules of the outer cell layers retain a slight amount of color.
Hydrolysis does not affect the staining of the cell structures.

8. Modified Erlicki Fixation. All the nucleoli are well stained,
but the chromatin shows no color. Hydrochloric acid treatment
renders the chromosomes deeply staining and the nucleoli only
erratically stainable. The center of each stained nucleolus is darker
than the periphery, some being stained only in the center, in contrast
to the light centers and dark margins of the unhydrolyzed tissue.

9. Chromic Sulfate-Formaldehyde Fixation. In neither un-
hydrolyzed nor hydrolyzed tissue are any of the cell structures con-
sistently stained. The nucleoli, especially those of the outer cell
layers, often show a faint color.

D. Safraniri'iodine stain

This stain reacts with the tissues after each fixation almost ex-
actly as does the crystal violet-iodine stain. It requires, however,
a stronger solution and longer time of staining, but differentiation
takes place in about the same length of time that it does with the
crystal violet-iodine. See pi. 9, figs. 11 and 12.

1936

Effects of several fixatives on staining reactions in Zea mays

113

Discussion

The Feulgen reaction, unlike the other stains, colors only the
chromatin, regardless of the fixative employed. This is doubtless
due to the chemical nature of the method ; for f uchsin-sulf urous acid
(Schiff's reagent) is a standard chemical test for detecting the
presence of aldehydes. Feulgen and Rossenbeck ('24) showed that
a partial hydrolysis of the hexose-containing thymonucleic acid frees
an aldehyde that will react with fuchsin-sulfurous acid to give the
deep violet color characteristic of the Schiff test. They demonstrated
further that pentose-containing nucleic acids do not react positively
to this test after the same hydrolyzing treatment, because of the
greater difficulty of effecting hydrolysis. What is of especial interest
to cytologists, however, is the adaptation by these investigators of
the Schiff test to staining fixed and sectioned cells. This method
is a valuable asset to the cytologist, in that it enables him to compare
a specific chemical color test with the ordinary histological stains,
i.e., iron alum-hematoxylin and the aniline dyes.

The time of hydrolysis with N HCl at 60°C. originally recom-
mended by Feulgen and Rossenbeck was 3-5 minutes. They expressed
the opinion that "das Optimum der Intensitat des hydrolytischen
Eingriffs bei alien Praparaten— gleichgultig, ob sie dem Tier- oder
dem Pflanzenreiche entstammen— gleich ist". Feulgen-Brauns ('24),
who investigated the optimum times for different temperatures, gives
4 minutes for 60°C. This optimum time was confirmed by Bauer
for the corrosive sublimate-acetic acid fixation employed by the above
authors, but he found that it was different for other fixatives. Boas
and Biechele ('32), working with plant tissues that were not killed
before hydrolysis whether sectioned or unsectioned, used a time of
2-3 minutes; for 4 minutes caused distortion of the tissues. They
found that some species react well and others not at all. As they
suggest, the latter might give a reaction with a different time of
hydrolysis. All this indicates that the optimum time of hydrolysis
varies with the type of fixation and the species of organism fixed.

Sax ('31) recommended the use of Newton's modification of the
Gram stain following fixation with Navashin in order to obtain a good
chromosome stain; this method proves to be equally specific for the
chromatin of the "resting" cell. That the selectivity of this reaction
is entirely conditioned by the fixative, however, is evidenced by the
effects of other fixing reagents on this staining reaction.

It is an interesting fact from the point of view of stain specificity
that the crystal violet and the safranin stains, when used separately,
exhibit an identical selectivity for the different cell parts. This

Cytologia 7. 1926

8

114

J. G. Carlson

Cytologia 7

would indicate that the differential staining effects attaching to these
stains when they are used together in the Flemming tri-color stain
are not due wholly to a peculiar affinity of each for a particular cell
part. The explanation of this possibly lies in the effect of one of
these stains on the other. Whether this influence is exerted through
the one replacing the other or simply through the color of one
obscuring that of the other is not clear. At least this points to the
danger of attaching very much significance to stain "specificities"
that are not known to be chemical reactions.

It is customary to distinguish between two chemically different
kinds of material in the ''resting" nucleus: chromatin, or basichro-
matin, which forms the reticulum, and plastin, or nucleolar sub-
stance. A careful study of the tissues fixed in nickel-chrome-
propionate indicates that the reticulum contains not only chromatin,
but also some substance with the staining properties of plastin. This
occurs in the form of scattered granules. Therefore, the globules to
which I have referred as reticular granules are of two kinds. An
examination of material fixed in nickel-chrome-propionate always
gives one the impression that the chromatin has been partially dis-
solved during fixation, and so altered in form to a greater or less
degree, and not that it is simply unmordanted. The fact that after
the Feulgen reaction the reticulum threads appear wide and fuzzy
instead of narrow and sharply defined and that the chromosomes are
stained in outline only, lends support to the correctness of this inter-
pretation. In this Feulgen-stained material there is no evidence of
the presence of reticular granules in the intermitotic nuclei, a condi-
tion to be expected if one supposes that they have been rendered
sufficiently fluid by the solvent action of the fixative to have lost
their normal form and become indistinguishable from the already
diffuse and partly dissolved reticular threads. Further, a large
granule is often seen lying in the reticulum near its point of juncture
with the nucleolus ; the hematoxylin stain shows it as a sharply defin-
ed, jet-black mass, while the Feulgen stain does not render it visible
at all. It must be largely, if not wholly, plastin or a plastin-like
substance.

Zirkle has pointed out that copper-containing fixatives render
the nucleoli stainable only if the pH at which fixation occurs is above
4.0. That this is a very delicate reaction is evidenced by the fact that
a fixative only slightly more acid fails to mordant the nucleoli. Yet,
strangely enough, this delicately adjusted fixation effect seems to
be an irreversible one; for if material fixed in this way is treated
with N HCl at 60°C. for 20 minutes, there is very little visible effect
on the stainability of the nucleoli. The same seems to be true of the

t

1936

Effects of several fixatives on staining reactions in Zea mays

115

mitochondria, only in the case of these it is probably a question of
preservation or dissolution rather than one of mordanting. With
the exception of the osmic acid-containing fixatives, those of pH
below 4.6-4.8 dissolve out the mitochondria ; after they are preserved,
however, the hydrochloric acid treatment has little or no effect. It
would seem, therefore, that once a cell structure is preserved and
rendered stainable by the fixative, subsequent treatment with solu-
tions of a pH far below that at which preservation and mordanting
would not have occurred during the fixation does not essentially alter
the original fixation image. Any theory that attempts to explain
fixation must be able to account for such a condition as this.

Summary

1. The Feulgen reaction maintains its specificity after each of
the nine different fixatives used ; of the cell components it stains the
chromatin only.

2. The intensity of the staining reaction of the root tip cuticle
by the Feulgen technique usually varies with the fixation as does
that of the chromatin.

3. For each of the other stains employed (Heidenhain's iron
alum-hematoxylin, crystal violet-iodine, and safranin-iodine) the
stainabilities of the different cell structures are conditioned directly
by the fixative.

4. Crystal violet-iodine and safranin-iodine exhibit no consistent
differences in their selectivity for the various cell organs after any of
the fixatives.

5. After certain fixatives the hydrochloric acid treatment used
for the partial hydrolysis of the thymonucleic acid for the Feulgen
reaction (N HCl at 60°C. for 20 minutes) affects the relative stain-
ing capacities of plastin and chromatin, when the iron alum-hematoxy-
lin, crystal violet-iodine, and safranin-iodine stains are used.

6. Only the Bouin- and Navashin-fixed material is affected by
hydrochloric acid treatment in the reaction to iron alum-hematoxylin :
in both the chromatin loses some of its staining capacity; in the
latter the stainability of the nucleoli is increased as well.

7. The effects of treatment with hydrochloric acid on the stain-
ing reactions of crystal violet-iodine and safranin-iodine is quite
pronounced for several fixatives: chromatin stainability is increased
after modified Erlicki, decreased after Navashin and copper bichro-
mate, and lost after Bouin fixation ; nucleolar stainability is acquired
after copper-chrome-propionate, increased after Navashin, copper
bichromate, and chromic sulfate-formaldehyde, decreased after

116

J. G. Carlson

Cytologia 7

1936

Effects of several fixatives on staining reactions in Zea mays

117

h

modified Erlicki, and decreased or lost after nickel-chrome-propionate

fixation.

8. Copper-containing fixatives that render the nucleoli stainable
with crystal violet-iodine and safranin-iodine cause the nucleolar
peripheries to stain darker than the centers, while after hydro-
chloric acid treatment this reaction is reversed, the centers staining
darker than the peripheries.

9. In some preparations mitotic nucleoli differ from inter-
mitotic nucleoli in their reactions to stains, which would seem^ to
indicate that the nucleolus undergoes some chemical or physical
change at the initiation of mitosis.

10. Evidence is presented in support of the view that plastin,
as well as chromatin, occurs in the form of granules in the reticulum
of the inter mitotic cell.

11. Nucleoli, which are mordanted by copper-containing fixa-
tives only if the pH is above 4.0, are but little affected in regard
to their stainability by subsequent treatment with hydrochloric acid
of a much lower pH.

12. Mitochondria, which are usually dissolved out by the more
acid fixatives, are not affected by treatment with normal hydrochloric
acid after fixation.

I want to express my appreciation to Dr. Conway Zirkle for

suggesting this problem, as well as for his help and advice throughout

the course of the investigation.

Department of Botany,
University of Pennsylvania

Literature Cited

Bauer, H. 1932. Die Feulgensche NuklealfSrbung in ihrer Anwendung auf cytolo-

gische Untersuchungen. Zeitschr. Zellf. Mikr. Anat., vol. 15, pp. 225-247.
Boas, F. and O. Biechele. 1932. t)ber die Feulgensche Nuclealreaktion bei Pflanzen.

Biochem. Zeitschr., vol. 254, pp. 467-474.
Feulgen R. and H. Rossenbeck. 1924. Mikroskopisch-chemischer Nachweis einer

Nucleinsaure vom Typus der ThymonucleinsSure und die darauf beruhende

elektive Farbung von Zellkernen in mikroskopischen Praparaten. Zeitschr.

Physiol. Chem., vol. 135, pp. 203-248.
Feulgen-Brauns, F. 1924. Untersuchungen uber die Nucleal-Ffirbung. Arch. Gesamt.

Physiol., vol. 203, pp. 415-435.
Lenoir, M. 1922. Les nucleoles pendant la prophase de la cinese II du sac embryon-

naire du Fritillaria imperialis L. Comp. Rend. Acad. Sci., vol. 175, pp.

985-987.
Margolena, L. A. 1932. Feulgen's reaction and some of its applications for botanical

material. Stain Tech., vol. 7, pp. 9-16.
Martens, P. 1925. Le cycle du chromosome somatique dans les Phanerogames. II.

Listera ovata. La Cellule, vol. 36, pp. 127-214.
Newton, W. C. F. 1926. Chromosome studies in Tulipa and some related genera.

Jour. Linn. Soc. (Bot.), vol. 47, pp. 339-354.

I

Sax, K. 1931. The smear technic in plant cytology. Stain Tech., vol. 6, pp. 117-122.
Van Camp, G. M. 1924. Le role du nucleole dans la caryocinese somatique (Clivia

miniata Reg.). La Cellule, vol. 34, pp. 7-49.
Zirkle, C. 1928. Nucleolus in root tip mitosis in Zea Mays. Bot. Gazette, vol.86,

pp. 402-418.

— 1928. The effect of hydrogen ion concentration upon the fixation image of various

salts of chromium. Protoplasma, vol. 4, pp. 201-227.

— 1929. Fixation images with chromates and acetates. Protoplasma, vol. 5, pp.

511-534.

— 1931. Nucleoli of the root tip and cambium of Pinus strobus. Cytologia, vol. 2,

pp. 85-105.

— 1933. Cytological fixation with the lower fatty acids, their compounds and deriva-

tives. Protoplama, vol. 18, pp. 90-111.

Explanation of Plates

Photographs were made at a magnification of 980 diameters.

Fig. 1.

Fig. 2.
Fig. 3.
Fig. 4.

Fig. 5.
Fig. 6.

Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.

Fig. 11.
Fig. 12.

Plate 8

Navashin fixation. Iron alum-hematoxylin stain preceded by hydrochloric

acid.

Modified Erlicki fixation. Feulgen reaction stain.
Chromic sulfate -formaldehyde fixation. Iron alum-hematoxylin stain.
Modified Erlicki fixation. Iron alum-hematoxylin stain preceded by hydro-
chloric acid.

Formic acid-acetaldehyde fixation. Iron alum-hematoxylin stain.
Copper-chrome-propionate fixation. Iron alum-hematoxylin stain preceded
by hydrochloric acid.

Plate 9

Bouin fixation. Crystal violet-iodine stain.

Navashin fixation. Crystal violet-iodine stain.

Navashin fixation. Crystal violet-iodine stain preceded by hydrochloric acid.

Copper-chrome-propionate fixation. Crystal violet-iodine preceded by

hydrochloric acid.

Copper bichromate fixation. Safranin-iodine stain.

Nickel-chrome-propionate fixation. Safranin-iodine stain.

Cytologia 7, 1936

Plate 8

^

i

Carlson : Effects of Several Fixatives on Staining Reactions in Zea mays.
Especially with Reference to the Feulgen Reaction

=1

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Cytologia 7, 1936

Plate 8

I

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Carlson: Effects of Several Fixatives on Staining Reactions in Z(<f }imys.
Especially with Reference to the Feuigen Reaction

INTENTIONAL SECOND EXPOSURE

Cytologia 7, 1936

Plate 9

\^

Carlson: Effects of Several Fixatives on Staining Reactions in Zea mays,
Especially with Reference to the Feulgen Reaction

Cytologia 7, 1936

Plate 9

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Carlson : Effects of Several Fixatives on Staining Reactions in Zea mui/s,
Especially with Reference to the Feulgt-n Reaction

INTENTIONAL SECOND EXPOSURE

Rrprintod from the American Jouhnal of Botany, Vol. 23, No. 7, 483-489, July. 193a

Printed in l'. S. A.

,t

'41

CYTOLOGICAL STUDIES OF TOXICITY IN MERISTEM CELLS OF ROOTS OF
ZEA MAYS. I. THE EFFECTS OF THE NEUTRAL SALTS ^

John K. Edwards

SiNCK THK classical researches of Ilinger (1SS4) on
the influence of saline media on certain fishes, much
work has been done on the biological efTects of varia-
tions in the salt environment of plants and animals.
Investigators in both i)lant an.l animal physiology
early noted the necessity of a balanced salt mechuni
for the normal functioning of living organisms. Ex-
cesses or deficiencies resulted in a retardation of
growth and develoi)ment, or, in some instances, death.
Plant i)hysiol()gists have generally explained the re-
sults of salt deficiencies as starvation. While this
e.\-i)la nation may be correct for hmg-time stiidios where
certain elements are deficient but not entirely al)sent,
the results of short-time studies are i)robably best
exjjlained as due to the toxicity of the remaining
elements (True, lll.-JO; Bamford, 1931).

The studies of True and his associates (18%, 1000,
1930) on the toxicity of single-salt solutions indicate
four general classes of inorganic ])lant poisons: the
acids and arid salts, salts of the alkali metals and
earths, compounds of the heavy metals, and salts with
toxic anions. Solutions of the acids and acid salts
are relatively toxic, dej)ending on the hvdrogen-ion
concentration. The neutral salts of the alkali "metals
and earths, weakly toxic in most cases, varv in their
toxicity as the cation is changed. With the heavy
metals, all extremely toxic, the metallic ion is the
cause of injur>-. The connnon anions most toxic to
plant protoplasm are those of the fluorides, bromides,
iodides, and cyanides.

The neutral salts of potassimn, sodium, magnesium,
and calcium are of esjiecial interest to the physiolo-
gist. All form a iiart of the normal environment of

^ Recoivod for imblicMtjon April 10. 1936.

\ wish to ('xpicss my .ippicciation to IVof. Hodiicy H.
Tnir. under whose diivctjon this iii\ ('slinatioii ' w.is
carrifMl {)ut. for his valuable stigjr,,sti,,ns and intcMcsl in
iiie (.n/oiein. i also wish to thank Prof. Conway Zirkle
and Dr. K. D. Douk for assistance in the preparation of
the manuscript.

plants and animals. >;odium is apparently the only
one of the grouj) not neces.sary in plant "^ nutrition;
yet Its close chemical affinity to jwtassium and its
universal presence in 'lature have caused it to be in-
cluded in most studies of this .'^eries. The majority
of papers reporting the various biological efTects caused
by these salts exi)ress results largely in terms of
gross mori)h()l()gy or as visible i)hysiological reactions
in living material. Conparatively little has been done
on the cytological reactions of the cells and tissues.
The present investiga^on was made in an effort to
determine the cytologi:-al changes in the root-tip cells
of 7.ca Mays followi ig toxic treatments with the
chlorides of iwtassium, .^odium, magnesium, and cal-
cium.

Various plant parts have been used in studying the
efTects of the neutral salts on protoplasm. Addoms
(1927) immer.«^ed root hairs of wheat in 0.1 N single-
salt solutions, and observed the effect on the proto-
plasm under dark-field illumination. The salts of
potassium, sodium, calcium, magnesium, zinc, and
aluminium all caused coagulation. Kerr (1933) noted
that SO iier cent of the root hairs of Limtwbium burst
after a few minutes in 0.02 M XaCl, the remaining
20 i)er cent dying witinn 30 minutes. Similar re.^nlts
with the salts of potassium have been reported by
Cholodnyj (1923). Kahho (1921), in a studv of the
efTects of neutral s,,lt« on cells of Trndesrniitin and
red cabbage leaves, found the following order for the
chlorides, beginning the series with the most toxic
K > XIT^ > Xa > Sr > Mg > Ra > Ca. The inabill
ity of the cells to plasmolyze in a strong sugar solution
after the salt treatment was used as an indication of
the coagulation anrl d-'ath of the protoplasm. Iljm
(1935), in a similar study, investigated the killing of
])lant cell-^ by immersing ei>idermal tis.«<ue in various
conr, ii^.tuuii^ UL KCl, XaCi, and CaCI... In the
non-toxic concentrations the cytoplasm and nucleus
were without visible structure, or, at the most, finely

484

AMERICAN JOURNAL OF BOTANY

[Vol. 23,

July. 1936]

EDWARDS — CYTOLOCJICAL STUDIES OF TOXICITY

485

granular. Stronger concentrations caused a granula-
tion and coagulation of the cell parts. The changes
in the cytoplasm were gradual, beginning in the region
of the nucleus and si)reading throughout the cell.
The tonoi)last was the last part to die, retaining its
semipermeable character in strong solutions long after
the other parts were dead. Seifriz (1928) found that
the salts of potassium, sodium, and calcium in 0.128 M
concentrations stoi)i)ed i)rotoi)lasmic streaming in
Elodea leaf cells after a few hours.

Injection experiments where the salt solutions are
placed in direct contact with the cytoplasm have
yielded interesting results. Skeen (1930) injected a
CaCK, solution (containing 20 p. p. m. Ca) into
Trianea {Limnohium) root hairs and noted that
cyclosis stopjied in five minutes, with death resulting
from coagulation in 10-15 minutes. Kerr (1983),
working with root hairs of Limnohium, found an
irreversible coagulation of the cytoplasm with CaCl.,
stronger than 0.04 M, whereas the coagulation pro-
duced by 0.1 M MgCl., was reversible. KCl and NaCl
in 1 M concentrations mixed freely with the proto-
plasm and stopped cyclosis for only five to ten
minutes.

Most cytological studies of cells showing chemical
injury have been made on material grown in unbal-
anced nutrient solutions. Reed (1907) found that
mitosis continued in root tips of Zea Mays in the
absence of calcium, but new cell walls were not
formed. Sorokin and Sommer (1929) made cyto-
logical studies on the roots of Pisum sativum grown
in nutrient solutions lacking calcium. They rejwrted
a material decrease in the amount of cytoplasm,
irregular mitotic as well as amitotic cell division, and
a failure of the embryonic cells to mature. In solu-
tions lacking both calcium and magnesium, the tissues
underwent a slower degeneration, but otherwise were
much the same as found whon calcimn alone was
lacking. This retardation of injury is explained by
the absence of microorganisms from the magnesiiun-
free solutions. Bamford (1981) observed the root
tips of wheat and corn grown in calcium-deficient
nutrients. Where calcium was entirely absent, growth
ceased almost immediately after the roots were placed
in the mediimi. Cell injury progressed from the
periphery of the root inward and was characterized
by a gradual " erosion " of the cytoplasm, followed
by degeneration and disappearance of the nucleus.
As wheat roots were normal after four days in dis-
tilled water, the pronounced injury at the end of two
days in the calcium-free solution was attributed to
the other constituents of the nutrient, especially mag-
nesium. Lutman (1984) grew various higher plants
in nutrient solutions in which either nitrogen, phos-
phorus, potassium, magnesiiun, or calcium was lack-
ing. Root-tip cells of secondary roots grown in these
solutions showed a premature vacuolization, inter-
preted by Lutman as indicating maturity and sen-
escence. The nucleus is reported smaller in relation
to cell size in the solutions deficient in nitrogen,
phosphorus or magnesium.

Yamaha (1927) has given a detailed account of
the etfeets of many different chemicals on the root-tip
cells of Vicia faba. The roots were placed in various
concentrations of the salts for a period of two hours,
followed by washing in running water and fixation
in Flemming's fixative. With the chlorides of potas-
sunn, sodium, and calcium in the higher concentra-
tions (2 M to 0.02 M), he reports coagulation of the
cytoplasm and certain mitotic anomalies.

Materials and methods. — Young seedlings of Zea
Mays L. (Golden Beauty) furnished the plant mater-
ial for the experimental work. When germinated in
moist sphagnum for four days at 19-22°C., about
40-50 per cent of the seedlings have primary roots
5-6 cm. long. As profuse laterals are not developed
until the sixth or seventh day, seedlings of such age
are very useful in short-time growth studies. The
experimental set-up was the same as that described
by Beall (1980), consisting of 500 cc. pyrex beakers
with perforated i)yrex Petri dishes supporting the
seedlings. Such a set-up insures immersion of the
entire radicle with the exception of the upper 10 nun.
The toxic solutions were prepared from 1 M stock
solutions of Merck's reagent CaCL, MgCL,, and KCl,
and Merck's c.p. XaCl. During the time of exposure
to the toxic solutions, the plants were kei)t in an
insulated dark room, with air and solution tempera-
tures varying from 19-22°C.

Preliminary growth studies were made with ten
concentrations of each salt, ranging from 0.1 ]\I to
0.01 M for KCl, NaCl, and MgCI.., and from 0.2 M
to 0.01 M for CaCl.,. The failure of the roots to
elongate in distilled water after 24 hours in the salt
solutions was used as a criterion of death, while addi-
tional growth in distilled water indicated survival and
recovery. On the basis of the i)reliminary studies,
the toxic treatment of material for cytological study
was limited to four concentrations of each salt, rang-
ing from strongly toxic through less toxic concentra-
tions to solutions where growth was only slightly less
than normal. The C^aCl, series was 0.2 M, o!l M,
0.088 M, and 0.01 M. For the potassium, sodiiun,
and magnesium salts, the concentrations used were
0.1 M, 0.088 M, 0.02 M, and 0.01 M. At the end of
the toxic treatments lasting 24 hours, the root tips
were fixed in each of four cytological fixatives. The
fixing fluids used were a chromic sulphate-formalde-
hyde mixture (Zirkle, 1932a), Zirkle's (1984) modifi-
cation of Erliki's solution, Bouin's, and Nawaschin's
fluid. The fixation time was 48 hours. Dehydration
in the butyl alcohol series was followed by embedding
in paraffin. The sections were cut eight micra in
thickness, and stained with iron-ahun haematoxylin.
No coimter stain was u.scd.

CeUs of the normal root tips of Zea Mays. — The
root tip of Zea Mails has the embryonic tissue organ-
ization typical of the majority of monocotyledons,
A distinct caly])trogen covers a single layer of initials
which immediately give rise to the ])lerome cylinder,
periblem, and dermatogen of the root. The cells near
the tip of seminal roots tend to be isodiametric,

T

r

•i»

I

1

I
4

measuring about 10-15 micra in size. The nuclei are
comi:)aratively small and spherical, normally occupy-
ing the center of the cell. Dividing cells are abundant
at all times, but are slightly more numerous during
the day (Karsten, 1915). Early prophase stages are
l)lentiful and show a well defined spireme. The
chromosomes are long and numerous and tend to be
fixed in clumps at metaphase and telophase. Al-
though the cells of the initial region are generally
re])orted to be non-vacuolate and rich in cytoplasm,
the work of Zirkle (1982a) shows the presence of
numerous spherical vacuoles, varying from four to
eight micra in diameter in the calyptrogen and
I)lerome, and somewhat smaller in the other tissues.
Although careful work on the nucleo-cytoplasmic ratio
has not been reported, it is quite likely that cells of
the i)rimary meristems resemble the cambium cells
observed by Bailey (1980), where the more rapidly
dividing cells, instead of being richer in cytoplasm,
are the more vacuolate.

Fixation in the normal meristem eells of Zea Mays.
— P>om his extensive work on fixation, Zirkle (1928,
1982a, 1982b, 1984) recognizes two definite types of
fixation image, the so-called " acid " and " basic "
images. The acid images are those produced by fix-
ing solutions containing acetic acid, and by chromic
acid and the bichromates on the acid side of pll
4.8-5.0. The basic images are set by formaldehyde
and by certain metallic salts on the basic side of
pH 4.8-5.0. The fluids pro(hicing the acid image are
the best preservatives of resting and dividing chro-
matin, nucleolar material, and spindle fibers. The
cvtoplasm, however, is fixed as a reticulum, or as
fairly coarse threads, and is referred to as " spongio-
j)lasm." All cytoplasmic detail is destroyed and the
resulting image, including the vacuoles, is wholly
artificial. The basic images are characterized by a
fixation of the cyto]ilasm as " hyaloplasm " — that is,
without any visible structure. Although the tono-
plast is dissolved, the vacuoles are preserved in their
true form. Mitochondria, the nucleolus, nuclear
lymph, and dividing chromatin are fixed, while rest-
ing chromatin is dissolved.

A critical study of the staining properties of the
root-tip cells following various acid and basic fixa-
tions has been made by Carlson (1936). He finds
that the basic fixations of chromic sulphate-formal-
dehyde and modified Erliki's solutions give similar
images and stain well with iron-alum haematoxylin.
He also finds that the acid fixations of Bouin's and
Xawaschin's fluids are much alike when followed by
the same staining technique.

Of the fixatives used in this study, two produce
the acid image and two the basic image. The chromic
sulphate-formaldehyde mixture preserves the cyto-
plasm much as it is seen in the living condition
(Zirkle, 1982a), and, except where otherwise indi-
cated, conclusions regarding the effects of the toxic
solutions on cvtoplasmic structure and vacuoles are
drawn from a study of material fixed in this medium.
The modified Erliki's fixative, although erratic at

times, fixes mitochondria better than the chromic
sulphate-formaldehyde solution, and is used to deter-
mine their presence. Injury to resting and dividing
nuclei is determined from material fixed in Bouin's
and Nawaschin's fixatives.

Results of cytological studies.— r//e eytoplasm
following basic fixation— The cytoplasm of root-tip
cells is markedly affected by strong concentrations
(0.1 M) of KCl and NaCl, but tends to assume a
normal appearance in the less toxic solutions. Basic
fixation following tleath of the roots in 0.1 M solu-
tions shows a tendency for the cytoi)lasm to pull
away from the cell walls and clump about the nucleus
(fig. 2, 3). The resulting vacuoles are large and dis-
torted, and probably represent a fusion of the small
spherical vacuoles of the normal cell. Potassium, in
the case of the 0.1 M solutions, produces a coarsely
granular cytoplasm, while sodium of the same strength
causes little change from the normal appearance of
the cytoplasm. The 0.083 M treatments produce an
image similar to that of the 0.1 M solutions, except
that true vacuoles, normal in size and appearance,
may be seen in quite a few cells. With the 0.02 m'
concentrations, a great number of large vacuoles
persist in an otherwise normal cytoplasm. All cells
ajipear normal after 24 hours in the 0.01 M solutions.
Although the sections were not destained especially
for observation of mitochondria, the latter are seen
in some cells of the root tips following all sodium
and potassium treatments.

MgCl, is very similar to KCl and NaCl in its
action on the cytoplasm (fig. 4). The 0.1 M solu-
tions cause a clum])ing around the nucleus, leaving
the fu.^ed vacuoles as large clear spaces. In most
cases the cytoplasm is fi.xed as a homogeneous mass
(hyaloplasm), but in some cells small dark-staining
globules are present. As normal nuclei are often
present in the same cells, no postulations are made
regarding the origin of this chromatin-like material.
The 0.083 M and 0.02 M concentrations cause much
the same type of fused vacuoles, comparatively few
being normal. Roots from the 0.01 M treatment
show cells with normal cytopla.sm and vacuoles.
Mitochondria were observed in all but the 0.1 M
series.

Although not so drastic in its killing action as
0.1 M KCl, the 0.2 M CaCl, affects the cytoplasm
in a similar manner. The cytoplasm is seen to contain
small evenly distributed granules, probably formed
by the salt solution prior to fixation. The vacuoles
tend to be large and distorted, and apparently repre-
sent a coalescence of the smaller normal vacuoles.
The non-killing concentrations produce a less gran-
ular cytoplasm and a higher percentage of normal
vacuoles as the dilutions become progres.sively greater
until a normal basic image is secured following the
0.01 M treatment. Mitochondria are present in all
cases.

Although the anfnofnni«tir ^fft^otv r%f f*nln'„,^^ <,», +i,^
injurious action of other salts were not thoroughly
investigated, the cytop asm and vacuoles appear nor-

IRREGULAR PAGINATION

July, 1936]

EDWARDS — CYTOLOGICAL STUDIES OF TOXICITY

Table 1. Summary of the effects of the neutral salts on various parts of the ccll.^

487

Fig. 1- 3. All figures represent und.fforontiatod colls from the region immodiafolv hohind tho initiil. The
magnification is 838 diameters. Fig 1-8 show tho basic fixation images of chromic suli.hafo-formal.l Vh Tlo follovnm'
the varioiK. indicated treanionts, while fig. 9-13 show tho acid images of liouin's fixative.-Fig. 1. (VI loni d

root tip grown in distilled water.— Fie. 2. 0.1 M Kr 'I —Fi.r q ni ai v.,oi i/... , nnn^\,\) J!, "^^""^jrmi'
M CaCL-Fig 6^ 0.033 M KCl.-Fig. 7. 0.02 M KCl.-Fig. 8. 0.033 'MScL-FigrorT^ll^lrimmli^a/n^o^ Hp
grown in distilled w^ter and fixed in Bouins.-Fig. 10. 0.1 M KCl.-Fig. 11. 0.1 M NaCl.-Fig. 12. 0.033 M KCl.

s

j

-' *

KCl

XaCl

MgCk.

CaCl2

M

M

M

M

M

M

M

M

M

M

M

M

M

M

M

M

Cell parts

10
CG

30
CG

50

100

10

30
CS

50

100

10

30

50

CS

100
X

5
CG

10
CG

30

100

Cytoplasm

X

N

CS

X

X

CS

CG

X

Vacuoles

D

D

DX

X

D

D

DX

X

D

1)

DX

X

D

DX

DX

X

Chromatin

CG

GX

X

X

CG

GX

N

N

CG

GX

X

X

CG

CG

N

N

Prophases

F

X

N

N

F

N

N

N

F

N

N

N

F

X

N

N

Metaphases

F

X

N

N

A

X

F

N

A

X

X

X

A

X

N

N

Anar)hases

F

X

N

N

A

N

F

X

A

X

X

X

A

X

N

N

Teloi)hases

F

X

X

N

A

X

F

N

A

X

X

X

A

X

N

N

Spindles

A

X

X

N

A

N

F

X

A

X

X

N

A

N

N

N

Cell i^lates

N

X

X

N

N

N

N

N

A

N

X

N

A

X

N

N

Nucleoli

N

N

N

N

N

N

N

N

X

X

N

X

X

N

N

N

Mitochondria

P

P

P

P

P

P

P

P

A

p

P

P

P

P

P

P

•> X, normal;

D, distorted;

C, clumped

; G, granular;

A, al

xsent; P, prese

nt; F

\ few

: S.

smooth.

mal in cells of roots grown in a solution containing
0.2 M KCl and 0.01 M CaCl,.

The cytoplasvi joUounnq acid fixation. — Root tips
grown in distilled water for 24 hours and fixed in
liouin's fluid show the typical acid image, with a
granular, evenly distributed cytoplasm (fig. 9). The
0.1 M solutions of KCl, NaCl, and ]\IgCl„ however,
produce images very similar to those described by
Sorokin and Sonimer (1929) and Iw Bamford (1931)
for calcium-starved roots. A majority of the cells
are practically devoid of cytoplasm, while the re-
mainder show large clear spaces, not typical of true
vacuoles (fig. 10, 11). In striking contrast to this
condition is the image obtained following treatment
with 0.2 M CaCl._, (fig. 13). Here the cytopla.sm is
granular and evenly di.^tributed throughout the cells,
much as the normal condition following acid fixation.
The weaker concentrations of the various salts do not
noticea})ly affect the acid fixation of cytoplasm.

Resting and dividing nuclei following acid fixation. —
The acid fixatives show well preserved nuclei in the
cells of roots grown in distilled water. The nuclear
lymi)h is dissolved, leaving a clear halo al)out the
well-fixed nucleoli. The resting chromatin is fixed as
granules of various sizes, generally in the peripheral
region of the nucleus. Division figures are abundant
and show normal spindle fibers and cell plates.

Only the more concentrated salt solutions cause
notice.ible injury to the resting nucleus (fig. 10, 11).
Following exjjosure to these, some nuclei are angular
or elongated, while others retain their normal spher-
ical shape. All salts react alike in causing a clumi)ing
of the chromatin, either as large granules or as dark-
staining amorphous masses around the ])eriphery of
the nucleus. P'ollowing immersion in 0.1 M KCl this
chromatic material appears to occupy most of the
space left by the dissolved nuclear lymph, leaving a
narrow, irregular halo around the nucleolus. Except
for a slightly more granular chromatin, the resting
nnolei nre nrnotif.'ilh- nr»rTTi,'il nffpr '^4 hnurs; in tho

0.0;« M solutions (fig. 12), while the 0.02 M and
0.01 i\I concentrations produce no noticeable changes.

The nucleoli appear normal in all cases, retaining their
staining cjualities and sjiherical shape.

The effects of the salt solutions on dividing cells
point to two tyjies of toxic action. Potassium and
.^odium kill root tijjs in 0.02 M concentrations, yet
allow normal mitosis in 0.033 M solutions. Calcium
and magnesium, ho\\ever, show a total ab.sence of
division figures in all roots subjected to killing con-
centrations of the salts. Of the two alkali metal salts,
KCl seems to have the least effect on mitosis. As a
.^tudy of table 1 indicates, even the 0.1 M KCl series
shows a few mitotic iigures, and all other tested con-
centrations allow normal cell divisions. Sodium re-
acts in a slightly different manner, as mitotic figures
are entirely ab.sent from the cells of roots subjected
to the 0.1 M concentrations, while the 0.02 M series
shows very few dividing cells. The effect of the 0.033
M treatment, however, is similar to that of potas-
sium in that normal mito.sis takes place. This .^scarcity
of dividing cells following exposure to the 0.02 M
XaCl was con.«!istent in all ol).served material, consist-
ing of four root tips fixed in Bouin's fluid, four in
Xawaschin's, two in the chromic sulphate-formal-
dehyde mixture, and two in modified Erliki's. With
the exception of the 0.02 M XaCl, the three less-
concentrated .solutions of each salt produce an image
very similar to that .shown for 0.033 M KCl (fig. 12).
The tendency for meta phase chromo.somes to clump
is typical for Bouin's fixative.

Resting and dividing nuclei joUounnq basic fixation.
— While the basic li.xntion image is not so useful for
nuclear study, all i)arrs except the resting chromatin
are preserved, making additional observations on the
elTects of the salt treatments on nuclei i)ossible. The
nucleolus, a.side from deviations from the spherical
shape, is practically the same in all cases. Xone of
the .salts destroyed the nuclear lymph, and the halo
around the nucleolus so typical of acid fixation is
missing. Divisi(m fig-ires, while not .so abundant as

with the n^'i^l f'lV'»fw»n.' -wnr^ /-jK^-or^-o^J iti anmo /ioc-r»c

following the weaker ^alt treatments. However, any
scarcity or absence o. dividing cells following basic

July, 1936]

iti.lls. Til.'
lOllowiinr

Fijr. 1-13. All |i-nn> icpioenr uiKliffcvnlial*-.! rr\\< fnmi tlio rojri.m i.unip.h;,(,.|v l.phin.l flu- in
.na,n.h.anon h S.Hduuu-U... Fi^ 1-S show fh. l.asi,- Hxatinn una^.s of Hnon.u- snlphatr-lonnaMrhvo,. .onoun..
IIh> xanon> ,n.lHa.r,l tn-a .....nK. vhil.. fiu. 9 13 >how ih,. an.l inmiros of Houin s fixalivr.-Fi- 1 (VIN" from nonu .1
root tip mcmn ,n<lisf,ll,.,| ^vaf..l•.-FiL^2. 0.1 M Kd.-Fiir. 3. 0.1 M \.a(1 -F... 1 (H..!:! M \l.('1 i'!... . no
M ra( l..-h,^^(l U.U33 M K^;i--1':>«- J- 0.02 M K(M.-Fi^. 8. 0.033 M CaCL. -F.u. 9. (VIls f.onr norn.aM-oo( lip
?'"''';V"n''V ;',''■;■ ^'"'' ^'^^'^ "* Jiouin ...-F,g. 10. 0.1 M KCl.-Fig. 11. 0.1 M XaCi.-Fi-. 12. 0.033 M KCl

it

o

■ ♦»

•/.►

1

EDWARDS — CVTOLOCIC AL STL DIKS OF TOXICITY
T.AHi.K 1. Siittntiiiii/ of tin ( IJi els of (Jic fu idral >i<ilts on rarioiis jxirls of IIk c( ll.'i

4S7

Cell part.s

Cytoplasm

V-ic'U()l(\s

Chromatin

Pro])ha.'^(\s

Mctaphascs

Anajihascs

Tt'loplia.'-ics

Spindles

Coil plates

Nucleoli

Mitochondri;

M

10

('(;

D

CG

F

F

F

F

A

N

X

P

KC'l

M M

30 50

(x; X

1) D.V

ex X

X X

X
X
X
X
X
X

p

X
X
X
X
X
X

p

M_

100

X
X
X
X
X
X
X
X
X
X
V

XaCl
M M M M

M M M M

C\iCL.

M M M M

10 30 50 100

10 30 50 100

10 30 100

c\s

D

C(\

V

A

A

A

A

X

N

P

1)

(;x

X
X
X
X
X
X
N
P

X

J)X

X

X

F

F

F

F
X
X

P

X
X
X
X
X
X
X
X
X
X

p

cs

1)

('(;

V
A
A
A
A
A
X
A

CS

I)
ex

X
X
X
X
X
X
X

p

cs

1)X
X

\

X
X
X
V
X
X

X
X
X
X
X
X
X
X
X
X

p

" X. normal; I), (li.<lort(>(l; C. clumped; (1. <rranidar; A. absent ; P. present; F. ivw; S.

^moo

CC

1)

CC

]''

A

A

A

A

A

X

P

th.

CC

1)X

CC

X

X

X

X

X

X

X

p

CC

nx

X
X
X
X
X
X
X
N
V

X
X
X
X

N
X
X

N
N
N
P

JIllU.'j)

inal in rolls of roots orown in a solution containinjij
0.2 M KCl and 0.01 M CaCl,.

The ('utopldsfti following acid fixnt'tou. — Hoot ti|)s
Krown in distilled water for 24 hours and fixed in
Houin's fluid show tlu» tyi)ieal acid iinajje, with a
granular, evenly distributed f'vtoi)lasni (tij;. 0). The
0.1 M solutions of KCl, Xa(i, and Mud.,, however,
])rodiK'e imajies very similar to tho.se de.scrihed l)y
Sorokin and Sonuner (PI2!») and by Bainford (P>31)
for calcium-starved roots. A nia.iority of tlie cells
are ])racticall> devoid of cyt()i)lasni, while tlie re-
mainder show larjie clear s])aces, not ty])ical of true
vacuoles (lis;. 10, 11). In strikin": contra.st to this
condition is the ima'ze obtained followini: treatment
with 0.2 M CaCL. (fi<r. i;;). Here the cyto])lasm is
granular and evenly distributed throughout the cells,
nnich as the normal condition following acid fixation.
The weaker concentrations of the various salts do not
noticeably afVect the acid fixation of cytoplasm.

Uvstnuj and dividiiu} inirlci followincj acid fixat'inv. —
The acid rixati\es show well preserved nuclei in the
cells of roots grown in distilled water. The nuclear
l>nii)h is dissolved, leaving a clear halo about the
well-fixed micleoli. The resting chromatin is fixed as
granules of various sizes, generally in the i)eripheral
region of the nucleus. Division figures are abundant
and show normal sj)indle fibers and cell i)late.s.

Only the more concentrated salt solutions cause
noticeable in.jtiry to the resting nticletis (fig, 10, 11).
Following exposure to the.se, some nuclei are angular
or elongated, while others ret.iin their normal s|)her-
ical shape. All salts react alike in catising a clumping
of the chromatin, either as large granules or as clark-
staining amori)hous masses around the periphery of
the nucleus. Following immersion in O.I M KCl this
chromatic material api)ears to occui)y most of the
space left by the dis-ohcd nuclear lym|)h, leaving a
narrow, irregular halo arotmd the nucleolus. FAcejit
for a slightl>' more granular chromatin, the resting
nuclei are jiracticallv normal after 24 hours in the
0.03.S M >()luti(.ns (fig. 12), while the 0.02 M and
0.01 M concentrations jiroduce no noticeable changes.

The micleoli ajjpear normal in all cases, retaining their
staining (|ualities and s])herical shape.

Tiie effects of the salt solutions on (hviding cells
jjoint to two types of toxic action. Potassium and
sodium kill root tips in 0.02 M concentrations, yet
allow normal mitosis in 0.0;>3 M solutions. Calcimn
and magnesiunt, ho\M'\-er, show a total absence of
division figures in al! roots .subjected to killing con-
centrations of the .salts. Of the two alkali metal salts,
KCl seems to have the least etTect on mitosis. As a
study of table I indicates, even the 0.1 M KCl .series
.shows a few mitotic figures, and all other tested con-
centrations allow normal cell divisions. Sodiiim re-
acts in a slightly di!T"rent maimer, as mitotic figures
are entirely ab.<ent fiom the cells of roots subjected
to the 0.1 M concentrations, while the 0.02 M ,<eries
shows very few dividing cells. The etTect of the 0.033
-M treatment, however, is similar to that of potas-
sium in that normal mitosis takes ])lace. This scarcity
of dividing cells following exposure to the 0.02 M
XaCl was consistent in all observed material, consist-
ing of four root tips fixed in Houin's fluid, four in
Xawa.schin's, two in the chronnc sulphate-form.d-
«leh>(le mixture, and two in modified Krliki's. With
the excei)tion of the 0.02 M XaCl. the three le.ss-
concentrated solutions of each salt ])roduce an image
very similar to that shown for 0.033 M KCl (fig. 12).
Tlie tendency for metapha.-e chromosomes to clump
is typical for Bouin's fixative.

Jh'stiufi and dividing nuclei joUawinq basic fixation.
-Wjnle the basic fixation image is not so useful for
nuclear study, all parrs except the resting chromatin
are j)re,served, makin» additional observations on the
effects of the salt treatments on miclei possible. The
micleolus, aside from deviations from the sjjherical
-hape, is practically the same iti all cases. Xone of
the .salts destroye<l tie nucle.ir lymjih, and the halo
around the micleolus so tyjiical of a(Md fixation is
missing. Division fig'ir<»s, while not .so abundant as
with the acid fixation^, were observed in some ca.<^s
following the weaker alt treatments. However, any
scarcity or ab.-ence o tiividing cells following basic

INTENTIONAL SECOND EXPOSURE

488

AMERICAN JOURNAL OF BOTANY

[Vol.23,

fixation may often bo due to slow penetration of the
fixative or to its dilution by substances already
l)resent in the cells.

Hoots grown in 0.2 ]\1 KCl antagonized with 0.01
M CaCL, showed an abinidance of mitotic figures fol-
lowing fixation in chromic sulphate-formaldehyde.

Discission. — The results reported in the present
l)aper, while not comi)arable to the cellular degen-
eration in roots after many days in calcium-free
nutrients, may i)roperly be compared to the results
re])orted at the end of one or two days in such media.
Many workers have rei)orted that calcium in small
amounts antagonizes all ordinary constituents of
])lant-nutrient solutions, which taken alone or in
mixetl solutions are toxic to i)lant i)rotoi)lasm. Since
root-tij) cells of Zea Mays are normal after one day,
and wheat after four days in distilled water (Bam-
ford, 19ol), it is evident that the seed furnishes
sufficient calcium to supi)ly the newly formed proto-
plasm and, at the same time, prevent marked injury
by any other salts made soluble by the processes of
germination and growth. With young seedlings it is
only when other salts are introduced into the ex-
ternal medium that additional c:ilcium from an out-
side source becomes necessary for the i)reservation of
the cells in a normal living condition. Consequently,
any exi)lanation of injury to eel's of seedlings grown
in calcium-deficient nutrients, when control plants
grown in distilled water are still normal, should take
into consideration the toxic effects of the remaining
salts. Some idea of the i^otential toxicity of a nutri-
ent solution lacking calcium may be gained by cal-
culating the total i^otassium and magnesium in the
solutions used by Sorokin and Sommer (1929). Their
solution without calcium contained a total of 0.57 gm.
of potassium and 0.049 gm. of magnesium per liter,
as well as traces of sodium, manganese, aluminium,
copper, iodine, and fluorine, all harmful to plants in
the absence of calcium. By way of comparison, the
amount of potassium in the weakest killing concen-
tration of KCT is 0.7S gm. per liter.

The cytological reactions ])ro(hiced in root-tips of
Zea Mai/s by single-salt solutions of KCl, NaCl,
MgCr., and CaCI.^ have been ilescribed. A comjiar-
ison of the effects of the salts (m the cytoplasm, as
shown by acid and basic fixations, is of interest,
especially since all jirevious workers have drawn their
conclusions from a study of material fixed in acid
fixatives. Sorokin and Sommer (1929) report a de-
crease in the amount of cytojilasm and the appear-
ance of large vacuoles as one of the first visible
effects of calcium starvation on roots of P/.s?/m sati-
vum. Tn a similar study with corn and wheat, Bam-
ford (1931) also records an early disajipearance of
the cytojilasm. He emphasizes, however, the toxicity
of the remaining elements of the culture solution,
while Sorokin and Sommer explain all injury strictly
in terms of calcium absence. They conclude that
calcium is necessary to the protojilasm. either as a
chemical constituent or because of its physical effects
on the colloidal system of the cell. Lutman (1934)

also reports less cytoplasm and increased vacuoliza-
tion when i)lants are starved for certain elements,
e;i^,ecially calcium, magnesium, and potassium, and
concludes that such cells have reached early maturity
and senescence due to starvation.

Although basic fixation shows that the cytoplasm
is definitely injured at the end of one day in 0.1 AI
solutions of KCl, NaCl, and ]MgCl., the injury is
evident only as clumping and increased granulation.
The many small vacuoles typical of normal root-tip
cells are fused into a few larger ones. No treatment
causes a definite loss of cytojilasm. On the other
hand, when roots from the same salt treatments are
fixed in an acid fixative, the results are strikingly
different. The cytojilasm, fixed in the normal cell as
a non-vacuolate sjiongioplasm, reacts with the salts
in such a way that it is either dissolved or is badly
shrunken by the fixing solution. The presence of
CaCL, however, even as a 0.2 M single-salt solution,
jiredisjioses the cytoplasm to the usual acid fixation
with the acid fixatives. It will also be recalled that
0.01 ]\I CaCL, is sufiicient to jirevent injury to the
cytojilasm, as is revealed by basic fixation. It would
seem, then, that injury to the cytojilasm is best inter-
jireted from material fixed in basic fixatives. Just
as the long-accej)te(l idea that meristem cells are
non-vacuolate has been based on observations of the
artificial cytojilasm of the acid fixation image, so the
apj)arent early loss of cytojilasm from cells subjected
to toxic treatments, or calcium starvation, is based
on an artifact of acid fixation.

Most investigators who have observed the effects
of the neutral salts on j)lant cells, whether as single-
salt solutions or as calcium-free nutrients, agree that
the nucleus is the last jiart of the cell to show signs
of injury. Reed (1907) studied the cells of several
tyjies of jilants after calcium starvation and found
no modifications in the form of the nucleus, or in the
amount, form, or arrangement of the chromosomes,
lie did, however, record a marked absence of cell
jilates in dividing cells of Splwgi/ra and Zea Mays.
Sorokin and Sommer (1929) rejiorted a disturbance
of the mitotic axes in dividing cells of Pisum sativum
grown from one to two days in a calcium-free nutri-
ent, while Bamford (193i) found that the nuclei
remain normal in wheat roots during the first few-
days in solutions without calcium, nuclear degenera-
tion being one of the late stages of cellular injury.
With the excejition of slight differences in the size
of the nuclei, Lutman (1934) found no fundamental
variations from the normal condition when unbal-
anced nutrients were used. Yamaha (1927) subjected
roots of Vicia jaha to salt solutions of various con-
centrations (2 .M to 0.000002 M) for a two-hour
jieriod. The stronger solutons kill all cells and leave
no definite structure; the weaker solutions, while not
killing in their action, are reported to cause various
tyjies of mitotic anomalies. Following 0.02 M KCl
treatment, there is a thickeninsr and shortening of tbp
chromosomes, a degeneration of the spindle fibers,
and an absence of cell plates in some cells. CaCl.,'

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July, 19361

EDWARDS — CYTOLOGICAL STUDIES OF TOXICITY

489

I

<«l»

o

i:

and NaCl cause reactions similar to those produced
by the KCl. Nothing resembling these mitotic aberra-
tions is found in roots of Zea Mays after 24 hours
in the salt solutions. The stronger concentrations of
the salts tend to jirevent cell division, while the
weaker solutions allow tyjiical mitosis. If the divid-
ing cells of Zea Mays are as definitely abnormal at
the end of two hours in the salt solutions as Yamaha
rej)orts for Vicia jaha, they are able to adapt them-
selves to such an extent that only normal mitosis is
taking jilace at the end of 24 hours. The granulation
and clumping of resting chromatin rejiorted in this
investigation is ujiheld by Iljin's (1935) observations
of the coagulation and dumjiing of the nucleus in
living material when subjected to toxic concentrations
of the neutral salts. Longer exjiosure of root tijis to
the toxic solutions would undoubtedly bring about a
degeneration of the nucleus, with a jiossibility of
jiseudomitosis or amitosis before the cell was com-
jiletely dead.

SUMMARY

The toxic effects of the chlorides of jiotassium,
sodium, magnesium, and calcium upon cells of root
tijis of Zea Mays L. were investigated. The roots
were exjiosed to various concentrations of the salt
solutions for 24 hours, followed by fixation for 48
hours in acid and basic fixatives.

The roots were dead at the end of 24 hours in
0.02 M KCl ami NaCl, 0.1 M MgCl,, ami 0.2 iM
CaCM.,, as shown by their inability to continue growth
in distilled water.

Basic fixation of root tijis after the toxic treat-
ments showed a clunij)ing of the cytojilasm about the
nucleus. The resulting vacuoles were large and repre-
sented a fusion of the smaller vacuoles of the normal
cell. A definite decrease in the amount of cytojilasm
was not observed.

The cytojilasm in cells of root tips exposed to
0.1 M concentrations of KCM, NaCl, and iMgCl._. was
either dissolved or badly shrunken by the acid fixa-
tives. The more dilute solutions of these salts, and
CaCL in all tested concentrations, had little effect on
the acid fixation image.

Excejit for a clumjjing of the chromatin by strong
concentrations of the salts, there were no funda-
mental changes in the resting nucleus.

MgCl._, and CaCl.. in killing concentrations jire-
vented mitosis, while KCl and NaCl were toxic in
0.02 M concentrations but allowed normal mitosis in
0.033 M solutions. Aberrant mitotic figures were not
observed.

Botanical LAnortATOKY,
LTnivkrsity of Pknn.sylvaxia,
Phil.\delphia, Tennsylvania

literature cited

JHlUSiniw

A C*«J *^ —

Addoms. R. M. 1927. Toxicity as evidenced by ch:ingos

in the pro'o|)l;isini(' structuro of root hairs of wheat.

Amor. .Jour. liol. 14: 147-165.
Bmlkv, I. W. 1930. The cambiuin and its dcrivativo

tissues. V. A roconnaiss:inco of the vacuonie in

living colls. Zoitschr. Zollforsch. 10: 651-682.
Bamford, R. 1931. Changes in root tijis of wheat and

corn grown in nutrient solutions deficient in calcium.

Bull. Torrey Bot. Club 58: 149-178.
Be.all, R. 1936. Plant Physiol. (In jircss.)
Carlson, J. (I. 1936. Cytologia. (In press.)
Cholodnvj, N. 1923. Zur Frajjo iibor die Bcoinflu.ssung

dos Protoplasnias (lurch mono- und bi\alonto Motall-

ionen. Jioih. Bot. Confralbl. 39 (Pt. 1): 231-238.
Iljin, W. S. 1935. Das Absterbon dcr Pflanzonzoll(>n in

roinen und balancicrtcn Sulzlosungen. Protoi)lasnia

24: 409-430.
Kahho, H. 1921. Zur Konntnis dor Noutralsdzwir-

kun>ion auf das Pflanzonplasnia. liiochoni. Zeilschr.

120: 125-142.
Kahlenbkrc, L., and R. H. Tkik. 1896. On the toxic

action of dissolved sails and their electrolytic dis-
sociation. Bot. Gaz. 22: 81-124.
Karstkn, (I. 1915. Vhcr ombryonalos Wachstuin und

seine Tagesi)oriodc. Zoitschr, Bot. 7: 1-34.
Kerr, T. 1933. The injection of certain salts into the

|irotoi)lasni and vacuoles of the root hairs of Limno-

bium .spoTif/ia. Prot()i)lasnia 18: 420-440.
Lutmax. li. F. 1934. Coll size and structure in i)l:ints

as atfoct(>d by various inorganic elements. Vermont

Agr. Exp. 8.a. Bull. 383.

Reed. H. S. 1907. Tho valuo of certain nutritive
elomonts to i)laiit colls. Annals Bot. 21: 501-543.

RiN(.ER. S. 1884. ConccMniny: tho influonce of saline
inodia on fi.><hos. fic. .Jour. Physiol. 5: 98-115.

Seifriz, W. 1923. Reaction of j)rotoplasm to salts and
antagonistic action of salts and alcohol. Bot. Gaz.
76: 389-402.

Skeex. J. R. 1930. Exporiinonts with Trinnca on an-
tagonism and al>sori)tion. Plant Phvsiol. 5: 105-
118.

SoROKix, H.. AXD A. b. Sommer. 1929. Changes in the
cells and tissu(\s of root tips induced by the absence
of calcium. Amor. Jour. liot. 16: 23-39.

Trie, R. H. 1900. The toxic action of a .series of aci<ls
and of their .sodium salts on Lupinus albus. Amor.
Jour. Sci. 9: 183-192.

. 1930. Tho toxicity of moloculos and ions. Proc.

Amor. Philosop. Sic. 69: 231-245.

Yamaha. G. 1927. I'.xperimontollo zytologischo Boitriige.
I. Mittoilung. Oiiontierung.svcr.suche an dor Wurzcl-
spitzon einiger I'H;inzen. Jour. Fac. Sci. Imp. L'niv.
Tokyo 2: 1-214.

Zirkle, C. 1928. T!io ofToct of hydrogon-ion concentra-
tion upon tho fi> it ion imago of tho various salts of
chromium. Prot >i)lasma 4: 201-227.

. 1932a. Vaci )les in primaiv mcristcms. Zoit-
schr. Zollforsch. Iti: 26-47.

. 1932b. Cytoiogical fixation with the lower fatty

acids, ihr'w con |)ounds and derivatives. Proto-
I)lasma 18: 90-11 .

. 1934. Amin ^ in cytological fixing fluids. Pro-

toplasma 20: 473 482.

\\\r*

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Lophotocarpus spongiosus in Salem County. New Jersey

John M. Fogg, Jr.

River about two mHes sSwe't Jw "^-f ^^^

front in this localiry has been rP If ^""'°"^•"^ Salem County. The river
a narrow dike wh h extenrnnHh * iT'* "'^"' ''''""'^ ''y *he erection of
to the mouth of M II Cr ek Att^ ^^ ^'u" ^'^'"""^ "^ ''''""t ^alf a mile
dike, flooding the lowS J ar^a beS Ttt ' "^ '"! •''°'^^" *'''''"«'' *»>«
water, not shown on anv m«n w^ k ' ^ '"^''*'"^ * '"^'^^ '"'""d body of

passages or " bre^acLrasTS Irt' kn^wnTr ^"' ^'^ '^^' ''^ ^^^"«''*

of mttr„t wSifeiXi; :s r,^:LTur? i' « -^^-^

meager vegetation, including cLerusH^^Zl k *k ^ ., "* ^^'"^ '"PP"*^ ''

Rich., ^i^oc^ana olivacea iS^ZZZaZ^^'iL) iZT 7"''' ''• ^'^'^
natm Michx.. ^cntda cannabma I s.Spu ^"^^'"'^'* <^^ ^unth, yuncu* aeumi.
with these was a plant whTchrffi '"f.^^^'^"' <^<^rnphorata (L.) DC. Associated
although its te.tlon«^ itl "^ !f t ^' """^ *^ ^^ « '^''''' °^ 'Saffi^'"'^.
unlikely ASSn'ot to us *'"'' '"•"'*^ "^'"'^ ""''^^ '* ^^ '^-^

disctv:r:?att:;X^^^^^^ - -^J^^h ^^ « ->«. to the southeast we

just above the upper bo„,er of thrrirr' T ""^^tf, """"^ ""'' ^^"'"^^''.

whio. must be inu^ndatera^^tS^ulX^'hi^VtiJ'''^ b^Lt^T
plants had suffered somewhat from +hp .ffl . , ; ^^*^ locahties the

to collect a representative sene^:Cedm^^^^^^ '"'' '"^'^' '"* ^^ ^'^ «"'«

sugJ's^Sat if SttlU:r^^^^ '"<^.*° ^'- ^«'^ ^-«. •>« -t once
definitely confirmed 'up?n^b^S2wn^;rt^' ' '"''^"^^^ ^'^''^ '^ '«*«,

for thf :uSr:s;iSt:g^^^^^^^^ "-'•^h^'^''*' ^'^ -- -^-

this occasion we investkadh!T ^^ , '°""^"'8 additional specimens. On
the second LoT^oZZ'^.^: ZSJ :^Ti^t Sfd a^"^ T

slough whic^h trave^Tirrrcitir ;dX t^^r^ ^^^^^^

gimca, scattered among which werp ornwm^ . / pr^ence of Peltandra tnr-

We had no means of e^mSgT: ^ZZTXtl.! UTT^'
in many places too deeply submerged to inX ^Z '*"! e^'^'^ ^°' '* *"
followed a winding course off acrossTmarBhlafdandS ^'^P'"'"*'""- but it
see. was occupied by a nearly solid SroX^P^r '/.VriZS ""''

31

^»

22

BARTONIA

Nearly a year later (October 8, 1935) I revisited the locality and succeeded
in crossing the marshes for a distance of perhaps an eighth of a mile. Upon this
occasion the water level was not so high as it had been the preceding autumn and
Lophotocarpus was observed to be restricted to the lower and more submerged
areas. That the colony must indeed be a considerable one was indicated by
the fact that beyond the farthest point to which I penetrated the plants might
be seen to continue in undiminished numbers.

Lophotocarpus spongiosus (Engelm.) J. G. Smith is a species of tidal shores
and brackish muds along the Atlantic coast of North America from New Bruns-
wick to Virginia. It has been definitely known within the Philadelphia local
area only from New Castle County, Delaware, where, between 1866 and 1894,
it was collected at several localities along the Delaware River (Cherry Island,
Wilmington and Delaware City). The herbarium of the Philadelphia Botanical
Club contains 10 sheets from the above stations collected by Commons, Martin-
dale, Canby and Tatnall.

All previous references to the occurrence of Lophotocarpus in southern New
Jersey appear to be unsubstantiated. In Britton's Catalogue^ the plant (as
Sagittaria calycina Engelm., var. spongiosa Engelm.) is credited to Camden
County, New Jersey, on the authority of C. F. Parker, who is supposed to have
collected it in tidal mud along the Delaware River. Dr. Witmer Stone has dis-
posed of this record, however, by insisting that it was actually based upon material
of a submerged form of Sagittaria graminea Michx.* Our collections from
Harrisonville, therefore, appear to constitute the first basis for admitting Lopho-
tocarpus spongiosus to the flora of southern New Jersey. The following specimens
are being deposited in the herbaria of the Philadelphia Botanical Club, University
of Pennsylvania or the Gray Herbarium: Adams, Nos. 1743, 1745, 1746, 1747;
Long, No. 45,271; Fogg, Nos. 7854, 10,003, 10,004.

It is of interest to note that Miss Larsen herself added another station to
those already known from New Castle County, Delaware, when, on July 20, 1934,
she discovered the plant on the northeast side of Pea Patch Island (Larsen No.
867). This was apparently the first time in forty years that Lophotocarpus had
been collected in Delaware. Pea Patch Island, it may be observed, is situated
in the Delaware River almost due west of our Harrisonville station.

University of Pennsylvania.

1 Britton, Cat. Plants N. J. 256 (1889).

2 Stone, Plants Southern N. J. 169 (1911).

4

I

I

1

MYXOMYCETES OF CLARK COUNTY, INDIANA

William D. Gray, University of Pennsylvania.

The Myxomycetes, a group of plants of approximately four hundred
species of world-wide distribution, and commonly known as slime-
moulds, include many species which exhibit characters that are of con-
siderable scientific and popular interest. On the one hand, at certain
periods of their life cycle they show features allying them with certain
lower forms of animal life, while at other periods they have definite
plant-like characters. To the layman, the bright color of some species
and the delicate make-up of the fruiting structures often prove of great
interest. Too often, however, because of their commonly small size and
the inconspicuous coloration of many species, they escape the notice of
many professional botanists as well as others.

Different species show a considerable range in habitat preference,
although a favorable habitat for many species appears to be very rotten
logs in cool, damp woods. Some species may be found on grass, rotten
leaves, soil, or cinders, and in one instance fruiting specimens of
Stemonitui fusca were collected on the painted ceiling of a dairy. Cer-
tain species, e. g., Physarum cinereum and Lycogala epidendrum, appear
to prefer one type of habitat, while others such as Fitligo septica grow
equally well in a variety of habitats.

The first reference to myxomycetes in Indiana which I have been
able to find was made by Underwood (1), who, in 1893, listed twenty-one
species from Putnam, Vigo, and Vermillion counties, those from Vermil-
lion County having been collected by Arthur; he followed this with a
report (2) of three additional species in 1894. In 1897, Olive (3) reported
forty-three species from the region about Crawfordsville; this list in-
cluded thirty species which had not been formerly recorded by Underwood.
Thomas (4), in 1900, recorded thirty species from the same region,
twenty of these being reported for Indiana for the first time. In 1901,
Whetzel (5) described in considerable detail seven species of Stcmotutis
occurring in Indiana and accompanied the descriptions with a key to the
species and some notes on plasmodial behavior; among the seven species
were S. pallida and S. axifcra, which had not been hitherto reported for
the state. In the same year Mutchler (6) listed thirty-eight species
which he had collected in the vicinity of Bloomington, adding nine to
the state list; later, in 1902, he presented a second report (7) on an
extensive collection which was, in the main, obtained near Winona Lake.
By this second list Mutchler added twenty-nine to the known species of
Indiana myxomycetes. In 1909, Barbazette (8) listed fifty-three species
based on collections of J. A. Niewland in 1905; of these, thirty-four were
collected in Indiana, and five species were new ones for the state. Dur-
ing the years 1911 to 1934, the late Professor J. M. VanHook (9, 10,
11, 12, 13, 14, 15, 16) reported various species to this academy at irreg-
ular intervals in conjunction with his reports on Indiana fungi. He in-
cluded forty-one species of myxomycetes in these various papers, adding
five to the state list, which raised the total number of species reported
for Indiana to one hundred and twenty-four.

None of these reports contained records of any species of myxomy-

(09)

RepHnted from the Proceedinga of the Indiana Academy of Science. Vol. V'. 1936

\

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IRREGULAR PAGINATION

70

Proceedings of Indiana Academy of Science

cetes collected in Clark County; in fact, Monroe was the southernmost
county in the state from which species had been reported. Clark County,
particularly that region along the Ohio River, presents malhy favorable
habitats for myxomycetes. In addition to woods along the Ohio, favor-
able localities are also to be found along Silver Creek, Fern Creek (Four-
teen Mile Creek), and Lacasagne Creek. So far, the most fruitful locality
has been in the Arctic Springs region (two miles east of Jeffersonville).
This work, which was started at DePauw University in 1933, repre-
sents collections made during the past three summers. Undoubtedly
more extensive search will reveal many species not included in this
report. Specimens herein referred to are deposited (with the same
numbers under which they are listed here) at the Herbarium of DePauw
University, Greencastle, Indiana. Doubtful specimens were referred to
Dr. G. W. Martin, State University of Iowa. Species marked with an
asterisk are those being reported for Indiana for the first time.

Subclass Exosporeae

Ceratio7nyxa fruticulosa (Muell.) Macbr.

On rotten locust tree in conjunction with sporangia of Hemitrichia
stipitata; woods near C. C. C. & St. L. yards (Jeffersonville), June 15,

1933 (No. 69). On rotten log, Arctic Springs, June 19, 1935 (No. 103).

*Ceratiomyxa fruticulosa (Muell.) Macbr. var. arbuscula.

(Berk. & Br.) Macbr. & Martin.

On rotten log in woods near C. C. C. & St. L. yards (Jeffersonville),
June 20, 1935 (No. 105).

Subclass Myxogastres

Arcyria cinerea (Bull.) Pers.

On rotten log, Arctic Springs, June 5, 1933 (No. 35); June 16, 1933
(No. 28); June 16, 1934 (No. 106); June 5, 1935 (No. 42). On rotten
log, Armstrong Bend, June 19, 1933 (No. 57); in conjunction with
Hemitrichia stipitata, June 19, 1934 (No. 109). Common in early sum-
mer.

Arcyria denudata (L.) Wettstein.

On rotten logs, Arctic Springs, June 16, 1934 (No. 50) ; July 19,

1934 (Nos. 95 and 96); July 23, 1934 (Nos. 3 and 4); June 19, 1935
(No. 29). On rotten stump, Jeffersonville, June 18, 1935 (No. 45).

Arcyria digitata (Schw.) Rost.

On rotten log, Arctic Springs, June 19, 1933 (No. 38). On rotten
locust log in woods near C. C. C. & St. L. yards (Jeffersonville), June 20,

1935 (No. 2).

Arcyria insignis Kalch. & Cooke.

On rotten log near main spring, Arctic Springs, June 30, 1933 (No.
6); on cinders, June 30, 1933 (No. 58).

Badhamia utricularis (Bull.) Berk.

On rotten tree along Fourteen Mile Creek, about 3 miles below
Beaver Hole, June 23, 1935 (No. 54).

Botany

71

I

\

Comatricha aequalis Peck.

On rotten log in damp ravine, Arctic Springs, July 18, 1935 (No. 91).

Comatricha irregularis Rex.

On rotten elm stump in woods near Lacasagne Creek (3 miles N. E.
of Jeffersonville), June 20, 1933 (No. 65).

Comatricha laxa Rost.

On rotten log in damp ravine, Arctic Springs, July 19, 1934 (No. 98).

Comatricha longa Peck.

On rotten sycamore log about one mile below Beaver Hole (Fern
Creek), June 23, 1935 (No. 55).

Comatricha typhoides (Bull.) Rost.

On rotten log, Arctic Springs, June 19, 1935 (No. 33) ; July 18, 1935
(No. 92).

Cribraria microcarpa (Schrad.) Pers.

On rotten log in woods near Arctic Springs, June 26, 1935 (No. 82).

Dictydium cancellatum (Batsch) Macbr.

On rotten log, Arctic Springs, June 19, 1933 (No. 40) ; July 23, 1934
(No. 62). On rotten log in woods about one mile n. of Memphis, June
8, 1935 (No. 107).

*Dictydium cancellatum var. purpureum Macbr.

On rotten log, Arctic Springs, June 16, 1933 (No. 67); June 16,

1934 (No. 13).

Diderma hemisphaericum (Bull.) Rost.

On rotten oak leaf, Arctic Springs, June 19, 1933 (No. 41).
Fuligo septica (L.) Weber.

On rotten tree stump, Jeffersonville, June 18, 1935 (No. 44).
Hemitrichia stipitata (Massee) Macbr.

On rotten log in woods near C. C. C. & St. L. yards (Jeffersonville),
June 15, 1933 (No. 47); in conjunction with Ceratiomyxa fruticulosa,
June 15, 1933 (No. 69); June 15, 1934 (Nos. 32 and 66). On rotten log
in damp ravine, Armstrong Bend, June 19, 1933 (No. 60). On rotten
log in drift pile, Fern Creek (3 miles below State Road 62), June 23,

1935 (No. 53). On rotten log, Arctic Springs, Sept. 9, 1934 (No. 110);
in conjunction with Comatricha typhoides, July 28, 1935 (No. 92).

Hemitrichia vesparium (Batsch) Macbr.

On bark of fallen tree. Tunnel Mill Reservation, April 14, 1933 (No.
56).

Lamproderma arcyriouema Rost.

On rotten log in damp ravine, Arctic Springs, July 18, 1935 (No.
102).

Lycogala epidendrum (L.) Fr.

On rotten ash tree, Arctic Springs, June 23, 1933 (No. 87).

72

Proceedings of Indiana Academy of Science

Lycogala cxignum Morgan.

On rotten ash tree in woods near C. C. C. & St. L. yards, June 15,
1934 (No. 31). On rotten log, Arctic Springs, June 26, 1935 (No. 80).

Physarum cinerenm (Batsch) Pers.

On living grass in City Hall lawn, Jeffersonville, August 25, 1933
(No. 5). The entire fructification covered a circular area approximately
one foot in diameter.

Physarum ghbuliferinn (Bull.) Pers.

On rotten log, Arctic Springs, Sept. 19, 1933 (No. 83); Sept. 19,
1934 (No. 85).

Physarum nucleatwrn Rex.

On rotten log in woods near C. C. C. & St. L. yards (Jeffersonville),
June 20, 1935 (No. 1).

Physarum nutans Pers.

On rotten log, Arctic Springs, June 19, 1935 (No. 43) ; July 18, 1935
(No. 90).

* Physarum pulcherrimum Berk, and Rav.

On rotten log in woods about 1 mile n. of Memphis, June 18, 1935
(No. 108). On rotten log, Arctic Springs, July 18, 1935 (No. 101).

Physarum vi)i(lr (Bull.) Pers.

On rotten log, Arctic Springs, June 19, 1933 (No. 39); June 19,

1934 (No. 97) ; June 19, 1935 (No. 84) ; July 18, 1935 (No. 94). On rot-
ten elm stump in damp woods near Lacasagne Creek, June 20, 1933 (No.
59). Common.

Stemonitis axifera (Bull.) Macbr.

On rotten ash tree, Tunnel Mill Reservation, June 14, 1933 (No.
14). On rotten log, Arctic Springs, June 16, 1934 (No. 12).

Stemonitis fusca Roth.

On fallen oak leaves in woods about 1 mile N. of Memphis, June 18,

1935 (No. 104).

Stemonitis pallida Wingate.

On rotten limb in Junior High School yard, July 14, 1934 (No. 26).
On rotten log, Arctic Springs, July 18, 1933 (No. 61); July 18, 1935
(Nos. 88 and 100).

Stemonitis smithii Macbr.

On rotten log, Arctic Springs, June 19, 1933 (No. 36); June 20,
1933 (No. 7); July 18, 1935 (No. 89). On rotten elm stump in woods
near Lacasagne Creek, June 20, 1933 (No. 30).

Stemonitis splendens Rost.

On rotten log in damp ravine, Arctic Springs, July 18, 1935 (No. 99).

Trichia favoginea (Batsch) Pers.

On rotten elm stump in woods near Lacasagne Creek (three miles
n. e. of Jeffersonville), June 20, 1933 (No. 16).

Botany

73

Acknowledgments

The writer wishes to thank Dr. Winona H. Welch, who originally
suggested this problem. Dr. Truman G. Yuncker, for invaluable criticisms
of the manuscript, and Dr. G. W. Martin for checking doubtful species.

Literature Cited

1. Underwood, Lucien M., 1894. Report of the Botanical Division of
the Indiana State Biological Survey. Proc. Ind. Acad. Sci. 1893:13-67.

2. Underwood, Lucien M., 1895. Report of the Botanical Division
of the Indiana State Biological Survey for 1894. Proc. Ind. Acad. Sci.
1894:144-153.

3. Olive, E. W., 1898. A list of the Mycetozoa collected near
Crawfordsville, Indiana. Proc. Ind. Acad. Sci. 1897:148-158.

4. Thomas, M. B., 1901. Cryptogamic collections made during the
year. Proc. Ind. Acad. Sci. 1900:121-123.

5. Whetzel, H. H., 1902. Notes on the genus Stemonitis. Proc.
Ind. Acad. Sci. 1901:261-266.

6. Mutchler, Fred, 1902. A collection of Myxomycetes. Proc. Ind.
Acad. Sci. 1901:291-292.

7. Mutchler, Fred. Myxomycetes of Lake Winona. Proc. Ind. Acad.
Sci. 1902:115-120.

8. Barbazette, L., 1909. Tentative list of Myxomycetes of north-
ern Indiana and southern Michigan. The Amer. Midi. Nat. 1 :38-43.

9. VanHook, J. M., 1912. Indiana Fungi. II. Proc. Ind. Acad. Sci.
1911:347-354.

10. VanHook, J. M., 1916. Indiana Fungi. III.
1915:141-148.

11. VanHook, J. M., 1921. Indiana Fungi. V.
1920:209-214.

12. VanHook, J. M., 1922. Indiana Fungi. VI
31:143-148.

13. VanHook, J. M., 1924. Indiana Fungi. VII. Proc. Ind. Acad.
Sci. 33:233-238.

14. VanHook, J. M., 1926. Indiana Fungi. IX. Proc. Ind. Acad. Sci.
35:233-236.

15. VanHook, J. M., 1930. Indiana Fungi. XII. Proc. Ind. Acad.
Sci. 39:75-83.

16. VanHook, J. M., 1935. Indiana Fungi. XIII. Proc. Ind. Acad.
Sci. 44:55-64.

Proc. Ind. Acad. Sci.
Proc. Ind. Acad. Sci.
Proc. Ind. Acad. Sci.

t

NOTES ON PLASMODIAL BEHAVIOR OF STEMONITIS

FUSCA ROTH

William D, Gray/ University of Pennsylvania

So far as can be determined from available literature on Myxomy-
cetes, little attention has been paid to the interesting phenomena of
structural and color changes which accompany the development of
sporangia from the mature Plasmodium. Some references are made to
the color changes which occur in the Plasmodia of various species, but
the changes which take place in the development of the sporangia are
generally not described. The genus Stemonitis is an example of one
in which comparatively little is known concerning the Plasmodia. Of the
seventeen species which belong to this genus, there are only eight of
which the Plasmodia are known. In a genus so difficult taxonomically as
is Stemonitis, every feature of each species should be known if possible.
The following is a report of such changes as have been observed to take
place in the development of Stemonitis fiisca from Plasmodium to mature
fruiting structures.

The Plasmodium of S. fusca, a pearly-white, tubercular mass, was
collected at 3:00 p. m., November 2, 1933. It was found on the lower
side of a rotting ash log in a ravine about one and one-half miles west of
Greencastle, Indiana. The Plasmodium was elliptical in outline, measured
2.0x2.5 cm., and was approximately 5 mm. thick.

The Plasmodium, together with a portion of its substratum, was
taken immediately to the laboratory, placed on a glass plate, and covered
with a large beaker which was sealed to the plate with vaseline. A small
dish of water was enclosed under the beaker to provide sufficient moisture
for the growth of the Plasmodium. During the daytime, the Plasmodium
received natural light from a north-facing window, and artificial light
from a seventy-five-watt electric lamp. With only one exception, when
the temperature fell to 12''C. for a few hours, the temperature of the
laboratory remained at approximately 19^ C. Under these conditions,
fructification was completed in eighty-nine hours. The following sum-
mary shows the changes that took place:

8:30 a. m. (Nov. 3): A portion of the Plasmodium had moved from
the side to the upper surface of the block of wood, and differentiation had
begun. Sessile sporangia, each with a bulbous enlargement at the top,
■were discernible. The color of the sporangia was still pearly-white. No
stipes were as yet evident.

12:30 p. m. (Nov. 3): Purplish-black stipes (2.5-3.0 mm. in length)
had developed. The sporangia had developed to a length of 4.5-6 mm.
and were faintly tinged with pink. As at the time of the preceding
observation, each sporangium was capped with a small bulbous enlarge-
ment. None of the Plasmodium proper remained.

5:15 p. m. (Nov. 3): The bulb at the top of each sporangium had
disappeared, the color was pale lavender, and the stipes had increased
in length to about 6.0 mm. The total height of the sporangia (which
were very moist with drops of water clinging to the almost black stipes)

made.

^Graduate student 'n Botany, DePauw University, when these observations were

(74)

Reprinted from the Proreedinps of the Indiana Academy of Science, Vol.

19.1 a

Botany

75

A,

varied from 12 to 14 mm. The sporangia were closely aggregated with
the outermost members of the group curving toward the center, thus
giving the cluster a rounded appearance at the top.

7:30 a. m. (Nov. 4): The sporangia had become very dark brown to
almost black in color. No changes in size or general form of structure
had occurred since the preceding observation. The sporangia were still
coherent because of their damp condition. During the preceding night
the temperature had dropped to 12°C., but no reaction to the lower tem-
perature was evident.

8:00 a. m. (Nov. 6): The fructification was completed, the stipes
were shiny black, and the sporangia had become rich chocolate brown.
The sporangia, spores, capillitium, and capillitial net showed no devia-
tion from those of a naturally developed fructification of this species.
The question as to whether or not each species exhibits individual
features during fructification can be answered only when many observa-
tions have been made. Macbride and Martin (1, p. 162) state that the
Plasmodium of S. fusca passes from white through blue and black, which
color changes differ from the observed color changes outlined above.
Whetzel (2, pp. 261-266), working with another species, S. splendens,
observed that the fructification passed from the pearly-white Plasmodium
through purple-black, dark brown, light brown, and finally, with the
shedding of the spores, purple-brown. Undoubtedly, further observations
on this species, as well as additional ones, will lead to interesting com-
parative results, which may, in a measure, serve to aid in the separation
and identification of members of this difficult genus.

Aside from its possible taxonomic value, the color changes accom-
panying the changes from Plasmodium to fruiting structures are also
of some interest physiologically. Seifriz and Zetzmann (3, pp. 175-179,
PI. Ill), working with the yellow Plasmodium of Physanim polycephalum,
which they have long cultured, have found the plasmodial color to be due
to a yellow pigment, which, they suggest, belongs to the group of respira-
tory ferments known as flavones, lyochromes, or flavins. These workers
have found that the yellow pigment is an acid-alkaline indicator, have
calibrated the pigment as a pH indicator, and have found that the
Plasmodium undergoes changes ranging from pH 8, when fruiting, to pH
1.6 (possibly 1.2), when a sclerotium is formed. The latter finding well
fits, and more or less partially substantiates the ideas of the late Pro-
fessor Macbride (4, p. 22), who placed the order Physarales as lowest in
the sequence of orders and inferred that the retaining of lime to the last
by the Plasmodium was indicative of lower rank. Whether or not the
color changes observed in Sfemovifis fusca are due to the pre.sence of an
acid-alkaline indicator must be determined by further work, which is
of course, hindered by the difficulty in obtaining Plasmodia of this species'
If Macbride's idea is the correct one. and an indicator is present in
Sfemovifis fuf>ca, it is to be expected that at time of fruiting the pla.s-
modium would exhibit a somewhat lower pH than that exhibited by
Physnnon polycephalum.

Acknowledgments

The writer is indebted to Dr. Truman G. Yuncker, DePauw Uni-

76

Proceedings of Indiana Academy of Science

versity, and Dr. William Seifriz, University of Pennsylvania, for their
criticisms in the preparation of the manuscript.

Literature Cited

1. Macbride and Martin, 1934. The Myxomycetes. Macmillan.
New York.

2. Whetzel, H. H., 1902. Notes on the genus Stemonitis. Proc.
Ind. Acad. Sci. 1901 :261-266.

3. Seifriz and Zetzmann, 1935. A slime-mould pigment as indicator
of acidity. Protoplasma 23:175-179.

4. Macbride, Thomas H., 1932. The North American slime-moulds.
Macmillan. New York.

i

I

f Reprinted from Phytopathology, March, 103(5, Vol. XXVT, No. 3, ])p. 293-294. J

I

J

A Metliod for Staining Rust Mycelium in Woody Tissues. — During the
course of a study of Scotch pine, infected with a species of Peridermium, the
author^ had occasion to use with good success the orseillin BB-anil in-blue-
staining procedure described by Strasburger, pp. 392 and 161.''

The staining of rust mycelium and haustoria in tissues was so effective
that the method suggested itself as valuable in the routine microscopic
diagnosis of the white-pine blister rust. The method perhaps most often
used for this purpose at present is the short method described by Colley'
with the use of safranin and light green. The Strasburger procedure has
been adapted to the shorter Colley method, so that sections may be stained
in a comparatively short time.

Sections of white pine bark and wood, cut on a freezing microtome or by
hand, are placed in water without previous fixation. The water is replaced
by a saturated solution of orseillin BB^ in 3 per cent acetic acid that is
allowed to remain for 10 to 12 hours. The sections are then rinsed in 40
per cent ethyl alcohol in order to remove excess stain. A saturated solution
of anilin blue in 3 per cent acetic acid is then added and allowed to remain
for 15 to 30 minutes. A rapid rinsing in 40 per cent ethyl alcohol to re-
move any excess stain is followed by the addition of 95 per cent alcohol for
2 to 5 minutes. This is followed by absolute alcohol, which in turn, is re-
placed by fresh absolute alcohol. The sections may then be transferred to
clove oil in which they may be examined if the mount is to be temporary,
only. The clove oil may be washed away by xylol, and balsam or gum
damar added if the sections are to be mounted permanently.

The most satisfactory results are usually obtained when differentiation
with 40 and 95 per cent of ethyl alcohol has been carried on, so that the
sections appear a rather light purple. It will be found necessary to alter the
time schedule somewhat according to the thickness of the sections and ac-
cording to the amount of resin the pine contains, as portions abundant in
resin stain heavily with orseillin.

In properly differentiated tissues the mycelium stains violet to blue,
lignified or suberized tissue stains red, parenchyma cell walls usually blue,

1 Colley, R. H. Diagnosing white-pine blister-rust from its mycelium. Jour. Agr.
Research [U.S.] 11: 281-286. 1917.

2 Hutchinson, W. G. Resistance of Finns sylvestris L. to a gall-forming Peridermium.
Phytopath. 25: 819-843. 1935.

3 Orseillin BB may be obtained from Akatos Inc., 55 Van Dam St., New York, N. Y.
It is not the same stain as orsein.

* Strasburger, E. Das botanische Praktikum. 7th Ed. 883 pp. G. Fischer, Jena.
1923.

i

I Rci)riiiti'a from Phytopatiioixjgy, March, 193(), Vol. XXVT, No. 3, i)p. 29:5-294.]

A Method for Staining Rust Mycelium in Woody Tissues. — During the
course of a study of Scotch pine, infected with a species of Peridermium, the
author^ had occasion to use with good success the orseiilin BB-anil in-blue-
staining procedure described by Strasburger, pp. 392 and 767."

The staining of rust mycelium and haustoria in tissues was so effective
that the method suggested itself as valuable m the routine microscopic
diagnosis of the white-pine blister rust. The method perhaps most often
used for this purpose at present is the short method described by Colley'
with the use of safranin and light green. The Strasburger procedure has
been adapted to the shorter CoUey method, so that sections may be stained
in a comparatively short time.

Sections of white pine bark and wood, cut on a freezing microtome or by
hand, are placed in water without previous fixation. The water is replaced
by a saturated solution of orseiilin BB^ in 3 per cent acetic acid that is
allowed to remain for 10 to 12 hours. The sections are then rinsed in 40
per cent ethyl alcohol in order to remove excess stain. A saturated solution
of anilin blue in 3 per cent acetic acid is then added and allowed to remain
for 15 to 30 minutes. A rapid rinsing in 40 per cent ethyl alcohol to re-
move any excess stain is followed by the addition of 95 per cent alcohol for
2 to 5 minutes. This is followed by absolute alcohol, which in turn, is re-
placed by fresh absolute alcohol. The sections may then be transferred to
clove oil in which they may be examined if the mount is to be temporary,
only. The clove oil may be washed away by xylol, and balsam or gum
damar added if the sections are to be mounted permanently.

The most satisfactory results are usually obtained when differentiation
with 40 and 95 per cent of ethyl alcohol has been carried on, so that the
sections appear a rather light purple. It will be found necessary to alter the
time schedule somewhat according to the thickness of the sections and ac-
cording to the amount of resin the pine contains, as portions abundant in
resin stain heavily with orseiilin.

In properly differentiated tissues the mycelium stains violet to blue,
lignified or suberized tissue stains red, parenchyma cell walls usually blue,

1 Colley, R. H. Diagnosing white-pine blister-rust from its mycelium. Jour. Agr.
Research [U.S.] 11: 281-286. 1917.

2 Hutchinson, W. G. Resistance of Finns sylvestris L. to a gall-forming Peridermium,
Phytopath. 25: 819-843. 1935.

3 Orseiilin BB may be obtained from Akatos Inc., 55 Van Dam St., New York, N. Y.
It is not the same stain as orsein.

* Strasburger, E. Das botanische Praktikum. 7th Ed. 883 pp. G. Fischer, Jena.
1923.

w

»>

and nuclei and cytoplasm red. Haustoria, penetrating tracheids and sur-
rounded by lignified callosities, stain deep red in contrast to the bluish
mycelium outside the cell wall.

Comparison of blister-rust-infected tissues stained by the above method
with similar tissues stained with safranin and light green shows that in the
former case the mycelium is mueh more obvious and clear-cut. Even in
dried woody tissues, it is a simple matter to recognize the rust mycelium. —
W. G. Hutchinson, Department of Botany, University of Pennsylvania,
Philadelphia, Pa.

^

3

Separately printed, without change of paging; from Bulletin of thk Torrky
PiOTANicAL Ci.uR 63: 477-481. November, 1936.

Wound responses of Ficus australis

John Austin Jump
(with six figures)

Upon reviewing the literature on wound reaction and cicatrization of
leaves, one gathers the impression that a leaf subjected to a traumatic
stimulus will give a constant type of reaction if the conditions under which
the experiments are performed are identical. Working with leaves of
Ficus australis Willd. {Ficus ruhiginosa Desf.) the writer has observed a
number of variations in response, even in the same wound.

The outstanding publications on cicatrization prior to 1930 have been
reviewed by Wylie (1930). The writer wishes however, to briefly mention
certain papers that have a specific bearing upon this investigation. Mas-
sart (1898) notes that in foliar wounds on Clivia miniata Regel the neigh-
boring cells proliferate and completely repair the wound. Nuphar luteum
Sibth. & Smith was found to react by filling the large lacunae with prolif-
erated cells in the vicinity of the wound. Certz (1918) is reported by Wylie
to have observed a marked hypertrophy of mesophyll cells bordering the
tunnels of leaf miners. Blackman and Mathaei (1901) in an extensive in-
vestigation upon the wound reactions of Prunus laurocerasus L., find that
the usual cicatrice is not formed when the material is kept moist in a beaker.
Buscalioni and Muscatello (1911) were apparently the first to work with
Ficus australis, but do not report the results that the writer has observed.
Their paper is chiefly remarkable for the variety of methods which they
employed in wounding many of the species which they enumerate. Krieger
(1935) called attention to mesophyll proliferation and hypertrophy in
Ficus australis but made no detailed study of it.

An understanding of the normal anatomy of the leaf of Ficus australis
is necessary for the interpretation of wound reaction (fig. 1). The upper
epidermis is multiple and heavily cutinized. The hypodcrm consists of a
layer of very large parenchymatous cells, one to two layers in thickness, in
which occur idioblasts containing cystoliths. These idioblasts usually
greatly exceed the normal hypodermal cells in size, and may be so large as
to crowd out the layers of palisade immediately beneath. The palisade is
double and contains numerous tanniniferous idioblasts which are usually
confined to the upper and more compact layer of the palisade. The spongy
tissue is very loosely organized and has large intercellular spaces. Tannin
and latex occur in the spongy tissue, usually being found in more or less
isodiametrical cells. Vascular structures show considerable variation, but
the collateral bundle seems to be the most typical. Beneath the spongy
tissue there is a low palisade, one cell in thickness, and a double layer of

477

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[VOL. 63

jump: wound responses

479

cells composing the epidermis. The outer layer of this epidermis is com-
posed of cells which are smaller and have thicker walls than those of the
inner layer and is heavily cutinized. Stomata were found exclusively in the
lower epidermis and sunken stomata are occasionally found.

The leaves upon which this study is based were wounded by cuts 4-12
mm. in length, using a sharp razor. The plant was approximately six years
of age and grew in an unshaded position in one of the greenhouses of the
University of Pennsylvania. The wounding was done during the late fall
and winter months, and the leaves were allowed to remain upon the plant
until collected for fixation. Material was fixed in Cohen's VIIc fixing fluid
(1935) and stained in safranin and light green.

A few of the sections showed a definite cicatrice and pseudo-cicatrice,
but in many cases a cicatrice failed to develop. In the latter instances the
pseudo-cicatrice was usually either lacking entirely or only partially de-
veloped, and hypertrophied spongy tissue filled the intercellular spaces. As
the proliferation of the spongy tissue continues, the new tissue fills the
space caused by the wound and the surfaces of the wound are again or-
ganically united (fig. 3). In some cases one surface of the wound prolifer-
ates as described above, while the opposite surface cicatrizes (fig. 4).

When a cicatrice is not formed, the first reaction is the proliferation of
cells in the spongy tissue. These cells are frequently hypertrophied and
filled with granules of various size (fig. 2). The microchemical tests em-
ployed by the writer in attempting to determine the nature of these gran-
ules tended to indicate that they were derived from one of the constituents

Explanation of figures 1-6

Key to Lettering

«, cicatrice; cu, cell from spongy tissue becoming cutinized; tv, cvstolith; ^/, dead

tissue; hd, hypoderm division; /. latex cell; ps, pseudo-cicatrice; ti, tanniniferous idio-

blast; A-, cells containing granules of undetermined nature; possibly a constituent of

latex.

Fig. 1. Normal leaf structure.

Fig. 2. Beginning of proliferation of spongy tissue following wounding
Fig. 3. Union of wounded surfaces. One of the cells of the spongy tissue in the re-
gion of the epidermis has secreted cutin on one side.

Fig. 4. Normal cicatrization on left surface of wound. Proliferation of cells with
only a small amount of dead tissue on the opposite surface.

Fig. 5. Wounded section showing hypoderm becoming meristematic
Fig. 6. Wound severing vascular bundle, showing spongy tissue filling the space
between the cut ends of the bundle.

The sections illustrated in Figs. 2, 4, and 5 are all from the same wound and were
fixed 32 days after wounding.

}ic cu

t

480

BULLETIN OF THE TORREY CLUB

f

I

[VOL. 63

of the latex. The above reactions may begin within five rfav« nf ^^

and continue for three weeks or more, until he woL^^Ji do^Th^e
spongy tissue often proliferates so vigorously that it MUtL u

the wounded surfaces ordinarily occupied by 'he pa^de andT T"

.c in a tew cases (hg. 5). Ihis suggests that if the growth of the snon.,,,

Zlite h"'r'""""^ P""^'' ^° ^^P'^^'^' -d ins'ert itself btweeHhe
opposite hypoderms and palisades, we might exoect m =„m» ^*'*'" ^"^

complete restoration of the original leaf stTuctuT ri '^''' ^ "''""'

noted that a mesophyll cell which ha'd gLTo^t toTplT;:; Z
epidermis apparently became cutinized preoaratorv tl f .•

v^civ Liun diso occurs It the leaf is vounff • the prla^^c r^f f k« i

in such cases, tending to gape excessively When a matnrfl ? '

part account for the ah^enr^ of o "^^- ^ ^^^ ^^y »"

the surface which ckatSd^ f T' u'''"'''' '" "'^''^'^ '^is occurred,
food supply by th wo nd as'.' '° ^''^ ''^"" ™^ °*'^ ^^^ '^^ —

very' ituz; ei°F:::t:: rr :hr r ^^^-^ i ---^ -^''- ••"

more detailed studies of othe 3^2 will rlvelT " '7'""'' '^ '''''''' '''''
responses. ^ '" "^''^^' '™''^'- unreported wound

The writer wishes to express appreciation to Professor H H Vn w
h.s interest and assistance in this investigation. ''' '°'

Botanical Laboratories,

University of Pennsylvania,
Philadelphia, Pa.

I

1936]

jump: wound responses

481

Literature cited

Blackman, F. F. and Mathaei, G. C. L. 1901. On the reaction of leaves to trauma-
tic stimuH. Ann. Botany 15: 533-546.

Buscalioni, L. and Muscatello, G. 1911. Contribuzione alio studio della lesione
fogliari. Malpighia 24: 27-153.

Cohen, I. and Doak, K. D. 1935. The fixing and staining of Linodendron tulipifera
root tips and their mycorrhizal fungi. Stain Tech. 10: 25-32.

Gertz, O. 1918. Kallushypertrofier och nagra i samband darmed staende anato-
miskt-fysiologiska forhallenden hos minerade blad. Bot. Notiser 3: 121-
139.

Krieger, S. 1935. A study of cicatrization. Master's thesis, University of Penn-
sylvania (unpublished).

Massart, J. 1898. Cicatrisation chez les vegetaux. Memoires couronnes et autres
memoires Academic Royale de Belgique 57: 2-68.

Wylie, R. 1930. Cicatrization of leaves. Bot. Gazette 90: 260-278.

^v

♦'-<-

•♦.

-^

i V

-» *■

»A

• « »

-'»•*

ROOT DEVELOPMENT OF PITCH PINE, WITH SOME COM-
PARATIVE OBSERVATIONS ON SHORTLEAF PINE '

By William Everett McQuilkin 2

Formerly jieU assistant, Allegheny Forest Experiment Station,^ Forest Service

United States Department of Agriculture '

INTRODUCTION

The development of the root systems of trees constitutes a rela-
tively unexplored field of botany. Most of the older references to
this subject have been based on casual observations of exposed roots
along roadside cuts and eroded stream banks, or of the upturned
roots of wind-thrown trees. Hence, descriptions of the root systems
ot trees generally have been incomplete, if not actually inaccurate.

Ihe extensive investigations of the roots of herbaceous and shrubby
plants, particularly those by Cannon (J) < and Weaver {26, 27, 28)
led naturally to a greater interest in the roots of trees. However, the
i-elatively large amounts of time and labor required have tended to
discourage work in this field. Problems arising in the nurseries and
the relative facility with which small plants can be studied have
directed research mostly toward the seedling stages. Tourney
described and classified the seedling root systems of many of the more
important eastern species {25) and stimulated manv other investio-a-
tions of a similar character. " ^

A few studies of the roots of older trees have been carried out in
this country. In general, the work done thus far has been of a
prelmunary character, and gives neither a complete picture of the
root growth of a species from seedhng to maturity on any particular
site nor an adequate account of the root reactions of a species to the
various sites on which it may grow. Woodroof's studies {30, 31) on
the pecan constitute an exception to this statement.

Somewhat more work has been done in the north-European coun-
tries. The German work has been summarized bv Biisgen and
Miinch (4). Perhaps the most complete studv of any one species
was earned out by Laitakari (77) in Finland on Scotch pine {Finns
■^l/lrestn.iL.).

The species chosen for the present investigation was pitch pine
iPfnus rigida Mill.), because of its wide distribution tliroughout the
Fastem States and its remarkable tolerance of a wide range of unfavor-
able situations. Particularly because of this latter qualit v, the species
promises to assume increasing importance in the fores tation programs
of the future, although it is not an important timber tree at present.
Pitch pine is prominent among the botanicallv interesting flora of
the pine barrens of New Jersey. The ability of the species to with-
stand fire evokes the wonder and admiration of all who know that
unique section.

• Received for publication Aug. 20, l!)3o; issued Fi-bruarv, 1936.

- Completion of this study was made iwssible througii fel'low ships granted by the Morris Arboretum and
tlie Lniyersity of Pennsylvania. Acknowledgment is due E. T. Wherry, H. H. York, and K. D. Doak
for helpful criticism and suggestions.

J Maintained at Philadelphia, Pa., in cooperation with the University of Pennsylvania

* Reference is made by number (italic) to Literature Cited, p. 1015.

Journal of Agricultural Research.
Washington, D. C.

29019-36-

-1

(983)

Vol. 51, no. 11
Dec. 1, imr>
Key no. F-74

--*•>

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IRREGULAR PAGINATION

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Journal of Agricultural Research

Vol. fil, no. 11

Dec. 1, t«39

Definite root studies of pitch pine have apparently never been

Te roottwZ'/'^^r"*** ^^'"^ ^T'^^ s?ateme/ts "concerning
the root development of the species, and pubUshed a small drawing
of a tree with its root system. His observations obviously were of f
most cursory character, and his figure is entirely inadequate as a
portrayal of a root system. lUick and AughanbLgh (15) mention

becomes relatively less sigmficant in older trees. They measured
the upturned root mass of a wind-fallen tree, recording 18 fe™t?s the
diameter of the root system of a specimen 82 feet talf Such figures
^u^^ ^V^^tionabe value; probably not more than one-fourth 7f the

W SeTr^^^^KirreTfT ^^^ ^"P'^^^''*^^ '" ^^« -^^ *-"

aJ^eVirS^fl^^^^^^^

pine, and of most other tree species. More precise knowledge of the
root habits of trees is requisite for the judicious plannbg of forest^!
ton programs. In this investigation, the major objective was To
study on one site a developmental series of root systems of pitch
pine from the seedling stages to maturity. SeconaaTobject ves
mcluded comparative observations of the root systemroTnormal an^l
weak trees., of trees growing on different sites, and of pitch pTne and
shortleaf pine (Ptnus echinaia Mill.) growing on the same site!

FIELD WORK
DESCRIPTION op SITE

The major part of the field work was done on the tract of the
Allegheny Forest E.xperimcnt Station in the Lebanon State Forest n
New Jersey. Some data were obtained from the Ockanickon area of
the experiment station near Medford, N. J., and from the Mont \lto
W», "'^ ■* '" «°" he™ Pennsylvania. Both of the locations Tn \ew

i^^o^L-^jy t Z^sfc"er' t^^r'"^ "^ ''''-'' '- ^-" ^-^^^

IT^ tf^^ ^'f 'y ''°P'"^- °''^'"''S« i^ S°°d ; cx-cax^tl^ns Vdeep
as 9 feet did not encounter water, even on the lower ground Tb^
soil of the ridge is typical Lakewood sand; on thefower ground h

SlvT'!^'""* A^V"^, *'"«. S'^^-fr''^ tvpes. At a few Xes a
hS^ harder dark-colored layer was encountered below «ie' 1

vefoned "^^1'^" "^ ^^^ ^*-- ''°''?^ ''''^''' b"* it was not weU de-
veloped. These minor variations in soil did not seem to evert «nt

appreciable du-ect effect on the behavior of the pine r^ots ^

A recent paper by Lutz (W) gives detailed data on the soils of tl,<.
section and a quantitative expression of tlieir poor nuahtv HU
determinations show that the pine-barren soils^re characterized hx-
^ l^J^^i^T*'*""**' "f ''^"^; (2) low percentages of clarand total

Root Development of Pitch Pine

985

<

4

The forest cover is a mixture of white, black, and chestnut oaks

itenes Tb if ^'^^^ Lam (?. j^rinu, L.) with pitch and short-
leat pmes The oaks are mostly sprout growth averaginff 4 to 5
inches m diameter breast high,^ and represent the growth since the
last cutting for charcoaL The pines are of all sizes^p to iTinches
1 An ^^ ^^ . ^^^^^^" ^^^^ ^^^ approaching, or perhaps exceed

100 years of age. Apparently the area has not suffered sever^eburnTng

abundXt ""' '^ ""^ '''' '^P^'''^ P'""^" ""^ ''^™"' '^^^^ ^^"^ ^«i^Jy

The ericaceous ground cover, typical of the section, is sparse on the
ridge and composed mostly of dry-land blueberry (Vacciniumvacl
lans Kalm) Lower down on the slopes, black huckleberry (^aX-
sacia baccata (Wang.) C. Koch) becomes predominant, wUh dande-
flatf ^ A I'nl''^ (L.) Torrey and Gray) appearing also on the lowest
fiats. A dense ground cover of the huckleberries rather effectivelv
excludes pme seedlings. The natural inference is that saS and
older trees growing among them got started following a ground fire
which had tempomily checked the huckleberries. The majoritv of
the pmes m seedling stages are now found on the higher and 'less
densely vegetated parts of the area.

METHODS

The root systems were exposed by the dry method. The first step
in the case of trees larger than saphng size was to uncover the root
crown to a depth of 1 foot or more and radially about 3 feet This
operation was of necessity, largely done by hand in "badger " fashion
With the. root crown thus revealed, the most suitable position for
making deeper excavations with the least destruction of roots could be
determined. A hole of sufficient size to allow a man to stand in it
and use a spade was then dug alongside the stump. Its ultimate
depth and the size necessaiy to permit spading movements and pre-
vent caving of the walls were determined, of course, by the depth of
the taproot. By careful caving of the wall toward the stump, the
root crown and taproot could be clearly exposed without serious muti-
lation. With the bases of the lateral roots thus revealed, represen-
tative ones could be selected and followed in any degree of detail
desired Exposure of the laterals also was largelv a hand process
although a trowel and small hand ax were useful.' An ice pick was
helpful in freeing the finer roots from the soil. Excavation of vertical
brandies of the large laterals necessitated digging a hole in the same
manner as for the taproot.

The dry method has several advantages over the alternative wet
method, m which the roots are washed from the soil with a stream of
water under pressure. The former can be used by an investigator
working alone, it does not require proximity to a source of water or
to a road by which water may be hauled, and it requires only the
simplest outlay of equipment. A spade, an ax, a trowel, a hand ax
an ice pick, a ruler, and a few small pails or jars of water for savinc
specimen roots constitute the essentials for field work To cet
complete detail of the finer root branches in sandy soils by drv exca-
vation IS not unduly difficult. The wet method is more suitable
However, for getting out seedling root systems, or parts of larger

* Four and a half feet above the ground; ahbrevi iff^.i "d. b. h."

J,

986

Journal oj Agricultural Research

Vol. 51, no. 11

systems, intact. Considerable breakage and desiccation of finer
roots are unavoidable by the dry method.

In this study, no attempt was made to obtain jcomplete intact
root systems except in the case of small seedlings. On larger trees
the taproot and only a limited number of representative primary
laterals were followed to the tips. Likewise, only representative
samples of the higher orders of branching were examined in detail.
Experience soon showed the general range of variation to be expected.
Excavation of all the roots would have consumed more time than the
results would warrant in a study of general root habit.

GENERAL FEATURES OF PITCH PINE ROOT SYSTEMS

The root system of pitch pine may be regarded as belonging to the
generalized type, i. e., it is both widely spreading and deeply pene-
trating, but does not attain extreme development in either direction.

4

— l-

6

— »-

TT"W*>

^•^f h fhkT;; ! »'^„^e-^i""«"s'<^":'l diuKrara illustrating the taproot and the positions of three primary laterals
H^ht. ofTh^^r^r^'^ secondary and tertiary branches. The development bf vertical roots is deSed at thi
right; at the left is shown a lateral of deeper origin obliquing upward; in the background is shown the horf.
zontal development with the topsoil removed. Modeled after a tree about 30 years old!

Insofar as we have knowledge of other pines, this type of root svstem
seems to be typical of the genus {0, 12, 17). It usually shows a definite
and lairly strong taproot which frequently is found to divide, below
a deptli of 2 to 3 feet, into numerous branches descending at acute
angles to the vertical. This feature becomes apparent only after
the tree has passed the sapling stages. The major part of the lateral
system originates from the root crown within 8 inches from the soil
surface. 1 hese primary laterals extend radially at depths of 2 to S
inches, rarely, if ever, turning downward appreciably. Where a
dense mat of ericaceous roots and rhizomes occurs, the pine roots
tend to run below it at depths of 5 to 8 inches; where the ericaceous
mat IS absent or poorly tleveloped, some pine laterals will be found
as close as 2 inches to the soil surface. The primary lateral svstem
IS almost entirely confined to the surface soil during the seedling and

[

1

f

n

1

Dec. 1, 1935

Root Development oj Pitch P'nn

987

sapling stages; later laterals originating as deep as 2 f^e^Ur^li^^
conspicuous size. Deep y originating laterals requei tly tbou< 1 no^
always oblique graduahy upward to the surface so 1^\ 1 ere^t^
proceed horizontally. The primary laterals give o« be Iver r d
Z filr'^'V'^ secondary branches, which rebranch co mon v to
the fifth and sixth orders. The term - vertical ", as here used "i nlies
only to roots grovving do^^nwal•d; those branches which <rw\nn^^^
from the more deeply situated laterals soon turn i ^ X^-h^^'t
direi-t-ion and are not distinguished from those which irrw^^^m^^
zonta ly from the start. The vertical branches 'don7 1 eS^.l ' f"
of a lateral may penetrate almost as deeplv as doos tL V r n
those of more recent origin on the distal halfof the lateralnit r V
are not so well developed (fig. 1). That the tree has contact vih
the subsod under the major part ^t>nuui witn

of the area covered by the lateral
system is a fact that is not general-
ly appreciated and that warrants
emphasis.

Many of the ultimate branches
are mycorrhizal. Lateral branches
of the higher orders frequently ex-
tend into the duff, where mycoj--
rhizal development may be profuse.
However, mycorrhizae are not
confined to the surface soil. In
the pine-barren soils, mvcorrijizae
seem able to develop under j)ra('-
tically all conditions that permit
growth of the pine root itself.
They were observed as deep as 8
feet in drained soils and more than
3 feet below the water table in
saturated soils.

L-i.f i: 7 ' ^"""'"■' "^ -''""* '■'«"'^« seedlings
taken in June from open woods. The leriL-i h of
the longest taproot was 5.5 inches

f ROOT DEVELOPMENT IN EARLIER STAGES OF GROWTH

SEEDLING ROOT SYSTEMS

systems'^TnLL^ "'? ""°n- '•«'!sP'cuons feature „f seedling ,o„t
systems raproots of seedlings m the r Hist year of growth varie,
in leng h from 3 inches to more than 1 foot (Hg. 2) The inl ,m nee
on root development of local differences in site is pro ha IK, e
clearly seen in young seedlings than at any later stage Te n xi
mum elongation of roots occurs on areas of clean, loose sand subjected
or traTlsViH^n ?pn°"- . ?'^ ^^ ^'"?'', "' "'frequently used roX y

The coinbitdLct^i ''f '7'' >"?•".'''' .'•^"Ptional root development,
ine combined lactors of shade, higher hiinuis content of the soil -ind

underground conipetition tend to retard root elonga ion o ;; "

mgs growing under the tree canopy. Tnder these condition* W

taproots sometimes penetrate scurcdV deeper thafthrouS he lavei-s

of duff and raw humus during their first year of growth Buri " f]

has shown experimentally that white pine seedlings produce the

strongest root systems when grown without shade, tl'e root "vstem

4.

988

Journal oj Agricultural Research

Vol. 61, no. 11

Dec. 1, 1935

becoming progressively weaker as shade is increased. Top develop-
ment, also, was markedly retarded in the plants grown in full shade.
In the relatively open woods of the pine barrens, conditions of full
shade rarely occur. Hence, top development gives little indication
of root development in the case of young pitch pine seedlings in nature.
By the end of the first year, recognizable lateral roots have ap-
peared along the upper 2 inches of taproot (fig. 3). Although the
strongest 2 or 3 laterals on well-developed root systems may be
as long as 6 inches, most of them do not exceed 3 inches in length.
Usually not more than 5 or 6 branches can definitely be identified as
potential laterals on the 1-year-old seedling. At this time branching
has proceeded to the second and occasionally the third order.

Root Development of Pitch Pine

Figure 3.— Siiecimens of Pinus rigida seedlings taken in August from an exposed roadside. The taproots

were about 8 inches long.

Mycorrhizae were already evident in June on the roots of seedlings
of the current season. Such seedlings could not have been much
more than 2 months old. Some apparently healthy seedhngs were
found on which the whole root system was sheathed in fungus my-
celium. Mycorrhizae were present in abundance on 1 -year-old
seedlings, with some clusters showing 4 or 5 dichotomies.

The taproot may be regarded as dominating the root system up to
the eighth or tenth year of the plant's fife, though after the first year
its dominance steadily decreases. Seedhngs 4 or 5 years old, growing
under conditions of moderate competition and partial shade on Lake-
wood sand, showed taproot penetrations of 15 to 24 inches and
maximum lateral extensions of 12 to 15 inches (fig. 4). Usually there

*

I

f
1^

989

spmdhng, decumbent or, at
best, only feebly erect.
The seedlings of both Pinus
rigida and P. echinata usu-
ally topple over during the
second or third year, and
remain in a semiprostrate
position for several seasons.
By the eighth year they
usually have again assumed
an erect position, but a per-
manent crook remains at
the ground fine. This crook
is ultimately obscured by
the thickening of the stem,
but frequently can still be
seen in saplings 8 or 10 feet
tall.

coSS^ \!r, is stdiitr lit ?5 tiHa^^ "-."^-'^

beginning to branch. Though plants of thi.Llf ' ""'^ "F®
fairly erect position, the st.^ s^rtk^i^it^'^ ^IZ^^^TA^.

Figure ^--Four-year-old seedlings of Pinus rigida with prao-
tically complete root systems. '"'Prao-

F.GURE 5.-Eight-year-old seedlings of Pinus rigida with practicaUy complete root systems

^iTum^'' TT.''''^r'^'^"H' increases with age, both above and below
ground. Taproot penetrations varying from n to '^4 \nrhc^J^^^Z

approximated 2 feet in length, though one was found which measured

Jf

990

Journal of Agricultural Research

Vol. 51, no. 11

'Dec. 1, 1935

Root Development of Fitch Pine

40 inches. The number of identifiable laterals averaged 10 or more
borne of the stronger ones bore secondary branches almost 1 foot long
hranchmg had proceeded commonly to the third and occasionally
to the fourth order, exclusive of mycorrhizae.

Some comparison was made with seedlings of Pinust echinata of the
same age and on the same sites. The development of the two species
was very siimlar; no difl'erences were noted which did not readily fall
withm the hmits of variation within each species. It is the writer's
opmion that detailed analysis of many specimens will be necessary to
determme any differences which may exist between the seedlimr root
systems of the two species. However, it was noted that the stems of
seedhngs of P. echinata tend to stand more rigidly erect than do those
01 P. rigida,

ROOT SYSTEMS OF SAPLINGS AND SMALL TREES

By the eleventh or twelfth year the pitch pine seedling has acciuired
a clegree of rigidity heretofore lacking, tlie growth rate has increaseil,
giving an aspect of vigor, and annual whoris of branches are beino'
regulariy developed. The young plant at this age begins to appear like
a tree Ihis acceleration of growth undoubtedly is coordinated with
root development; it could not take place until an adequate absorbinc^
system had been formed and sufficient contact made with the subsoil to
insure a constant water supply. The dimensions attained by the roots
indicate an even more striking acceleration of growth underground
than that displayed by the tops. The eighth to tenth years mark
the inception of a period of rapid root elongation. This period also
witnesses the appearance of seconchirv branches of conspicuous size

It should be borne in mind that, although it is convenient to speak
ol these younger stages in terms of years, age probably is relatively
insignihcant. The development of the plant is more properiv to be
viewed as aii orderly process, the speed of which is governed ,%vithin
limits, by the environmental conditions. Thus, a stage of develop-
ment ascribed to 10-year-old trees in New Jersey might be reached
several years earher, or later, under other conditions

A fairly typical sapling, 1 1 or 12 years of age, was 4 feet tall, and
0.7o inch in diameter at the ground line. The current season ^s leader
was 8.5 inches long, and that of the past season, 7 inches The tap-
root reached a depth of 4 feet, the longest lateral root reached out S
eet, and 3 other laterals exceeded 4 feet in length. Ten major
laterals had originated within 6 inches from the surface of the -^oil
and o more were present at greater depths. The longest lateral
bore 12 secondary branches ranging from 1 to 2.5 feet in length The
strongest secondary branch descended vertically from one of the other
pnmary laterals to a depth of 3.5 feet.

The appearance of greater numbers of branches, both primary and
secondary as the seedling grows older does not mean that new
branches have originated at points remote from the tip of the parent
root. As vyill be discussed later, branch roots normally originate only
in tip regions before the advent of secondary thickening. Many
branches never become more than a few inches long, and eventually
die and disintegrate. Certain ones, however, which at first are un-
hJtnTfpntn from th ephemerais, later undergo increased growth
s^rittnrir^Th^'"'^ thickness and become recognizable as permanent
structures These account for the apparent increase in the number
oi major branches.

b- -.

, «

W

J

i

I

i

991

brandies. The tSr of com4 re^ "^ PT""-^ '"'^'•^'

organ of central anchoraee ann= tl,. 1 ? *^ importance as an
center of the root system It, L\tl "«<= '^"'^f ' """"^ Pl'/^iological
which all liquids must nass Tt^Tn ^'^ '' "'^ "?"<'"■' t'"-o"gh
however, becomes more Td more inskn'ificant T M^'^'^'T "'P"'
\ertica secondary branches or »fnW^ ? /^ *'^^ ^^^^ develops,
relieve the taproot oTksearhervifT; ?"'?'"* *'" '"''^°" ''"d th"«
bv drought. Ultimatelv ! ,„l! lole-insurance against death

bmnchesfmaynoT^Sevce.-r?r'; """^^^'^S ^'^ descending
to the amounfof abforbing area ''"^"' '"'''"'^ ""'^^ '^'P'"'^

th;\rp?:otntX ^S^^'^^^^^^i:^^^ r '"^ of

^ atKsi^t? tiTefir^- ^}^'Si:'tTc

eKLn^ons^Vt'LflaS^t^^^^^ observer can predict the gen-
by ring count, 7 5 feet talT nn.^ r -i^... ^ 1?''"^ "*' '^ y*^*"^ o'd
line. ;The taproot reached a depth "of "nly f^Zf'','' '*'« «T"o^

fengLVolSsSrotlef^ I^»ot^!;t iLl^^nlfs"
while th™Sr rTgll ?rom > io sleeT T*^"^"'' ^ ^"'' ^" '«"g*'''
than 2 feet were not cSeml TuJoi ^"T.'^ '«'?■■«'« shorter
about 30 seconda";' bZfc ap^ /n^oT: ce^d^^^ TVtZ

saariL7eaSdTLx^s,yoTob. tl ^ ^-^^

&^5"mtChi.^-"- "-' I— ^edt'tSiXoS, S

were dead, or apparently dormant, and del3even?ualS to sil^l

loot sj stem. A branch which has attained a lens-th of i f««* =
growmg, or previously has grown, more vigorous^t^ ts ba al nar s

been ZtZZTir'^'f'^:- ''t'f'T^' ''■>'• «>« epidermi ' ha
oeen leplaced by the characteristic flaky bark, eivine the nsnert nf «

permanent organ. A reasonably close correla ion exits between tho

caTb^eTtimTted"titLo.'ft"^"" "/ T" ^°°*^- ThS t&LVtt
^uii ue esumated without complete exposure Branrhino- tn fi,^

KrtSo'nel.*'""^''*'"'''^- "'"''' •'^•'^"- "" latemlbranThe's^tht

primafv eSt^, ^r" f "f """*''": '" '"creasing numbers as the
ft ma7hpSt-40^iSre: ? f-^S o^g'^^T^.^allVrS

o nethlu"rre7er£f ""%' V"=^°"'"'' ""'.' ^''^-aiSoneX'th

from a roof in t«„ V ^^''*' '^ Pnmaiy lateral thus has developed

tL„^ 1 '"*o a root system in itse f. Its zone of absorotion ev

late^llt"'"" ^, the.moister sands of the subsoil! and sevemlfeet

'^yr :"''"■ '"^' '" "'« *°P^°"- Subsequcnt'growth of Ihe pri-

992

Journal oj Agricultural Research

Vol 51, no. 11

Mil 's simp y the extension, verticaUy, laterally, and linearly

of this zone ol root occupancy. However, downwarl^ growth does
not keep equal pace with the lateral and linear advance. Se the
ffv. n^f "■ °'"="Pr'^y,°f "^ P""''''-y I'^teral undergoes a grfdualrela!
inl nrnn^'J-"^ as development progresses, and a 1;onstantfy "nereas-

sufface'fnn t/ ^'.™''* Y'*"" *' ^ ^''"l" ^^^^^^ localized fn the
surface soil. The data on larger trees illustrate this tendency

A specimen tree, 14 feet tall, 2.5 inches d. b h and about ^^' vpnr=
old, may be cited. This individual had originated as a sprout" and

Krfrom t'jd""s' -f ^hf '''^"^ '"' ''' 1^' than tre:sXvolo;i"g
airectiy trom seed. Suitable specimens of seedling oriein of tlii«
general size were almost enth-ely lacking in the locality Tl,» y^L !
was rather atypical. It divided about'^O indtes bdot tlil uSof
the soil the two roots subsequently uniting in a naTuraJ CTa/rnnd
then redividing. One branch ended at a depth of 3 feet Tlie other
after forking twice more, reached a depth of sligldy more than ? fePt
Twenty-two primary laterals had originated at depths of less than i
foot, and several small ones were found at deener levek %n?,l lit .
exceeded 15 feet in length, the longest beine 19 feet A fiff i ^^^^^^
ured 12 feet; the other! were of°S.s sKrtngtht ''t hHumW
of secondaiy branches varied from 40 to 75 on thi 4 largest iSls
Horizontal secondaries as long as 6 feet were fnnnY <,t , u- . •
tertiaiy branches of conspicuous size weTe present ' None' o'f "Z
vertical secondaries or sinkers, reached de^erTan 5 feS ""

nnl^^ r^ ^f"^^^^ sequence of development is followed bv the sec
m±7 ,''5*-"''''f ""^ >y primaries. Their early growth is a^elv
mamfested in elongation, most of the side branchefS smaKi
icate, and ephemeral. When a length of 4 or 5 feTLs hef n Tt f L.J"
permanent branches of the next higher order firs become coZlJ

oTstill ddtrrooT^yrr "'''' "^ '^P^'^*^^ '''"^ »>>' *-««^ ^^^^^

wJtSrdevdiXirtin4 '^P*^-"^ ' ''''■■ ^^S>pT;.rt '
soilsurfaclLSShtiSX^n^el^S^^^^

1

Dec. 1, 1935

Root Development of Pitch Pine

993

LSnrsi 5rt1i.!i t^i^'twrfs ^of'Te"^"'^^ "^i^^^

branches chstributed within a c^X^e: ibout 2^1* i'lLmS
Afig- 6). Most of them did not extend deenpr tli«n 7 w i diameter

longest ones reached depths of 8.5 anS 9 S ' GrowLrt Ltei^'fa h??

^ abundant, and active mycorrhizae were collected as deenar? w/^

There were 20 sizable laterals, 13 of which originated' at deSs of

less than 6 inches; the other 7 were found nt o^moto!. ,i A '^'v'""* ^^

the 2.foot level. The longest laTLl ml^ rf^l" 3 wl? t ot '4

count were estimated
to be about 8 feet
long.

A few, of still smaller
size, were not recorded.
Secondary branches
were not extensively
excavated. One of
the larger secondary
laterals was 9 feet long,
another was exposed
for 11 feet without
recovering the tip ;
several others were
estimated to be of
equal, or greater,
length. One sinker
was observed to end
about 5.5 feet deep,
but others probably
reached deeper. The
largest sinkers gener-
ally may be expected
to approximate the
taproot in depth.

On any tree second-
ary branches that can
be classified as large
in proportion to all
secondaries are rela-
tively few. The ma-
jority of them are

mo^^ rfnl T** '",??••?"?. 1' IW in group 2, and 222 in group 3 The
cZsed li'^groun"'/"^,^""'''^' "T 'Z ««<^«."daries, of^XSi 6 Jere
prima^ii^rXka^d^o-sSa^ierf^lllV^nfr,!;,^!. '''^ -'«"-

** T ^ "fl? T^^P""^^ exposure of a 30-year-old tree. The excavation «•««

994

Journal oj Agricultural Research

Vol. 51, no. II

GENERAL CONCLUSIONS FROM STUDY OF ROOT SYSTEiSs IN EARLY STAGES

The plan of this paper thus far has been to present general mor-
phological data from the developmental point of view. The growth

ooniffiZ f?''*^ ''?' ''"? ^""""""^ ^™'» tl^« ^««dUng stages to tAe
conditions found m immature trees 20 to 25 feet tall Subsequent

^W?It :ilffl?r;i- ' "' •""^t'-«ted by larger trees, proceeds along some-
what different Ines. Hence, certain generalizations applicable to
the earlier growth stages are now presented.

Proportions of Vertical and Horizontal .Ikcondary Bra.\che.s

A summation of data with respect to the proportions of vertical
and horizontal secondary branches is given in table 1 Since the
horizontal category includes branches on botii lateral sides of the
Z'e?IlT*' l^ '' «"dent that the primary puts out braLcl^s in hree

flTr!! of r'*.'?i' ""^u ''"^ *'"^ ''"'^^'"^ represent appro.ximately one-
third of the tota This three-way distribution of branches cannot
be correlated with the anatomy of the protostele. Although the
prnnary xylem is not infrequently triarch in the roots of pUch pine
the diarch condition is most common. ^ '

Increasing Prominen(e of the Lateral Svste.m

Mention has been made above of the increasing prominence of the
lateral system of roots as the tree develops. In the pine barrens

he iXr o?""/.'-'' '" ""', ^T^'f »"'' '»»«* conspicuo,^ root up to
the eighth to tenth years of the plant's life. Soon after that time the
s rongest laterals linearly outstrip the taproot, and eventuairneariv
all of them e.xceed the taproot in length. Mention also lias been
made of the tendency of the root system to flatten out and become
re «tively more localized in the topsoil as deveoprnt progresses
This change comes about largely through the more ranid Growth of
the primary laterals as compared with tlfat of the rproo"^^ „„!l s bk^^^

It ha^;eSiS -leS ^o1 7^7^:X '""' """ ''^-^^=^-

Table 1. — The

proportions of vertical and horizontal secondary branches of three

pitch pines ''

Tree no.

Primary
roots
tabu-
lated

Secondary roots

1 I.
3«.

Nu mber
3
9
6

Total

Vertical

Horizontal

f

NumbfT
80

284

X umber
37

7.5

Percent
46
34
26

\ A° """^V?'^' ^"'^«'"'>"s lO-year-oM sapling.
, I'-year-o d free cited in te.vt.
* JOyear-old tree cited in text.

Xii mber
43
190
2aSi

Percent

54
6<i
74

.iJSii.^s^„'fct°Li„tisrr<,JS:,i-=f

I

Dec. ], 1935

Root Development oj Pitch Pine

995

S r a™hSLre''dso"uS& ^'^

in that column ProL^^t^SS^SSotrot^nt t^^^ S iT

Height of tree

Age of
tree

1.5.

3.0-
11-
22..

Inches

2.7.
4...
4.5.

Feet

14..
22.5.

Months
2-3

y'ears
1
4

8

10
12
11
10
14
17
30

Depth of
taproot

Depth of

strongest

sinkers

Inches

8
18
27

Feet

Length of
longest

primary
lateral

Feet
3.
4
2.

3"
3
5
9

' No data.

1

3.

3

3

4

5

Inches
1.5

3.0
15
33

Feet
5.5

8
12
14

10.5 I
19 I
31

Primary

laterals

longer

than

taproot

Number

Ratio of
longest
primary
lateral to
taproot

(')

3
5
13
12
12
12
17

0.3

.4

.8

1.2

1.7
2.0
4.8
4.7
3.5
3.8
3.4

Length of
longest

secondary
laterals

Ratio of
secondary

laterals
to sinkers

Feet

1

2.5
4

2.5
3.5
6
12

1.0

. I

1.3

.8

.9

1.2

1.7

Relatio.x op Lateral Spread of Roots to Height of Tree

of such a criterion must be recoo^nizprl Rn^^ / • u ^^n^^^^tions

tions and variations dul 1^ trelv^^rlne^ aT^m^ Lf cl^^

tions of root development are impossible when based oAlv on nW-]"

It IS even more difficult to estimate the depth of taoroots TI.»
figures in the first and third columns of table 2 indicate hat T e
stem and taproot attain hnear equality somewhere be ween -'an 1 4

knee fh!lT'^'-' *'r '"P'r* ""* '°°Ser; after the interval of eni^va-
lence, the stem is always longer. Nine feet appears to be nen flfa
maximum depth attained by pitch pine taprootsb Hie localiu"

•l-AHi.E 3.~Da.a .houing ihc relation of the maximum lateral .pread of roots to

n eight of tree

Height
of tree

Inch IS
11
II
18
18
20

F(d
2.5
2.8

Laterals
exceed-
ing the
stem in
length

Xiimbfr

Length

of longest

la t end

Inchfs
13
15
2»i
27
40

F»,1
4.7
.5.3

Ratio of

Laterals

longest

lateral to

height

Height
of tree

exceed-
ing the
stem in

j of tree

length

! 1

F((t

Xumber

1.2

2.8

4

1.4

4.0

4

1.4

4.0

4

1.5

7.5

O

2.0

7.0

3

7.5

4

1.9

14.0

4

1.9

t

22.5

6

Length

of longest

lateral

Feet

6.3

8.0

12.0

y.o

14.0
10.5
19.0
lil.O

Ratio of

longest

lateral to

height

of tree

2.3

2.0

2.7

1.2

2.0

1.4

1.4

1.4

996

Journal of Agricultural Research ^

Vol, 51, no. 11

Dec. 1, 1935

Relation Between Root Diameter and Length

Little has been said in the preceding discussion concerning the
diameters of roots. From a physiological standpoint, diameters are
of far less significance than lengths of roots and volume of soil oc-
cupied. Basal diameters, particularly those of horizontal roots, are
roughly proportional to lengths, and can be used to some advantage
in estimating the lengths of laterals. In cases where there is appre-
ciable enlargement at the point of origin, measurement should be
taken beyond the region of pronounced tapering. For a general
estimate of the length of a root, the basal diameter is measured in
milhmeters; the length expectancy is an equivalent number of feet
l^or example, a root of 5 ram in diameter at the base is about 5 feet
{^ng; a root 30 mm m diameter at the base is about 30 feet lono-
JNaturally, there are rather wide variations, with a somewhat greater
tendency of lengths to fall short of, rather than to exceed, the expec-

Root Development of Pitch Pine

''"'''" JfUie'laS ''^Kit.ulZlKT^'"', ''^^ l'->'^«'--c,ld pilch pine showing the ropelike character
or ine laterals. The tools mark the ends of roots. The root on the right was 19 feet long.

tancy according to this simple formula. The length is generallv
mversely proportional to the degree of branching, i. e., the parent
root IS shorter when well branched than when poorlv branched.
1 he diameters m millimeters and the lengths in feet weVe tabulated
tor 53 primary laterals from seven trees. The ratio of the lencrth
hgure to the diameter figure was calculated for each root The
average of these ratios was 0.85. This expresses the tendency,
noted above, of lengths to fall slightly short of the simple ratio of 1 0

Jt can be seen that these lateral roots are rather slender structures
femce branches usually are distinctly smaller than the parent root"
and on older roots are relatively sparsely distributed, the largei^
lateials may aptly be described as "ropelike" in appearance (fie 7)

During the younger stages sinker roots maintain the same propor-
tions as laterals, but after they reach a depth of 3 or 4 feet, elongation

I

997

no reliable indication of its length diameter of a taproot furnishes
ROOT DEVELOPMENT OF TREES APPROACHING MATURITY

Whit SeSiS^fS; zi SrSrLrr^ ^r --
:S^rfe!irfe:^^dLfe^^^^

spread of roots, 30 to 35 feet- and n^p\n .r!! ^^' f ^^^^^^l radial
ment involves 'a rather stTady^n^^^^^^^ W ]^f develop-

vertically, during which the ropelike character oH.^^^^^^^ ""^^

, tamed without marked thickenincfnrit? ^ . ^ laterals is main-

' tions exist between extent o^^^^^^ general correla-

diameters and lengths of rootV 41 "^^f^^^^^^Ps and between

nothing to the raTat^^^^ ^T^^ ^'^^'A''^^ or

M), in a footnote, indicate that a s^?nilYr se^uen^p'ff'')^''"'^^ ^"^"^
has been observed in ^""uar sequence of development

Europe (probably on

Pinus sylvestris).

Instead of further

elongation, a marked

thickening becomes

apparent along the

basal 1 to 3 feet of the

stronger primary lat-
erals. The smaller

primaries, and to some
extent, the secondary
laterals, continue to
elongate, thus increas-
ing thedensity of roots
within the 30-foot
radius. Ultimately,
the originally smaller
primaries attain
lengths approxima tely

as great as those of the
strongest laterals;
they then cease elon-
gating and undergo
basal thickening.
Further penetration

of the subsoil by the . .^.

taproot and the stronjr figure s.-The central ront« nt .. r^ . TT ~~

of tko ioSemoi, ScS ^Et , E. T''"'«° " "l'"" »"' ''J' """^

998

Journal of Agricultural Research "

Vol. 51. no. 11

)ec. 1, 1935

Root Development oj Pitch Pine

The basal thickening of the primaries very frequently is morp nm
nounced in the vertical plane, resulting in ^'plankhke- roX Th;
asymmetry rarely extends outward more than 3 feet UnToubtedlv
It is a mechanically efficient means of bracing the tree «P-Ain«7 Ik.
increased stresses of a larger crown. Rigg and Harrar (22 v\^^
noted an extreme development of this tendency S the shallow ^oo
systems of trees growing in sphagnum peat, and, apropos of the mp
chanical efficiency of such modifications, stke that^Tsti^^^^^^^^
beams of varying transverse sectional shapes approximatL tSu«r^^
oUhe vertical dimension when the horizo^ntal ^dLien^™^^^^^^^

These plankhke roots frequently are decidedly eccentric- that i^
the morphological center of the root is below the^actual center, La

result of greater thick-
ening on the top than
on the under side (fig.
0). Thus, shallowly
situated laterals ofteii
grow out of the ground
near the stem base,
producing the familiar
buttressed effect.
Although many decid-
uous trees display this
feature more m'^ark-
edly, it shows plainly
in pitch pine and prob-
ably is a wide-spread
habit. Laitakari {17)
mentions it in discuss-
ing the roots of Pinn.^
■^ylrestris. Vertically
narrowed roots and
buttresses attain strik- ,
ing dimensions in some
tropical trees. Obvi-
ously, buttresses are *
mechanically eflBcient
structures; the same
amount of material
would give greater
support in that posi- ^
tion than in the strictly j
horizontal position o'f f
the original root. In-
vestigations of tropi-

^''the^aDrooV'^'rift'S"''' ''^^'"''^ ^""^ '«^«^«' •'oois cut dose to the

oal ti-ees indicate, however, that buttresses cannotlfways^be explS
as a s mple response to mechanical stress. They may develon when

nfl^ttiaH^^^^^^^ j'' -^! -«^- relatioLtpTear To'b

innuentiai lactors (7, 9). The formation of the oliaracteristir hut

,hnwn%°^*^''>'"''''j=yP'"''^^ (r«xorfi«m distichum (L. rIi, ) has been
The tu'J'F^'^^r' r " co'^Wnation of water plus ah-' ( V)
the buttresslike development of nteral roots of mature ererf trp«-

suggests that the roots function mechanically as props' or braces!

999

^ "riSySucSftL'g^o^!^^

V - roots also function mechanicX^/?r-l Undoubtedly, supporting
.1 is minor and is secondary to the nrll''^ P^ ''T^' b»' *»'»' role

) pointed out the tendencTo^ScotcS^ ^''''''^'''i ('^) ^as

J laterals on the side opnosite th«t Atw?. P'^^^Hce more and stronger

^ propping position To check thk to ' ir^'*" '"^u^'"^^' '• « - '« the
leaning trees were examined Tht *f"''''."fy ^U'ther m pitch pine,

! such individuals k dkUnct and constant"" Of f ''^ ?" "'^ ™''*^ "^
i 3 showed a very marked conPontT,- . PI ^ ^"""^ ^""^^^ examined,
the inclined tr^n^ ^th Z!i^awZ(tlTf''\'''' **'«, ^'^^ ""^^r
side (fig. 10). This indfcates ea^fy t^arthose rooLIn ft ""^"^''^
position are thickened in response'totr ssT'th^tin" hel[°yS

i

dTand dtappear'"'''"' '"' """' ^'^'^^'^ ^^^^ under the strain,
relatively srirgbches d.*b h lTlfiel'i''Jf%r'''''' T^

was not given particular attention in this investigation Hnw^it^
ioT^T'" '""'""' "'"'"' ''''" alf rXr weak slow^gT^dng

998

Journal oj Agricultural Research

Nol. "il. no. 11

Dec. 1, 1935

The basal thickening of the primaries very frequently is more nrn
nounced m the yertical plane, resulting in -planE^ Vots The"
asymmetry rarely extends outward mo^ than 3 fee^ Und^ibtedlT
It IS a mechanically eflicient means of bracing the tree aSth\;
increased stresses of a larger croNvn. Rigg and Ilarrai C^^Tvm
noted an extreme deyelopment of this temlency n [he shallow roo
systems oi^es growing in sphagnum peat, and, apropof o^^^^^^
chanical elhciency of such modifications, stite that 'X^tre Ith of
beams ol yarying transyerse sectional shapes appr<>xima L the^^^^^
ol^the yertical chmension when the horiZtnlluo^^n IZ^

These phinkhke roots frequently are decidedly eccentric- th.t is
the morphological center of the root is below the actual center La

result of greater thick-
oning on the top than
on the under side (fig.
0^. Thus, shallowly
situated laterals often
grow out of the ground
near the stem base,
producing the familiar
bu t tressed efiect.
Although many decid-
uous trees display this
feature more niark-
t'clly, it shows plainly
in pitch pine and prob-
ably is a wide-spread
habit. Laitakari (17)
mentions it in discuss-
ing the roots of Phnis
si/frestris. Vertically
narrowed roots anil
buttresses attain strik-
ing dimensions in some
tropical trees. Obyi-
ously, buttresses are
mechanically efilcient
structures; ^the same
amount of material
would giye greater
support in that posi-
K,. ,HK.. rr . , tion than in the strictly

^^^i^'>^^^^^^^^^^^ Imrizontal position of

t«o lower ones show ec^n.ricthUkerunK on (he u,/,^^^^^^^^ th(> original root. Ill-

cnl trees indicates howeyer, that buttresses canm!f K^e exnlliTf
as a simple response to mechanical stress Tl ey n a vfh '

T . I I'' '''■,P^""'"^"" 'I co.nl.ination of wafr plus a\v(w')
s.iIo«, T 1''' 'l"^<^'»P"'<'nt „f hnornl roots of n a n'e e r'^ t trees
suggests that tl.c roots function .Meehanieully as props' or br.nees

Ji"ot Development t,j Pitch Pine

999

• and that the tliickenine develons i>. ,);..„„.
stresses produced by the sro X c,- ''.^fPonse to the increasing

- roots also function niccimniMlv^as n -l ^-"^o" '"''•l.v. supporting
is minor and is secon(arvZtl,en,l •''''' ""^ ""'''"• l"" "'«' ''ole
pointed out the tended o^Scotdn^^^^^^^ '"'"T' '""'"'^'"•' "') '""^
laterals on the side opposite thti^ •"'"•""' '"•""' ""*' ""■""^'"•

propping position '^1,0,1/1,'' '"^ I»''^-n""!-Vvin,ls, i. e., in The
leaning M'ees were ex , mine, T '^''"'''.»fy, H'ltlier in pitch pine

such iudivi.lualsiLi"i ril coZZT'V f'''! r" ""^ ■•»°'^ "'
3 showed a very nnnk.>.l c, nrenn ,r . ^ ^ H'""'' *'''^''s oxaniined,
the inclined tr.mk \ no ^'^ h,":;!',,"' '"^/■"■^"n the si.le unde,:
side (fig. 10). Thi; in ^at^s cl,:^ 1 . H r;r '"'''™'^ •^" V" "W"*'''"
portion are thickene., in resisi\^''.^:,.^^rt.r ^in" ^

propping

Sits dtppear'"'"'' '''' '""^^ ^"-'''^ ^'-^ under the strain,
weaken this conchision A " *""' ''°°,^ development, di<l not

INTENTIONAL SECOND EXPOSURE

1000

Journal of Agricultural Research

Vo». 61, no. 11

specimens. The number and size of the primary laterals and the
thickness of the taproot were distinctly inferior in comparison with
those of normal trees of the same size. In view of the open character
of the forest, it is doubtful that the weakness of the specimens was
due to suppression by larger trees. It seems most probable that both
the leaning and the general weakness of the tops had resulted from,
or were coincident with, an originally weak root system.

It seems quite probable that the processes of maturing and senes-
cence are correlated with the tree's inability to extend the root system
into new territory. It is obvious that the demands for support placed
upon the inner root system increase enormously as the top attains
mature stature. The material requirements for strengthening near
the root crown may be an important factor in precluding further root
extensions. In any event, a fertile field for speculation is offered by
the question of whether maturing and senescence are wholly internal
and protoplasmic, with the cessation of root extension as a result, or
whether the mechanical difficulties of support and of transportation
through an ever-widening root system are contributory causes of
those processes.

ORIGIN OF ROOT BRANCHES

The endogenous origin of the branches of roots is one of the funda-
mental facts of plant anatomy. Normally, branches emerge along
the maturation zone of the root tip, and along the tender parts farther
back which have not undergone marked secondary thickening. The
completion of a solid ring of secondary wood generally precludes
further branching. On an actively growing root, this probably takes
place during the second year; on roots gi owing less vigorously, either
of two types of behavior may occur: (1) Secondary thickening may
be indefinitely delayed, and the period of possible branching equally
prolonged; (2) secondary thickening may advance to withm 5 mm
of the tip, resulting in thick, blunt-ended roots with a barelj^ per-
ceptible tip of tender tissue. The former is usually associated with
dormant lateral roots; the latter condition is most often found in
vertical roots deep in the subsoil.

The growing tips of the major extending roots are thick and succu-
lent, about 3 mm in diameter. The tips of branches at emergence
usually are distinctly smaller. In cases where the parent tip has
died (a surprisingly common occurrence), a replacement tip emerges
which, from the start, has the attributes of the parent root. It
resembles the latter in size and vigor of growth, and at once assumes
a forward rather than a lateral direction (fig. 11). Sometimes two
such tips emerge opposite each other; the immediate result, while the
dead parent tip persists, is an apparent trif urcation ; ultimately the
dead root disintegrates, leaving a bifurcate condition.

The formation of replacement tips was not observed to take place
when dying-back had extended into the region of complete secondary
thickening. In that event, an existent branch may undergo greater
than average growth and after the disintegration of the dead parts
appear to be the normal continuation of the parent root. This is
nearly always what has happened in those occasional cases in which
a major lateral root appears to become vertical, or vice versa. The
abrupt change of direction marks the origin of a branch which assumed
dominance after the death of the parent root beyond that point.

Dec. 1, 1935

Root Developjuent oj Pitch Pine

1001

>-.:

^(

The roots of many herbaceous plants, which do not undergo exten-
sive thickening, may produce branches at any point. Moisture fre-
quently stimulates abundant branch production on such roots in
regions far removed from the growing tip. The fact that the woody
root, in some species at least, is only slightly or not at all capable of
such response warrants considerable emphasis, particularly in respect
to the taproot. With such species, of wliich pitch pine appears to
be one, it means that the original complement of primary branches on
the seedling taproot never can be increased in later life. It is highly
probable that an originally poorly branched taproot, or one deprived
of its branches in any way, will re-
sult in a weak root system and pre-
dispose toward general weakness of
the tree. The best cultural care
is, to some extent, wasted on a tree
with an inadequate root system.
Of course, when the number is re-
duced, roots may become somewhat
larger and branch more profusely
than usual, but it is doubtful that
such development can be fully com-
pensatory in respect to absorbing
area. Certainly, reduction in the
number of primary laterals weakens
the support of the tree and predis-
poses toward wind throw in later life.

The above statements do not
appl^ with equal force to all tree
species. Adventitious roots, aris-
ing both from stems and from other
roots, are mentioned occasionallv
in the literature. Luncz {19)^ m
reviewing some European work,
cites Austrian pine and cherry as
examples of trees whose roots may
give rise to adventitious branches,
and states that such branches are
not morphologically distinguish-
able from the original roots. If
adventitious roots ever arise on
either pitch or shortleaf pines,
they must require a more powerful
stimulus than normally occurs in
nature. Examination of scores of
roots showed that branches of all sizes, on both species, universally
extended inward to the protoxylem of the parent root, and therefore
had originated when that region was young (fig. 12). The only pos-
sible exception found was in cases where the roots had been severed
smoothly >vith a sharp instrument. Some small branches had subse-
quently arisen exogenously near the cut ends. Examples of this
were found at the scene of stump-grubbing operations of the preceding
winter. No observations were made of exogenous branches of
greater age.

Figure 11.— a young replacement tip developing
after the death of the terminal portion of the
parent root. Note its relative size and direction
of growth in comparison with the remnants of
normal side branches.

1002

Journal of Agricultural Research

Vol. 61, uo. 11

Dec 1 . 1935

Root Development of Pitch Pine

1003

GROWTH OF ROOTS UNDER WATER

Probably the most striking single feature of pitch pine roots
observed in the course of this investigation is the fact that they
develop extensively below the water table in saturated soils. So far
as the writer is aware, no similar observation on a mesophytic tree

has been reported.
Such hydric species as
Taxodium distichum un-
doubtedly root under
water, but in practically
all descriptions of the
roots of mesic trees, and
even of such bog inhabi-
tants as American larch
and black spruce, they
are reported to develop
entirely above the satu-
rated zone. Textbooks
generally state that
root penetration ceases
when the water table
is encountered (4, 24),
and much has been
written of the delete-
rious effects of poor
drainage on various
economic plants. Rigg
and Harrar {22) found
that none of the six
conifers growing in the
bogs of Washington
extended roots below
water. According to
Woodroof (30), the roots
of the pecan cannot
tolerate submergence .
The longleaf pine
{Pinus palustris Mill.),
which in some respects
is the southern ecological
equivalent of pitch pine, is
reported by Hey ward (12)
not to penetrate below
the water table on poorly
drained sites. However,
Hesselman (11) notes the
occurrence of vigorously growing spruce stands in Sweden in the
vicinity of springs, even where the water level comes to the surface.
Although no data are given on root development, the trees obviously
do produce roots under water in such situations. Adamson (1)
describes an Indian tree {Terminalia arjuna Bedd.) which inhabits

Figure 12.— Cross-sectional views of roots showing the central
origin of branches: A. A section from a taproot. 5.5 inches in
diameter: B, section from a primary lateral, illustrating the
origm of a very small branch, show n about natural size.

river banks and extends roots out into the saturated soil of the

river bed. • ^i • j t \

The root behavior of white cedar {Chamaecypans thyoides (L..)
Britton, Sterns, and Poggenberg), which is the typical bog tree of
southern New Jersey, has not been investigated, but in view ot the
sites supporting it, it probably does root below the water level
Pitch pine, however, is most at home on higher, well-drained sites, and
intermingles only slightly with the white cedar in the narrow dividing
ecotone. The pine can live among the cedars only by overtopping
them; once the cedar canopy closes, any pine beneath it is doomed.
The occasional pine that towers among the cedars points clearly to
the fact that it is not the site itself, but the competition, that generally

excludes them. i i *

Pitch pine is to be regarded as a member of that wide-spread plant
fraternity which, demanding little except a place in the sun, must find
its place as a rule in the left-over, unfavorable areas that will not
support the '^nobler" species. Such are the sterile, fire-scorched
sands of the Coastal Plain, or the wind-swept ridges of the mountains,
both of which are typical pitch pine sites. It invades burned-over
areas, abandoned fields, and clearings, where indmduals may persist,
but where, if conditions are favorable, reproduction soon becomes
impossible in competition with other species. Although its tolerance
of competition is low, its tolerance of a wide range of site factors is
remarkable. Sandy soils or clay, fertile or infertile, drained or
saturated, situations xeric or mesic or hydric— all are acceptable to
this undiscriminating species. ^i i i ^i ^

Three trees were examined with respect to root growth below the
water table.« The largest one was 33 feet tall and 5 inche^s d. b. i.
It was located on a steep incline that bounded the valley of a creek.
The water table was bout 3 feet below the surface at the spot where
the tree stood. Lateral roots extended up the inclme into typical
Lakewood sand; on the lower side, they extended into the flat valley
bottom where water was only about 6 inches below the surface, with
sphagnum moss and cinnamon fern {Osmunda cmnnmomea L.) to
testify to the saturated condition. The laterals were of the usual
tvpe, and put down sinkers regardless of the proximity to water (hg
13, A). They were aligned generally parallel with the incline of the
soil surface, both uphill and downhill.. The taproot Penetrated the
zone of saturation as a strong shaft, 4 inches in diameter at the le^ el
of submergence. It broke up into a fan-shaped mass of brancheb
along its fifth foot. This fan, which was about 2 feet wide and 4 to
6 inches thick, was a most peculiar tangle of roots of vanous size,
unlike anything found on drained soils (fig.. 13, B), As a mass it
ended at a depth of 6.5 feet, though a few mdividual roots reached
slightly deeper. Many of the tips were alive and growing sIoa^Iv.
A large proportion of the ultimate fine branches were mycorrhizal,
a^Sn which was quite unexpected. The literature contains

0 The method of securing the ^ntral root ^y^Jef « i'Jfj^f »y,^«^^^^^^ wS

removed around the root crown down o the ^atf/, «";^,JJ^/f„^J'''^ WithThi<f 3 or 4 m^n could pull stumps
Sf ?KeVd^erb^d^.^"p^SSi^ t^r^l^:^^^-^ tX-;^d' was held enmeshed in the root
''n'^:^^nrm^^^.^^C.cT^ll these roots has been verified by K. D. Doak, of the Allegheny Forest
Experiment Station.

1004

Journal of Agricultural Research

Vol. 51, no. 11

Dec. 1, 1935

Root Development of Pitch Pine

1005

Figure 13. — Central roots of trees from saturated soil: A, Preliniiiuirx oxcnvntion of the largest tree, with
water standing in the bottom of the hole. B. Under-wafer portion of the taproot of the same tree. It
reached a depth of 6.5 feel. C, Roots of a smaller tree, fr< in ;> site s* here the water table was only S inches
below the soil surface; these roots reached a depth nf .' Uwi

very few definite statements with respect to the occurrence of mycor-
rhizae in nature on submerged roots. The negative findings of Bondois
(^) constitute the only direct observations on tliis topic known to the
writer. The fact that mycorrhizae had not been reported on sub-
merged roots and that good drainage seems essential for the successful
synthesis of mycorrhizae in culture (21) has led to the general opinion
that they do not develop under saturated conditions. In some re-
spects the presence of mycorrhizae far below a permanent water table
is even more remarkable than the growth of the pine roots themselves.
Interesting questions are raised as to the identity and physiology of the
fungus involved— questions to which there are at present no answers.
The other two trees taken from saturated soils grew on a lower site,
in what probably was originally the ecotone between white cedar and
pine stands. (Clearing had disturbed the natural vegetation.) The
water table was about 8 inches below the surface in dry weather; for
several weeks after a rainy period it remained only about 4 inches
down. The site was not a true bog as known in glaciated regions. The
surface soil to a depth of 6 to 10 inches was composed largely of fibrous
plant debris, hummocks of cinnamon fern, and sphagnum moss. It
was densely interwoven with living roots and rhizomes. This fibrous
mat rested on sand, with a sharp line of demarcation.

The specimen trees were 4 inches d. b. h. and about 20 feet tall,
somewhat smaller than the one discussed above. Both were vigorous
and healthy in appearance, the more recent internodes being 12 to
18 inches long. One of these trees was essentially taprooted, though
the shaft forked into two almost equal and parallel parts and divided
extensivelv into a fan-shaped mass of many-branched and contorted
roots (fig/l3, (7). It ended at a depth of 4 feet, with a few mdividual
roots going about 1 foot deeper. The finding of two such fans indi-
cates that their formation is a normal response to the saturated condi-
tion. Why this is the case must, at present, be left to conjecture.
The branches did not originate wholly on opposite sides of the tap-
root, which would seem to eliminate diarchy of the protostele as an
explanation. In fact, many of the roots were triarch. This tree,
also, showed numerous growing tips and mycorrhizal clusters.

The second smaller tree had no taproot, but was anchored by four
strong sinkers located a few inches out from the stump. They ended
at depths of 2.5 to 3 feet. One of these obliqued downward at an
angle of about 45°, which is notable as the only example found on
any tree of a major root following such a course. (Major roots are
almost always either definitely vertical or definitely horizontal.)
This tree had developed no fanUke structure.

These three trees demonstrate that pitch pine can thrive on satu-
rated soils ; that it will root soUdly and deeply on such soils, with
no more than the usual danger of windthrow; and that mycorrhizae
can develop under conditions of saturation. . .„ , .

Before applying these conclusions generally, it will be necessary to
examine trees in other regions and on other sod types. It is possible
that conditions in the pine barrens permit a greater degree of aeration
of ground water than usually occurs elsewhere. As is well known,
many herbaceous plants that are intolerant of saturation will grow
in water culture if properly aerated. Hole (M) has shown that some
trees, at least, respond in the same manner. Hesselman (11) attrib-
utes the growth of spruces in the saturated soils near springs to the

1006

Journal of Agricultural Research

Dec. 1, ia35

Root Development of Pitch Pine

1007

Vol. 51, no. 11

unusual aeration of the water. Further investigation is necessary to
determine wlietlier it is the species or the site that is most unique in

southern New Jersey. . i !•«.

Plant roots growing under water often show anatomical differences,
particularly in the development of air spaces. Adamson (1) reports
the presence of lacunae in both the primary and secondary cortex of
the submerged roots of Terminalia arjuna. However, no air spaces,
nor marked looseness of the cortical cells, can be discerned in the
submerged roots of pitch pine. Hesselman (11) mentions that neither
pine nor spruce roots develop special air passages when submerged.
Bondois, in a paper dealing with anatomical features of the roots of
mesic trees grown in water (2), noted that spruce (the only conifer
studied) did not develop cortical air spaces, though this did occur in
several hardwood species.

ROOT DEVELOPMENT ON HEAVIER SOILS

Conditions did not permit extensive comparative studies of Pinus
rigida on widely different soil types, though such studies are much to
l)e desired. Some observations and partial excavations of root sys-
tems were made near Mont Alto, Pa., where pitch pine grows on the
higher ridges in the mountains. The soils are raw, podsolized, stony,
and high in clay. They are rendered so sticky and compact by the
clay that spading is almost impossible without preliminary loosening
with a pick. These soils offer far greater physical resistance to root
penetration than do the sands of the Coastal Plain and in all prob-
abiUty retard root elongation irrespective of other factors.

Two small trees, 11 and 14 feet tall, were examined. The following
differences, in comparison with New Jersey specimens, were noted:

(1) Vertical roots were generally weaker in both thickness and
length. Taproots of these trees reached depths of 3 to 3.5 feet.

(2) Primary laterals were shorter, the longest ones scarcely exceed-
ing the stem in length.

(3) Laterals frequently ran closer to the soil surface than in the
New Jersey sands. This probably is a result of the greater moisture-
retaining capacity of the clayey soil.

(4) Laterals followed more tortuous courses through the soil owing
to the resistance of the soil itself and to obstructing rocks.

(5) Secondary branches were generally more numerous per linear
unit of primary root.

(6) Primary laterals had undergone much more basal thickening
than is usual on New Jersey trees of the same size. This is undoubt-
edly correlated wath the weakness of the taproots and the consequently
greater burden of support placed upon the laterals. Stronger pre-
vailing winds, also, may be a factor.

These generalizations, though derived from a limited number of
observations, are in essential agreement with statements in the litera-
ture relating to tree roots. Hence they carry more weight than could
be granted to them if they were not thus supported. Of particular in-
terest are the observations of Rigg and Harrar {22) on the roots of con-
ifers growing in sphagnum peat. They found markedly greater root
elongation in this loose substratum than in more compact and resistant
soils. They agree with Biisgen and Miinch U) that mechanical ob-
structions rather than conditions of nutrition retard root elongation.

ROOT FUSIONS

Frequent references are made in the literature to root fusions, both
within the same system and between different systems. The occa-
sional persistence of life in coniferous stumi)s is usually explained as
resulting from intersystemic fusions with the roots of a living tree
ffig. 14). In this investigation, such natural roots grafts were found
only rarely, and with
one exception involved
root of the same tree.
Some were found on
both pitch and short-
leaf pines. Close ag-
gregation of trees is
most conducive to root
grafts. The fact that
sucli situations were
avoided in choosing
specimen trees prob-
ably accounts for the
small number observed.

ABSORBING AREA

AND THE PROBLEM

OF ABSORPTION

This report is con-
cerned primarily with
the gross features of
root systems. The de-
scription and interpre-
tation of the ultimate
branching patterns and
of functional areas of
absorption entail more
detailed study, involv-
ing tip-by-tip analyses
and laboratory experi-
mentation.

In a general way, the
terminal parts of ver-

Fn.uKE 14— A iialural root fusion ou Pinus echiiintn.

tical roots are coarse and sparsely branched (hg. lo) ; those of horizon-
tal roots are more slender and much more profusely branched. On
vertical roots, the diarchy or triarchy of the protostele often can be
determined readilv bv the alinement of the branches hg. lb); on
horizontal roots, the alinements usually are obscured by the profuMon
and contortion of the branches. Roots feeding in the surface huimis
are most intricatelv branched; the network often is so complex that it
is next to impossible to obtain any sizable portion intact and free trom
(.rcranic debris. The intricacy is enhanced further by mnumerab e
nivcorrhizal clusters. Though mycorrhizae are found at all levels
(fig. 15), they occur in greatest abundance m the surface organic
lajers of soil.

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Journal oj Agricultural Research

Voi. 61, no. 11

Dec. 1, 1935

Boot Development oj Pitch Pine

1009

The amount of functional absorbing area is, in the final analysis,
one of the most significant features of root systems. The extent and
degree of branching of the skeletal framework ol a root system is
important only insofar as it brings the absorbing membranes of young
tips into contact with the necessary water and nutrients, bxact
knowledge of the root parts wherein absorption occurs is very meager.
It is not known with certainty how much of the terminal region of a
root actively absorbs, or whether dormant tips can absorb, or whether
mycorrhizae are important as absorbing organs. According to the
usual concept, absorption takes place primarily in the zone ot exten-
sion found between the apical meristem and the suberized mature
region of a growing root. Scott {23) has reiterated this general idea,

Figure 15 -Tip regions of two vertical roots taken 7 feet deep in drained soil. The clusters are mycor-
rhizae. Xote the paucity of permanent branches.

and pointed out that, with retarded growth due to drought or other
unfavorable conditions, suberization processes continue and encroach
on the absorbing regions of the root. The tip ultimately may be
completely enclosed by a suberized covering. •£• i u u

It has been observed by various investigators, and verified by the
writer in respect to the species studied, that the roots of pines do most
of their growing in the spring and fall. Finding a growing tip was
a rare occurrence in July, August, or early September. The dormant
tips found generally during that period were brown and shrunken as
compared with growing ones. Free-hand sections showed the cortical
cells to be in a flaccid condition, and the cell walls were brown inward
to the endodermis. They appeared to represent thoroughly that
state described by Scott wherein the root tip is completely enclosed
by a suberized covering. If it is admitted that active absorption takes
place only in growing tips, and if Scott's statement is accepted that

''the distribution of suberization may be used as an indication of the
delimitation or curtailment of the absorbing region of the root ,
then the incongruous situation is presented of absorbing area being
reduced to a minimum when transpiration stresses are greatest.
Scott supplies no answer to this question. Perhaps that actually is
the case and the tree obtains its water during the late summer months
solelv through the few vertical roots whose tips are not completely
dormant. Though many of the tips found deep in the subsoil also are
dormant, and the number of growing ones seenis who ly madequate to
supply the needs of the tree, the fact remains that the tree passes
through the dry summer period annually without visible deletenous
effects. The prevaihng paucity of root hairs on pitch pme roots ren-
ders the mode of water entry
even more uncertain. The
whole problem of the loca-
tion and mechanics of water
absorption in the pine offers
a fertile field for further re-
search.

SIGNIFICANCE OF THE
ROOT HABITS OF PITCH
PINE

The relation between ini-
tial root habit and the abiUty
of seedUngs to maintain
themselves on various sites
has been emphasized by
Tourney {25) and is gener-
ally recognized by foresters
and ecologists. To establish
itself on a xeric site, the
seedling must prompt.ly
make contact with the more
permanent reservoir of mois-
ture in the subsoil. Those
species which push their out-

Dosts farthest into the mid- _ , r- u.,

western prairie are notably deep-rooted. Quercus rnacrocarpa Michx
ZTjugfans nigra L. may extend their taproots .^own mor^^^^^^^^^
4 feet during the first season of growth {13). Species with inflexibly
shallow root! are restricted to sites where the topsod is always loist^
The excessively drained sands of the pme barrens constitute a
relatively xeric situation; hence the native pitch pine might be ex-
pected to put down a vigorous taproot during the first season As
Lted in the discussion of seedUng root sys ems, l:y^^r-«l{\.Pl^^^^^^^
reach a depth of 1 foot or more on exposed ^/^^uations Alt^^^^^^^
this is not a striking development, it does bring the ^o^ts into contact
with sands wherein the water content only ^^^^y rarely is de^^^^^^^^
the point of the wilting coefficient. In comparing ^^is grow t^^^^^
that of xeric broad-leaved trees of the Middle \A es , ^^^^^^^ *^^^^7^^^^
in mind that the latter undoubtedly transpire much more water than

FK.iKE Ifi.-Line drawing of two tips of vertical roots The
one on the left was triarch and that on the right, diarch.

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Journal oj Agricultural Research

Vol. 51, no. 11

do needle-leaved seedlings in the more humid atmosphere of the
eastern seaboard and therefore require a deeper and more extensive
root system.

The literature indicates that seedlings of pines, in general, develop
deeper root systems than most other native gymnosperms, but, at
best, fall far short of the growth attained by such hardwoods as
black walnut and the more xeric oaks and hickories. Pinus palus-
trls, growing in the southern sandy Coastal Plain, ordinarily does
not penetrate the soil deeper than 1 foot during the first season {18).
P. ponderosa Laws., the dry-land tree of the Rocky Mountain area,
does not develop unusually deep seedling root systems (8). Evi-
dently the successful ecesis of pines on dry areas cannot be wholly
ascribed to root adaptations. Other features, including reduced
transpiration, undoubtedly are significant. Pitch pine is equaled
or surpassed, with respect to depth of seedling root systems, by
several other pines and by various hardwoods. From an ecological
viewpoint its seedUng root habit is only one of the complex of char-
actei-s by means of which the plant is adapted to xeric situations.
Though it would be presumptuous to account for the prominence
of the species in the flora of the pine barrens on the basis of anj^ one
character, resistance to fire probably is most decisive (W).

With respect to the extent of root systems of mature trees, pitch
pine conforms generally with other species for which information
is available. The similarity between the root systems of pitch and
shortleaf pines has been noted above. The root development of
jack pine (Pinus banksiana Lamb.) evidently closely parallels that
of P. rigida and P. echinata (6). Longleaf pine and Scotch pine, at
least under certain conditions, may considerablv exceed pitch pine
in spread of roots. Lengths of 70 to 80 feet are reported (12, 17).
Laterals of the bur oak attain lengths of more than 60 feet (29).
The horizontal roots of the pecan may extend laterally 30 feet on
12-year-old trees, and generally maintain lengths of about twice
the lateral spread of the crowii {30, 31). Such figures, of course,
are not strictly comparable, since the various species grew under
widely differing conditions of soil and climate. They show, however,
that despite soil conditions generally conducive to root elongation
the extent of root systems of pitch pine is not exceptional. Further-
more, they show the fallacy of the old notion that the spread of root
systems of trees approximates the spread of the crowns. In trees
beyond the sapling stages, the spread of roots is nearly always at
least twice, and in many cases four or more times that of the tops.
Equivalence of spread above and below ground is to be regarded as
the exception rather than the rule.

Depth of roots is more generally responsive to soil conditions than
is their lateral spread. The figures reported for pitch pine in this
pai)er certainly do not approach the maximum depths attainable
by tree roots. Even on the conservative basis of a depth of 8 feet
and a lateral spread of 30 feet, the root system of a tree is seen to
occupy an enormous volume of soil. Trees in nature nearly alwavs
are spaced so closely that considerable interminghng of root systeins
results. With these are associated the roots of all the subdominant
shrubby and herbaceous plants of the forest floor. In general, the
underground parts of the plants of the forest are more intimately

I

Dec. 1, 1935

Root Development of Pitch Pine

1011

>

)

1

associated with each other than are the tops. Particularly on dry
or infertile soils, underground relations probably determine the
density of the vegetation. \Mien viewed with a full appreciation
of the extent and interrelations of root systems, it may justifiably
be contended that the complexity of the forest community attains
its greatest expression underground.

From a silvicultural standpoint, several suggestions are warranted
on the basis of this study. They may be listed as follows:

(1) Only seedlings with strong, well-branched root systems should
be planted. A root system poorly branched in early life will always
be weak, both as a supporting and as an absorbing structure, and
will result in a weak tree.

(2) Pitch pine is not recommended for planting in shallow soils
overlying soUd rock. The tendency to develop fairly deep vertical
roots is strongly inherent in the species. Complete inhibition of
the vertical development would result in weakly anchored trees, and
probably would cause physiological disturbances.

(3) In the sandy soils of New Jersey, pitch pine may be planted
on sites where the water table is as close as 8 inches to the soil sur-
face. Development of vertical roots under such conditions is not
markedly inhibited, and the trees are firmly anchored. Planting
on sites where the water table remains at the soil surface is not rec-
ommended without further study. Aeration of the surface soil
where the majority of the lateral roots are found may be essential
to the health of the tree. Also, planting on heavy soils with a high
water table is not recommended until the reaction of the si)ecies
on such areas has been investigated.

(4) Pitch pine probably can best be used in mixture with other
species. The natural oak-pine mixtin-es cliaracteristic of the pine
barrens constitute a pertinent suggestion. All observations indicate
that the greatest productivity of the soil is to be realized by folio ^ving
this hint from nature. Pitch pine roots extensively, but not inten-
sively. Some intermingling of the roots of adjoining trees may take
place without initiating marked competition between them. In the
case of such intolerant species as pitch pine, crown closure does not
necessarilv hidicatc complete closure underground. The mixed
planting, in which the intolerant pine is given a start over more
tolerant associated species, thus promises the greatest return.

COMPARATIVE OBSERVATIONS ON PINUS ECHINATA

Shortleaf pine {Pimis echinata) is an important component of the
forest on drained soils of the Lebanon area. It occurs in mixture
with P. rigida and the oaks, or, locally, in almost piu'e stands. How-
ever, it does not follow the pitch pine into low areas of poor drainage.
Presumably, it cannot tolerate a high water table. In the Lebanon
area, shortleaf pine appears to be even more intolerant of reduced
light than pitch pine. The trees are remarkably well self-pruned,
and if at all crowded tend to become spindling and weak.

Although this investigation was centered around pitch pine,
observations sufficient to establish certain similarities and differences
between the two species were made. It has been pointed out that
no constant differences are apparent in the younger stages of growth

1012

Journal of Agricultural Research

Vol. 51, no. 11

(up to 10 years of age). Differences are apparent in the roots of
trees approaching maturity, but the stage of development at which
those differences become conspicuous was not determined. The
most striking difference is in the development of the taproot. Pinus
echinata produces a much more powerful and massive central shaft
(fig. 17) that maintains its thickness to greater depths and displays
less tendency to divide into an array of descending branches. The
depths attained by the vertical roots of the two species are essentially
the same.

The root systems are similar in general form and extent. The
same general relationships between spread of roots and height of

tops, and between lengths and
diameters of laterals, seem to
prevail as in pitch pine. Also,
the growth of laterals follows
the same sequence, i. e., elonga-
tion and the maintenance of a
ropelike character during the
first decades of the tree's life,
followed by retarded elongation
and the initiation of basal thick-
ening as the stresses incident
to an enlarging crown increase.
Thickening, however, is less
pronounced in shortleaf pine,
inasmuch as the taproot has
assumed the greater burden of
support, and for the same reason
strong supporting sinkers near
the root crown are few or
wanting.

These comparative general-
izations concerning Pinus
echinata are based on examina-
tions of three specimen trees.
Two of them were somewhat
weak and spindling, 25 feet and
31 feet tall, respectively, and
4 inches d. b. h.; the third was
a mature tree, 8.5 inches in
diameter, 45 feet tall, and about
85 years old. It may be noted,
incidentally, that the longest
root excavated during the entire investigation was found on the
somewhat atypical 25-foot specimen of P. echinata. The taproot
of this tree gave off only 2 sizable laterals, the larger of which was
10 by 5.5 cm in cross section at the base, and 50 feet long. The
root was unusually well branched; obviously, this was somewhat
compensatory for the paucity of primary laterals. It is very doubt-
ful, however, that the 2 primary laterals, even though exceptionally
well developed, could equal the usual complement of 15 to 30 pri-
maries in absorbing area. Certainly such a root system is mechanic-
xilly weak as a structure for anchorage and support.

Figure 17.— Taproot of an 8o-year-oI<J sliortleaf pine,
showing its massive character.

f

t

Dec. 1. 1935

Root Development oj Pitch Pine

1013

)

The suggestions given for silvicultural management of pitch pine
hold equally well for shortleaf pine, except that this species cannot
tolerate situations where the roots reach saturated soil.

SUMMARY

The root system of Pinus rigida was studied by the direct, dry
method in the pine-barren section of New Jersey. It is to be classed
in the generalized type. It attains a moderately extensive develop-
ment both vertically and horizontally. The basic plan consists of a
taproot from which 15 to 30 horizontal branches originate and extend
radially in the surface layers of soil. From these primary branches
horizontal and vertical secondary branches develop; these in turn
give off tertiary branches, and so on. Normally, the root system
occupies a roughly circular area of topsoil; in all of this area except
the marginal parts it has extensive contacts with the subsoil through
the development of vertical, or sinker, roots.

Seedlings are prominently taprooted. They reach depths varymg
from 3 to 12 inches during the first season of growth.

The taproots of seedlings 4 to 5 years old reach depths of 15 to 24
inches, and the strongest laterals approximate the same length.
Plants of this age are semidecumbent ; a permanent crook persists
at the ground line long after they have become rigidly erect.

The taproots of plants 8 to 9 years old reach depths of 1.5 to 2.5
feet, and the strongest laterals approximate the same length. Plants
of this age usually are erect.

Saplings 12 years of age are beginning to display regular whorls
of stem branches, and are assuming the aspect of a tree. Taproots
reach depths of 3 to 4 feet, and the strongest laterals extend 6 to 8
feet. The period between the eighth and twelfth years of the young
tree's life witnesses the following developments:

(1) An acceleration in the growth rate of primary lateral branches,
as a result of which the radial lateral spread of the root system be-
comes approximately twice the length of the taproot, and 1.5 to 3
times the length of the stem. Hence, the lateral roots displace the
taproot as the most prominent feature of the root system.

(2) Retardation of the growth rate of the taproot as it penetrates
the subsoil below the 3-foot level.

(3) The attainment of linear equivalence of stem and taproot,
prior to wliich the taproot was longer, and after wliich the stem is
always longer. The difference in length between stem and taproot
constantly increases with the growth of the tree.

(4) The attainment of conspicuous size by secondary and tertiary
branches on the stronger primary laterals. .

Root growth between the ages of 12 and 30 years, or, on the basis
of stature, between the heights of 4 and 25 feet, is characterized as

follows: , . ... ,, •

(1) Continuous elongation of the primary laterals until lengths ol

25 to 35 feet are attained. .i *

(2) Maintenance of a ratio of about 1.5 to 1 between the length of
the strongest laterals and the height of the tree.

(3) Maintenance of a ratio approximating 1 mm to 1 foot between
basal diameters and lengths of lateral roots. , , . . ,

(4) Continuous elongation of branches of the higher orders, in-
creasing the density of roots within the occupied volume of soil.

1014

Journal of Agricultural Research

Vol. 51, no. 11

D9C. 1. 1935

Root Development oj Pitch Pine

1015

(5) Continuous elongation of vertical roots, Nvliich, however, after
reaching depths of 3 to 4 feet, grow much more slowly than the
surface laterals, and practically cease to grow at depths of 8 to 9 feet.

As a result of the processes listed above, a constantly increasing
proportion of the root system as a whole becomes localized in the
surface soil.

After trees have reached a height of about 25 feet, and a diameter
breast high of about 4 inches, root development enters gradually
upon a different phase. It is characterized as follows:

(1) Cessation of elongation by the stronger primary laterals and
by the taproot and stronger sinkers.

(2) Marked thickening of the basal 2 or 3 feet of the stronger
primary laterals. The increment usually is greater in the vertical
plane, forming narrowed, planklike roots; in manv cases, the incre-
inent is added almost entirely on the top side, resulting in eccentrically
thickened, buttresslike supporting roots. This basal thickening
increases the mechanical strength of the central root svstem, and in
all probabihty is initiated by the stresses incident to*^ an enlarcrincr
crown . '^

(3) Continued elongation of the smaller primary laterals until
they approximate the length of the stronger ones, after which the\'
too, undergo basal thickening.

(4) Some continued growth of branches of the higher orders
resulting in increasing density of roots in both the topsoil and the
subsoil.

(5) A tendency toward marked thickening of the innermost sinker*,
buch thickening occurs in response to the requirements of the tree
tor support, hence the development is inverselv proportional to the
strength of the taproot.

Lateral roots function mechanicallv as props or braces, and onlv
incidentally as guy wires. Thickening is stimulated by compressi(»n,
and not by tensile strain. This is conclusivelv demonstrated bv the
eccentric development of the root svstems of leaning trees.

Poor development of tops is associated with inferior root svstem*.

Koot branches, irrespective of size, can be traced inward to the
primary xylem of the parent root, thus indicating that they originated
when that region of the parent root was young. Hence, a root svstem
poorly branched in youth will always be poorlv branched, and will
predispose toward weakness of the tree. Adventitious root branrhe*
appear to be absent in pitch pine, except that in rare cases thev niav
develop in wound tissue following clean severance of a root

Keplacement tips develop following the death of the terminal
portion of a root only when death has not extended back into the
region where secondary wood completely encircles the primarv xylem

1 itch pine is capable of extensive root growth below the water
table in saturated soils. The descending branches of taproots under
water sometimes are arranged in a peculiar and characteristic fan-
shaped mass, the explanation of which is obscure. Trees crrowinL'
on saturated sods appear to be in full health and vigor

On sloping sit^es, the primary lateral branches generally parallel
the soil surface, both uphill and downhill. ^

On heavier soils, root development tends to be less extensive, both
Pldn""" vertically, than it is in the sandy soils of the Coastal

Root fusions occur occasionally, on both Pinus rigida and P.
echinatay between roots of the same or of different systems.

The fine roots of the higher orders, through which most absorption
presumably occurs, are most profusely developed in the upper soil
layers, and frequently extend into the raw surface humus. The
ultimate branches of vertical roots are generally coarser and are
produced in much less profusion. Nearly all root tips become dor-
mant, and many die, during the dry periods of midsummer. Growth
takes place mostly in the spring and fall.

Mycorrhizae are a conspicuous feature of pitch pine roots. Their
period of growth coincides with that of nonmycorrhized tips. They
are found at all depths on both drained and saturated soils, but
attain their greatest profusion in the surface organic layers.

With respect to adaptations to xeric sites, the root systems of
pitch pine show no apparent superiority over those of many other
species of trees.

Certain suggestions are offered with respect to silvicultural practices.

The root development of Pinus echinata in New Jersey is funda-
inentally similar to that of P. rigida. The most marked difference
is the tendency of P. echinata to develop a much stronger and more
massive taproot, with wliich is correlated a weaker development of
secondary supporting roots. This species does not invade areas
where the water table is high.

LITERATURE CITED
(I) Adamson, R. S.

1910. NOTE ON THE ROOTS OF TERMINALIA ARJUNA, BEDD, XeW Plivtol.

9: 150-156, illus.
(2j BoxDois, G.

1913. CONTRIBUTION A L'ETUDE DE L'INFLUENCE DU MILIEU AQUATIQUE

SUR LES RACINES DES ARBRES. AllM. ScI. Nat., Bot. (9) 18: 1-24,

illus.

(3) Burns, G. P.

1914. studies in TOLERANCE OF NEW ENGLAND FOREST TREES. I.

DEVELOPMENT OF WHITE PINE SEEDLINGS IN NURSERY BEDS.

Vt. Agr. Expt. Sta. Bull. 178, pp. 127-144, illus.

(4) BusGEN, M., and Munch, E.

1929. THE structure and LIFE OF FOREST TREES. Traiisl. from German
by T. Thomson. Ed. 3, rev. and enl., 43G pp., illus. New York.
(5j Cannon, W. A.

1911. THE ROOT HABITS OF DESERT PLANTS. 96 pp., illus. Washington,

D. C. (Carnegie Inst. Wash. Pub. 131.)

(6) Cheyney, E. G.

1932. THE ROOTS OF A JACK PINE TREE. JouT. Forestry 30: 929-932,
illus.

(7) Francis, W. D.

1931. the BUTTRESSES OF RAIN-FOREST TREES. Kcw Bull. Misc. Inform.
1931 (App. 1): 24-26, illus.

(8) Haasis, F. W.

1921. relations BETWEEN SOIL TYPE AND ROOT FORM OF WESTERN

YELLOW PINE SEEDLINGS. Ecology 2: 292-303, illus.

(9) IIaberlandt, G.

1914. PHYSIOLOGICAL PLANT ANATOMY. Transl. from 4tli German ed. by
M. Drummond. . . . 777 pp., illus. London.

(10) Harshberger, J. W.

1916. THE VEGETATION OF THE NEW JERSEY PINE-BARRENS; AN ECOLOGIC

INVESTIGATION. 329 pp., iUus. Philadelphia.

(11) Hesselman, H.

1910. om vattnets syrehalt och dess inverkan pa skogsmarkens
FoRSUMPNiNO OCH SKOGENS VAXTLiGHET. Meddel. Statens
Skogsforsoksanst. [Sweden] 7: [91J-125, illus. [In Swedish.
Resume in German, pp. [.\iii]-xvi.]

1016

Journal oj Agricultural Research voi. 51. no. 1 1 , Dec. 1 , 1935

(12) Hetward, F.

1933. THE ROOT SYSTEM OP LONGLEAP PINE ON THE DEEP S\NDS OF

/ION XT A WESTERN FLORIDA. Ecology 14: 136-148, Hlus.

(13) HoLCH, A. E.

1931. DEVELOPMENT OP ROOTS AND SHOOTS OF CERTAIN DECIDUOUS TREE
SEEDLINGS IN DIFFERENT FOREST SITES. EcologV 12: 259-298

illus.

(14) Hole, R. S.

1918. RECENT INVESTIGATIONS ON SOIL-AERATION. PART II WITH

SPECIAL REFERENCE TO FORESTRY. Agr. Jour. India 13: 430-
440, illus.

(15) Illick, J. S., and Aughanbaugh, J. E.

1930. pitch pine in PENNSYLVANIA. Pa. Dept. Forests and Waters
/,^x T^ Research Bull. 2, 108 pp., illus.

(16) KuRz, H., and Demaree, D.

1934. CYPRESS buttresses and knees IN RELATION TO WATER AND \IR

, ^, ^ Ecology 15: 36-41, illus.

(17) Laitakari, E.

1927. mannyn juuristo morfologinen tutkimus. the root system

OF PINE (PINUS SILVESTRIS); A MORPHOLOGICAL IN'VESTIGATIOX

Acta Forest. Fennica 33: 1-380. illus. [In Finnish. Summary
„„, ., in English, pp. 307 -380.

(18) Lenhart, D. Y.

1934. INITIAL ROOT development OF LONGLEAP PINE. Jour. Forestrv
32: 459-461.

(19) LuNcz, G.

1931. RECENT RESEARCH WORK ON THE ROOT SYSTEMS OF FOREST TREES

/oAx T TT Internatl. Rev. Agr. 22: 239-243.

(20) LuTz, H. J.

1934. ECOLOGICAL RELATIONS IN THE PITCH PINE PLAINS OF SOUTHERN

(21) McArdle, ^^^y^^^^^- Yale Univ. School Forestry Bull. 38, 80 pp., illus.

1932. THE RELATION OP MYCORRHIZAE TO CONIFER SEEDLINGS. Jour \cr

/«ov T. ^ Research 44: 287-316, illus. " ^

(22) RiGG, G. B., and Harrar, E. S.

1931. THE ROOT SYSTEMS OF TREES GROWING IN SPHAGNUM. Amer. JoUr.

/«„x « , ^^^- ^°- o91-397, illus.

(23) Scott, L. I.

1928. THE ROOT AS AN ABSORBING ORGAN. II. THE DELIMIT -VTION OF

(24) TouMEY, j''^/««°««iN« ZONE. New Phytol. 27: 141-174, illus.

1928. FOUNDATIONS OF SILVICULTURE UPON AN ECOLOGICAL BASIS V 1

illus. New York and London. '

(25)

1929. INITIAL ROOT HABIT OF AMERICAN TREES AND ITS BEARING OV

728 Tlus''"''''* I"t^^"atl. Cong. Plant. Sci. Proc. l": 713-

(26) Weaver, J. e'. *

1919. the ECOLOGICAL REI.ATI0N8 OF ROOTS. 128 pp., ilhis. Washington,
^' ^' (Carnegie Inst. Wash. Pub. 286.)

(27)
(28)

(29)

— and''BRUNER!'''w "e. ""^ '''^'^'' ''''''^^' ^^^ P^' '""'' ^^^^^' Y^^^"

1927. ROOT^DEVELOPMENT OF VEGETABLE CROPS. 351 pp., ilhlS. NeW

and Kramer, J.

1932. ROOT SYSTEM OF QUERCU8 MACROCARPA IN RELATION TO THE
(30) W00DR00F,TT" '' ^"""" ^°*- ^^^- ''■ ''-''' ^"-•

1933. RELATION OF THE ROOT SYSTEM OF PECAN TREES TO NURSERY ^ND

(31) and^^^rR^o^ptT^J: ^^- ^'^P*- ^''' ^""- '''' '' PP-f illus."'

1934. ^=^;^f_«^OT GROWTH AND DEVELOPMENT. Jour. Agr. Research 49:

U. S. GOVEUHMENT PRINTING OFFICE: 1928

y-^

STUDIES IN THE TAXONOMY AND DIS-

TRIBUTION OF THE EASTERN NORTH

AMERICAN SPECIES OF LOBELIA

A THESIS

IN BOTANY

PRESENTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE

UNIVERSITY OF PENNSYLVANIA IN PARTIAL FULFILLMENT

OF THE REQUIREMENTS FOR THE DEGREE

OF DOCTOR OF PHILOSOPHY

ROGERS McVAUGH

• »»

•4

Rhodora, Vol. 38, pp. 241-263, 27&-298, 305-329, 346-362.

Uhodora

Plate 435

^J

.#^

V. - i ' ' '^- ^'^'''''^•■'•'•"; "'•<•• 5, I^. Caxbyi; fk;. 6, L. spmaia, var < ukjina ,s■

1.. (.LAMHLosA, PKj. U, L. flokiuaxa; fk;. 12, L. inflata; fk; i;i L Doktmawa'
FK. 14, L. amoexa; fig. 15, L. siphilitica; fig. 16, L. Cakdinalis. ^""''•^'^'^^'^a,

?

(

7

STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES OF LOBELIA

Rogers McVaugh

(Plates 435 and 436)

The present paper is intended as a study of the distribution of the
species of Lobelia native to the eastern part of North America, with a
view to establishing better understanding of the relationships within
the group, the possible origin of the various species, and the relation
of this group of species to the worid-wide genus Lobelia. Eariy in the
study it became clear that the identities of various species were much
in doubt, which necessitated considerable taxonomic work, in an
effort to clarify the situation so that significant distributional studies
could be made.

Most of the work has been carried on at the Botanical Laboratory
of the University of Pennsylvania during the years 1933-1935. Two
summers have been spent in this time in eastern New York, largely
in botanical studies, so that the writer has been able to gain acquaint-
ance with all the northeastern species in the field. The remaining
species have been studied only from herbarium material.

During the course of the project about 7000 sheets of dried material
have been examined. This has been made possible through the
generosity of the following gentlemen, to whom the writer wishes to
express his sincere thanks: Dr. R. M. Anderson, National Museum
of Canada, Ottawa, Can.; Dr. W. C. Coker, University of North
Carolina, Chapel Hill, N. C; Dr. E. L. Core, West Virginia University,
Morgan town, W. Va.; Mr. C. C. Deam, Bluff ton, Ind.; Dr. J. H.
JEhlers, University of Michigan, Ann Arbor, Mich. ; Dr. N. C. Fassett,

Kliodoni

Plate 43r)

Si.:ki>s <,F LoHKiiA, X 12. Fk;. 1, L. Clikfou iiana; vu.. '_'. I. F.xwsx- ii, 'K I
»i<.. /, i>. MKAiA. \:ii. (AMI-AM lata: ik;. s, F. ithkiu la; kk;. 1», F Km mm- ,.„ in

H<.. 14, J.. AMOK.NA, H(.. 15, L. SIPHIUTKA: FKi. 1 (), L. CaHDINALLS.

STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES OF LOBELIA

Rogers McVaugh

(Plates 435 and 436)

The present paper is intended as a study of the distribution of the
species of Lobelia native to the eastern part of North America, with a
view to estabUshing better understanding of the relationships within
the group, the possible origin of the various species, and the relation
of this group of species to the world-wide genus Lobelia. Early in the
study it became clear that the identities of various species were much
in doubt, which necessitated considerable taxonomic work, in an
effort to clarify the situation so that significant distributional studies
could be made.

Most of the work has been carried on at the Botanical Laboratory
of the University of Pennsylvania during the years 1933-1935. Two
summers have been spent in this time in eastern New York, largely
in botanical studies, so that the writer has been able to gain acquaint-
ance with all the northeastern species in the field. The remaining
species have been studied only from herbarium material.

During the course of the project about 7000 sheets of dried material
have been examined. This has been made possible through the
generosity of the following gentlemen, to whom the writer wishes to
express his sincere thanks: Dr. R. M. Anderson, National Museum
of Canada, Ottawa, Can.; Dr. W. C. Coker, University of North
Carolina, Chapel Hill, N. C; Dr. E. L. Core, West Virginia University,
Morgantown, W. Va.; Mr. C. C. Deam, Bluffton, Ind.; Dr. J. H.
Ehlers, University of Michigan, Ann Arbor, Mich. ; Dr. N. C. Fassett,

#

i

INTENTIONAL SECOND EXPOSURE

242

Rhodora

[July

University of Wisconsin, Madison, Wis.; Dr. H. A. Gleason, New
York Botanical Garden, N. Y.; Dr. E. H. Graham, Carnegie Museum,
Pittsburgh, Pa.; Dr. J. M. Greenman, Missouri Botanical Garden,
St. Louis, Mo.; Dr. E. M. Gress, State Botanist, Harrisburg, Pa. J
Dr. H. D. House, State Botanist, Albany, N. Y.; Mr. Bayard Long'
Academy of Natural Sciences, Philadelphia, Pa.; Dr. J. C. McKee'
State College, Mississippi; Dr. W. R. Maxon, U. S. National Museum'
Washington, D. C; Dr. Aven Nelson, University of Wyoming
Laramie, Wyo.; Dr. H. J. Costing, Duke University, Durham, N. C.j
Dr. F. W. Pennell, Academy of Natural Sciences, Philadelphia, Pa.j
Mr. J. H. Pyron, University of Georgia, Athens, Ga.; Dr. C. O,
Rosendahl, University of Minnesota, Minneapolis, Minn • Dr R R
Tatnall, 1100 W. 10th. St., Wilmington, Del.; Dr. T. M.'c. Taylor*
University of Toronto, Toronto, Can.; Mr. C. A. Weatherby, Gray
Herbarium, Harvard University, Cambridge, Mass.; Dr. K. M.
Wiegand, Cornell University, Ithaca, N. Y.

Thanks are also due to Mr. S. Savage of the Linnean Society of
London, through whose kindness several photographs of Linnean
types were secured, to MM. F. Gagnepain and M. Humbert of the
Museum National d'Histoire Naturelle of Paris, who gave information
concerning specimens in the herbarium of Lamarck; to Dr. K. D Doak
of the University of Pennsylvania, who gave his time to take the
photographs of seeds; to Dr. J. H. Barnhart of the New York Botan-
ical Garden and Miss Ruth Sanderson of the Gray Herbarium, who
supplied bibliographic information; to Professor M. L. Fernald of the
Gray Herbarium, who placed at the writer's disposal the facilities
of that institution.

Finally, to Dr. Edgar T. Wherry, who gave many helpful sugges-
tions and contributed a number of his personal collections for study
and to Dr. John M. Fogg, Jr., whose cooperation made possible the
taxonomic part of the work, the writer is deeply grateful.

Historical Introduction

The genus Lobelia was unfamiliar to the eariy European botanists
as only two species are represented in Europe, and these are widely
dissimilar m appearance, and not co-extensive in their ranges It is
not until the second half of the 17th century that related species of the
genus are consistently grouped together.

The first mention in literature of a species of Lobelia appears to be

i

1936] McVaugh,— The Taxonomy and Distribution of Lobelia 243

that made by Charies I'Ecluse (Clusius) in 1611 (10). This is a
description of L. Dortmanna L., and is copied verbatim by Ray (61).

A North American Lobelia is brought to attention by John Parkin-
son in 1629 (51) ; he had plants of L. Cardindis L. from France, which
had come originally from the St. Lawrence valley. No distinction is
made by the eariy writers between this species and the Mexican
L. splendens Willd. and L.fulgens Willd. (cf. Hernandez (27)).

By the end of the 17th century the campanulaceous affinities of
Lobelia had come to be well recognized: in 1623 Bauhin (4) had
included L. mens L. among the Mustards, but in 1686 Ray (60) places
all the Lobelias known to him (except L. Dortmanna L.) under
Rapunculus, which included most of the Campanulaceae. Moreover,
Plukenet (53) distinguishes Lobelia (Rapunculus) from the rest of the
Campanulaceae.

The greatest advance in classification, before Linnaeus, is made by
Tournefort (73), who defines sharply the genus Rapuntium.

The name Lobelia is first used by Plumier (54) for a related genus,
Scaevola L. Plumier dedicates the genus to Matthias de Lobel
(1538-1616), the Flemish doctor and Botanist to James I of England.

After Plumier's use of the name Lobelia, it is taken up by Linnaeus
for the genus known by the name at the present time (35-41).

General Discussion

As understood today, the genus Lobelia comprises between 200 and
250 species, widely distributed. The great majority (neariy 9/10) of
the named species are native to Australia and South and Tropical
Africa, with a large number in South and Central America and
Mexico, as well as the Pacific Islands. Several species are found in
China and eastern Asia. The genus is represented in western Europe
by two species, and is absent from the rest of the northern Eurasian
continent except in the extreme east. The North American Lobelias,
as treated in the present paper, consist of 27 named species and
varieties.

In summarizing, it may be seen that Lobelia is largely a genus of the
Southern Hemisphere. The same may be said in general of the whole
group Lobelioideae. Bentham (5) (1875) makes the following spec-
ulations :

"That the primitive race (a hypothetical ancestor of the whole
family Campanulaceae) flourished very early in some region in con-

IRREGULAR PAGINATION

244

Rhodora

[July

nexion with Africa. That the Lobeliae were first developed at a time
when the geological or other conditions afforded some general means
of communication between South Africa and Australia, between
Australia, New Zealand, and Antarctic America, between South
Africa and extratropical South America." These speculations were
made largely because of species or genera common to New and Old
Worlds. In addition, Bentham says,

"From thence (the place of origin in the Southern Hemisphere)
Lobeliae appear to have spread in several distinct directions into and
beyond the tropics, without any transverse northern connexion
between the several lines."

It should be pointed out here that too much stress must not be laid
upon such evidence as the above, in determining the origin of the
North American species, for the following reasons: In the first place
the genus as a whole is evidently highly advanced from an evoluiionary
standpoint, as shown by the structure of the flower and fruit Some
authors have assumed that the ancestors of the highly advanced
Compositae must be sought among the Lobelicndeae (Delpino (13)-
bmall {68)). Secondly, the worid-wide distribution of the various
species combined with the great diversity of vegetative structure,
types of inflorescence, types of seed coats, and flower color, points to
the conclusion that the group has enjoyed a long evolutionary, or
geological, history.

Rock (1919), in his monograph (63) of the Hawaiian Lobelicndeae,
says, m speaking of the Hawaiian genera,

"That their age is enormous and that they form with the Com-
positae the oldest element in our flora may be judged from their
numerous species and their distribution over the whole group

The present writer hopes to show later that several of the North
Amencan species are in an old or decadent condition, which is favor-
able to the assumption of a great age for the genus as a whole.

If It be granted for the moment that the genus actually is a rela-
tively old one, it is logical to assume that groups of species in various
parts of the worid may have arisen from several sources, which have
now disappeared. In other words, two species geographically con-
tiguous at present may have come from two widely separated ances-
tors^ both of which have died out in the intervening geologic time.

Ihis situation seems to be the one now existing in North America;
the species here designated as ''North American" {Eulobelia and Hemi-

1936] McVaugh,— The Taxonomy and Distribution of Lobelia 245

pogon, in part, of Bentham and Hooker (6) (1876)) form a distinct
group, apart from the Mexican and South American species, some of
which occur naturally or as weeds in Florida, Texas, New Mexico
and Arizona. Aside from purely structural characters, most of the
species native to the United States and Canada may be shown to have
characteristic geographical ranges and probable points of origin which
definitely relate them as a group, and separate them from the tropical
species now native in Mexico and southward.

The North American Lobelias were included by Bentham and
Hooker in their "Genera Plantarum" (6) in two sections of the genus,
Eulobelia and Hemipogon; the first of these all North American
(except the east-Asiatic L. sessilifolia Lamb.), distinguished by the
large short-pedicelled flowers in lax terminal racemes; the section
Hemipogon including species of Europe, Africa, America and Australia,
characterized by slender, simple or branching stems and few flowers.
These divisions of the genus now seem somewhat artificial. Appar-
ently no adequate classification can be devised, if based upon habit
and appearance alone.

Characters used by eariier taxonomists as natural ones, and as
criteria of aflfinity, such as pubescence of the anthers, or the presence
or absence of tufts of bristles, seem to be of no great absolute value.
The same may be said of the shape of the capsule, which may vary
considerably in the same species. The writer has been able to find
one character only, by which to separate the North American species
from those of other geographical areas: the mature seeds of this group
are peculiariy foveolate-reticulate, some more than others, according
to species, and indicating several distinct lines within the limits of the
group. The seeds of L. sessilifolia, which was formerly included in the
section Eulobelia, are smooth with prominent wings, while the seeds of
apparently related Mexican species such as L. gruina Cav., L. fenes-
tralis Cav., and the tropical L. Cliff ortiana and its relatives are per-
fectly ovoid, smooth and shining. No Mexican, Central American or
West Indian species seen by the writer has a type of seed even ap-
proaching the roughened ones of species of the United States. It is
of course obvious that a single character, however fundamental, is
never wholly trustworthy in determining relationships. It seems,
nevertheless, that an entirely consistent feature such as the above,
in conjunction with the evidence from geographical distribution,
points to a common origin for our species. Any connection with

246

Hhodora

[July

an ancestor in the Southern Hemisphere, such as that suggested by
Bentham (5), must have been very remote in time and before the
development of our present forms.

Discussion of Geographical Distribution of Species

The region under consideration is mostly eastern North America,
west to the Mississippi Valley; two species of Lobelia cross the con-
tinent, north of the moraine, and will be considered separately in
detail; phases of L. siphilitica and L. spicata push westward to
Colorado and Saskatchewan, respectively; L. Cardinalis and its very
close relatives occur west to California and well south into Mexico.
With these exceptions, all the forms concerned are confined to the
eastern half of the continent.

In the first place, it is necessary to consider briefly the geological
history of the area in question. During Cretaceous time the general
land level in eastern North America was lower than at present, so that
the present Coastal Plain was submerged, and the shore line followed
the present (16) "Fall Line," which runs through New York, Phila-
delphia, Washington, Richmond, cuts off the eastern third of North
Carolina, passes through Columbia, S. C, Augusta and Columbus,
Ga.; swings west and north in Alabama, leaving about two-thirds of
the state in the Coastal Plain; follows the course of the Tennessee
River north to its mouth; passes across southern Illinois and south-
eastern Missouri; southwesteriy through Arkansas, leaving about
half the state below it; cuts off the southeastern corner of Oklahoma
and passes southward near Fort Worth, Austin and San Antonio.

Upon the elevation of the Appalachian system and emergence of the
Coastal Plain, a considerable area was thus thrown open for coloniza-
tion by plants. We need consider no eariier major geologic changes,
since existing species were not then represented on the earth; the only
other factor that must be taken into account is that of glaciation.

The latest (and in eastern North America the best marked and
usually most extensive) glacial period was in Pleistocene (Wisconsin)
time, ending roughly 35,000 years ago. The terminal moraine {2)
reaches from Nantucket across Long Island to Pennsylvania, southern
Ohio, Indiana, and Illinois, then north to Minnesota and west roughly
along the 48th parallel. It was formeriy held that all land north of
this line was covered by a solid sheet of ice, and that all plants now
living in this area had migrated from south of the moraine since

1936] McVaugh,— The Taxonomy and Distribution of Lobelia 247

Wisconsin time. It has been shown recently, however, by Fern aid
and others (11, 17), that certain areas, such as parts of Newfoundland,
were wholly untouched by Wisconsin ice, and students of phyto-
geography, as well as glacial geologists, are coming more and more to
believe that such places as the Bruce Peninsula represent examples of
(perhaps much more numerous) tongues of land which were partially
unglaciated. If such be the case, many Canadian plants may have
persisted in these areas during the ice invasion.

Of the 27 species and varieties here considered, nine are, so far as
known, confined to the Coastal Plain, and one reaches above the Fall
Line only into the prairies of Arkansas and Oklahoma. Two species
(L. Dortmanna L. and L. Kalmii L.) are northern, and reach non-
glaciated country only rarely. The remaining fifteen plants comprise
ten rather distinct entities all of which are found in the Appalachian
region of the eastern United States (although not necessarily confined
to it), and five varieties, which, if not found in the Appalachian
region, are clearly derived from the species found there.

The case of the two northern species may be discussed first. Z.
Dortmanna and L. Kalmii are clearly very distantly related to other
American species, and to each other. The former differs from all other
species by the combination of the scapose habit and hollow linear
leaves; corolla smooth and slit only part way to the base; anthers all
tufted; pedicels ebracteolate and seeds dark, with prominent square
base. L. Kalmii also differs from all other species, having rather large,
smooth flowers, in loose racemes, pedicels bracteolate in the middle,
and very finely reticulate, acute-fusiform seeds. Both species are
found, in suitable habitats, north of the moraine, from Newfoundland
to British Columbia. The former is found also in western Europe
(Great Britain, western France, Belgium, Denmark, northern Ger-
many, western Russia, and Scandinavia, as far north as 68°, according
to the Illus. Fl. Mit.-Eur. (26)) ; the European and American forms are
apparently identical.

All theories as to the origin of these species must remain largely
theories only. The modern range of L. Dortmanna suggests a circum-
polar range in pre-glacial times. The extremely rare occurrence, both
of L. Kalmii and L. Dortmanna, south of the glacial moraine seems
to point to a survival within the glaciated area, rather than south of it;
however, the scarcity of suitable habitats such as calcareous bogs and
sandy ponds in unglaciated country may account for the distribution.

\l 1/

/

248

Rhodora

[July

Whatever the explanation of the above, it is sure that any past con-
nection with the remaining species is very remote.

With the exception of the two species just considered, the group as a
whole is rather uniform in character and rather closely related,
although divisible into the following sub-groups:

a) Species with a smooth lip, small flowers, delicate stems; in
Piedmont, Mountains and northeastern Coastal Plain represented by
L. Nuttalli; this giving way in Florida to L. Feayana, which is not
surely separable from it by any one character.

b) Species with larger flowers, usually in spikes, a hairy lower lip to
the corolla, and an entire corolla-tube (except for the dorsal slit).
This is the L. spicata complex, which has apparently given rise to
L. inflatay L. Canbyi, and possibly to L. Boykinii.

c) Two species very close to the above, but with characteristic thin
and smooth leaves, and secund racemes: L. Gattingeri of the uplands
of Tennessee, giving way in the Coastal Plain to L. appendiculata.

d) Four species evidently related to the last two, but with a
tendency to reduction of the stem-leaves, and to a fenestrate corolla
(one in which the two upper petals have separated from the corolla-
tube near the base) : L. flaccidifolia, L. Halei, L. floridana, L. paludosa.

e) Large, coarse, smoothish species with showy flowers, the corolla
with a smooth lip, fenestrate: the seeds roughly ridged and long rather
than ovoid; L. Cardinalis, L. siphilitica, L. amoena, L. elongata,
possibly L. glandulifera.

f) Coarse species with large flowers, fenestrate corolla, smooth lip.
Seeds rather small, smoothish, ovoid, resembling those of L. spicata
and its allies : L. puberula and its forms. From a form like L. puhervla
may have come the two species L. brevifolia and L. glandulosa.

The sub-groups may now be discussed in detail :

a) Inspection of the maps (Figs. 27 and 28) will show the apparent
relations : L. Nuttalli or its immediate ancestor seemingly migrated in
post-Cretaceous times, southeastward onto the emerging Coastal
Plain, where it gradually extended its range both northeastward and
southward. It did not enter the Florida peninsula, but gave rise there
to the rather similar L. Feayana.

b) In the Appalachian and Ozark regions the dominant repre-
sentative of this sub-group is L. spicata var. leptostachys, which is not
found elsewhere, except on the immediately adjacent portions of the
Gulf Coastal Plain (Fig. 14); it is not found on the Atlantic Coastal

i

i

1936] McVaugh, — The Taxonomy and Distribution of Lobelia 249

Plain, being there in part replaced by the var. scaposa (Fig. 18) ; in
the northeastern states, west to the Great Lakes, it gives way to the
var. originalis and the var. campanulata (Figs. 15 and 17). Westward
and northwestward, from Illinois and Missouri, the var. hirtella
appears (Fig. 16). Where the ranges of these varieties overlap, a host
of intermediates is found. These cannot be referred with certainty
to any of the named forms, and constitute the best reason for reducing
L. leptostachys from the rank of species, and for postulating that it or a
plant similar to it may have been the ancestor of all the varieties of
this sub-group (Fig. 19).

It is unfortunate for the sake of clarity that the rules of priority
make the northeastern L. spicata Lam. the type of the species, for it
seems to have been derived from the var. leptostachys by the dis-
appearance of the auricles (which sometimes reappear in individuals)
and by adaptation to a somewhat more mesophytic habitat. Accord-
ingly, it seems best to refer the northeastern phase to var. originalis.
The var. hirtella, on the other hand, may have arisen from the var.
leptostachys in a more xerophytic habitat.

Of the three remaining members of this sub-group, L. Canbyi is not
separable from the spicata complex by any character of the corolla;
its seeds are almost exactly similar, also. It has seemingly spread
from the Appalachian region to the Atlantic Coastal Plain (Fig. 21).
L. Boykinii is a rather distinct species of the Atlantic Coastal Plain,
whose affinities are obscure; it resembles no other species very closely,
and seems to approach L. Canbyi only through the habit and the
slightly hairy lip (Fig. 22).

The final species, L. inflata, is very distinct, but seems to show its
connection to the spicata complex by the spicate character of the
young inflorescence, the hairy lip, the seeds, which are very similar
to those of L. spicata and varieties. It is possible that some connection
may be shown through the reduced number of flowers and the sub-
inflated capsule of L. spicata var. campanulata, but this is only a
speculation. L. inflata evidently has spread from the Appalachian
region; it has, however, been unable to enter the Coastal Plain very
extensively (Fig. 20).

c) Logic similar to the above would demand that L. Gattingeri be an
ancestral type, and have given rise to L. appendiculata (Fig. 23).
However, the range of the former is so restricted that such a con-
clusion is largely guesswork; with the reservation that, as stated

\

250

Rhodora

[JXJLI

\

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elsewhere, the two are difficultly separable in the final analysis other
than through geographic range. It would seem to be taxing the power
of coincidence to postulate two forms having had exactly parallel
development, but no relationship. This is especially true in such a
region as the one under discussion, where the evidence seems to be for
development away from the Appalachian region, in radial directions.
d) On the southern Coastal Plain occurs this sub-group of four
species. They show their relation to L. Gattingeri and L. append-
iculata by a tendency to develop auricles, and by the characteristic
bell-shaped capsule. They show their connection to sub-groups (b)
and (c) by the spicate habit and the hairy lip. Two of the species,
L. flaccidifolia and L. Hcdei, of the southeastern and southwestern
Coastal Plains, respectively, are set apart by the large, usually green
bracteoles near the middle of the pedicels (It is possible that L.
flaccidifolia is only a habitat form: cf. discussion under this species).
Both show an increase in flower size from the spicata complex, and
there is a tendency for the corolla to become fenestrate. The leaves
in L. Hold (Fig. 24) may be rather bunched near the base of the
stem, as is sometimes seen in L. spicata var. hirtella.

In L.floridana this reduction of leaves has gone further, so that they
are nearly all basal; the prominent auricles of L. Halei have vanished,
and the bracteoles are inconspicuous. This is a species mostly (so
far as known) of the Gulf States, from west Florida westward along
the coast. It is also known from Wilmington, N. C. (Fig. 25).

Apparently the most advanced of this sub-group is L. paludosa,
which has developed in peninsular Florida, and west about to the
Apalachicola River. The leaves are entirely basal, there are no
bracteoles on the pedicel, and the corolla has become plainly fenestrate
(Fig. 26).

e) This sub-group is a rather composite one, based in part upon the
patently artificial character of size; the species seem, however, to agree
well in seed characters (Plate 435).

In the first place, L. Cardinalis is a wide-spread species, very
different in form of flower, as well as in color, from other Worth
American ones; its color suggests Mexican affinities (although parallel
development in the case of color is not by any means rare), as does
the fact that it is represented throughout the Southwest by the plants
passing as L. splendens WiUd. (Fig. 4) and L. fulgens Willd. Its
recent spread in this country may well have been, nevertheless, from
the Appalachian region, where it is now common (Fig. 3).

1936] McVaugh,— The Taxonomy and Distribution of Lobelia 251

The second species, L. siphilitica, is abundant in the Appalachian
region, but has not spread to any extent into the Coastal Plain, nor far
northeast into glaciated country. It has, however, migrated west-
ward as the var. ludoviciana (Fig. 6), as far as Colorado. In
Wisconsin and Minnesota, southward through Missouri, many inter-
mediates appear; the typical form is not uncommon in the Ozark
region (Fig. 5).

What passes for L. amoena Mx. is a much misunderstood plant of
the Appalachian Mountains and Piedmont (Fig. 7). Small (71)
gives this as a Coastal Plain species, which is obviously an error,
probably based upon the misidentification of plants of L. glandulifera
from Florida. The species itself is confined to the uplands, but the
closely related L. elongata has developed in the eastern Coastal Plain
(Fig. 8). The position of L. glandulifera Small is not wholly clear:
it combines the flower-size and glandular calyx-lobes of X. glandulosa
with the smooth corolla and general smoothness of L. amoena. The
capsule is intermediate (where seen) between those of L. amoena and
L. puberula. It seems best to regard it as a distinct species, close to
Z. amoena, perhaps also related to L. glandulosa. Its range is both
Appalachian and Coastal Plain (Fig. 9).

f) L. puberula is separated from the species in the preceding sub-
group because of the seeds, which seem closer to those of the spicata
complex (Plate 435). It has (in one of its phases) an Appalachian
range (Fig. 12), almost identical with that of L. spicata var. lep-
tostachys. This is evidently the ancestral type; it grades freely into
several forms: one on the Atlantic Coastal Plain (Fig. 13), in which
development has been in the direction of a hirsute calyx, large and
leafy bracts, broad calyx-lobes and obtuse leaves. Southward the
species becomes nearly smooth (Alabama, Mississippi); from Florida
little material has been seen, but it seems to approximate the Ap-
palachian type or that of the Atlantic seaboard. Westward and
southward (Louisiana, Texas, to Arkansas) the species grades into
the form with strongly dentate leaves, large long bracts, but rather
smooth calyx. In other words, the Appalachian or central type of
L. puberula seems to pass into several forms which radiate, as it were,
from a central one.

The two remaining species, L. bremfolia and L. glandulosa, are
closely related to each other, as shown by flower-structure. The
common ancestor, if any, is, however, very much in doubt. The most

r.

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252

Rhodora

[July

probable suggestion seems to be that both have been evolved from a
plant related to i. j^ervh; all have a similar type of pubescence on
the calyx, besides the fact that a number of plants have been seen

rn;rcat:xtsC °' '• ^^^"'" '''' '■ *^^^*' ^••'^•^ -^

Phobable Relations of Species
It seems well at this point to discuss the features of this part of the
genus IoM,„ wh,ch seem important as indicators of rJationshTp
as well as those characters which appear to be primitive or advanced.'

A. INDICATORS OF RELATIONSHIP.

1. Seeds. So far as can be determined, this is the most important
smgle character. Species like L. Kalnui and L. Nuttalli, which we^
confused by the earlier botanists, and considered closely related Z
separated by several good characters and are evidently no ve"
c osely connected. The seed-differences alone, in this case arlso
strikmg as to make this obvious (Plate 435)

of !h Ji?'''!"'' °^ ^°"'"^- '^^^ '°°^' "' '^"^ ""t °f h-i^ ^i the base
of the lower hp m many species, considered in conjunction with other

corolla-characters is often of great help in taxonomy of the group

For example i. Car^yi, |o„g considered close to L. Nuttalli because

str^'etre oftr "', •"'""'; °' ^°"*^' '^'^ '""^ ^^^^ "P »«! «--
structure of the sptcata complex.

3. Other Corolla-Characters. Size of corolla is a weak character in
general, as .s the degree of external pubescence; this applies Iw ,' "
the s.ze and degree of pubescence of the stamens. However the
length, of the corolla, anther-tube, and filament-tube areluv 1
constant m the same species within small limits, and often served
good specific mdicators when used with other characters. Fenestrate
corollas have appeared separately in several groups (including the
Mexican L.f^.,ralrs Cav.), so that this is of general importance oJly
4. Calyx-Characters. General shape of calyx and degree of in^

:rshou,;rb 7 °!, --■''-'">'^ -p-t-e. but^::: ich

stress should not be placed upon them, as they vary even between
different flowers of the same plant. The general form of the X
lobes .s to be considered, but their length is very variable (w dthTlsoT

Character. The presence of auricles at the base of the calyx-lobes is

\

Rhodora

Plafr 436

«

i

L - _d
'■A -

I

I

ll|-|;|l Mi < I'MUS

ll'-ilM.r II.-

'J'ypk of LoBKLiA spicATA Lam.

r»i

f

m

i

19361 McVaugh— The Taxonomy and Distribution of Lobelia 253

evidently an ancestral character which has persisted without any
apparent correlation with other features.

5. Pubescence. The absolute amount of pubescence present is so
variable in the same species that it makes little difference, but the
character of this pubescence is in some cases important. For example,
in the sub-group of L. spicata and its relatives, the base of the stem is
densely short-pubescent. In some forms of L. puberula there is found
very nearly the same type of hairiness, which is further evidence of the
relation suggested by similarities in range and in seed-characters.
Furthermore, certain plants found in Arkansas and Oklahoma are to
be distinguished from L. spicata only by the fact that the stem is
slightly hirsute in lines only, just as in L. appendiculata.

6. Leaf -Characters are so easily influenced by environment that they
are relatively of little importance. The best example of this is fur-
nished by L. Kalmii, in which the leaves vary from linear-filiform to
broad elliptic, depending on the habitat.

7. Inflorescences and Branching. Most of our species have a
definite central axis with subordinate lateral branches, but the degree
of branching is sometimes helpful.

B. ADVANCED OR PRIMITIVE CHARACTERS.

1. Plants with leaves all cauline are considered more primitive in
this respect than those with basal rosettes only; the latter all possess
vestigial cauline bracts, which in some cases develop into leaves.

2. The same reasoning applies to the bracteoles usually found on the
pedicel. It seems logical to assume that they are the remains of more
or less leafy bracts, and are gradually disappearing; this is confirmed
by their absence in such highly specialized spgcies as L. Dortmanna
and L. paludosa.

3. The Occurrence of Auricles. In general, this is probably a
primitive character, in respect to this group. This is confirmed by
their presence in forms like L. spicata var. leptostachys, L, pvberula,
and L. Hald, and their subsequent loss in related and obviously
derivative plants such as L. spicata var. originalis, L. floridana, etc.

4. Separation of Petals from the Corolla-tube. The condition of a
"fenestrate" corolla seems to be an advanced one. If, as is now
mostly accepted by taxonomists, freedom of flower-parts is a primitive
condition, then the corolla-tube of Lobelia must have come from once
separate petals. It is hard to see how the fenestrate condition could
have arisen without the tube once having been entire (Fig. 1). The

254

Rhodora

[July

Fig. 1. Flower of Lobelia, showing Fenestration.

character. * " "' "^ P^^^feally no use as a taxonomic

niot^Sanc^ThlTCer^'''''^' ^* ""* "~"'^' "-»y
of flowers than do the o^rand T"' t^' """ '^■^^"'*^ '" f°™

plant which e JtedTiT™:^,^^^^^^^^^ " ^^^^'"'^ —*-'

time, probably in the rerion^T ^'^*^'=~"« °^ early Tertiary
matter of fact there mavT t '°"*'™ Appalachians. As a

the rest of the group mS^riierth "'" '''''"""^ "'"^'''*^'' ''"^

be designated Lgh'ly L thT :^nt:erdT^^^^^^^^^^^ ""'^

In the latter, L. Cardinal^, ;, , ..° . ''^ f™ the large-flowered.

existed unchanged!::*"?^;;:^^^^^ 'rlT''"''- """^ '"' ^''^^^
L. nphilUica, while L. am^^al 7 , /T ""^ "^ ^^^^ "^
well have «>n,e from a ZZ:. ancetr '"' "^^ ^''"^''^'^'' -''^

flowed Tear trS TZ^T' ''''' T' '^^ "^ "'-^ ^^'^
hairy lower lip, single racCe inV' "'"'■'«»-»'''*« ^o^Ha with
lobes, and broad, cauHnTlers "S- '^^^f^' '""''^'^^ »" ^^e calyx-
one of these devdoped"L;rfll .'"'Z '^'^"^'^ '""> t'^" '■nes;

oped large flowers and a fenestrate corolla, withou

ma, McVaugh,-The Taxonomy and Distribution of Lobelia 255
the hairy lower lip (Z. p^,^„ J i .

source of L. glandulosa and L. irm^fe Th. Tr" ^P°'''"*'
"'■«J»/ofto. Ihe second line failed to

F'O. 2. Diagram of Probable Relationships of North American Spe^

of Lobelia.

develop large corollas, but branched out in several wavs (Fia 2^
The fonns of L. ,^eata make up one branch, a second b:Sfi by

256

Rhodora

[July

1936] McVaugh, — The Taxonomy and Distribution of Lobelia 257

I'

I

L. Gattingeri and L. appendiculata. A third branch is that culminating
in L. paludosa. The line of L. Nuttalli and L. Feayana shows some
resemblances in leaf and capsule to L. spicata, and very possibly is a
minor offshoot of this complex.

Further evidence for the above, although not wholly satisfactory, is
afforded by the fact that supposedly older types, such as have given
rise to wide-ranging varieties and species, are confined in several cases
to restricted ranges; they are not aggressive. It may be that such
plants as L. Gattingeri, L. amoena, L. spicata var. leptostachys, and the
Appalachian representative of L. pvberula are old species which have
passed the colonizing stage of their existence.

General Conclusions

The partly unsupported conclusion reached in this paper is that
from one or more ancestral types living in the Appalachian region of
the Southeastern United States, in Tertiary time or before, have
come a majority of the species of Lobelia now native in this region.
Secondly, that these changes have been brought about by the natural
radial spread of the original species, so that closely related plants are
seen to be occupying different radii of the same hypothetical circle.
These relatives are usually not to be considered cases of simple linear
development, but of parallel development from a common ancestor.
There are several excellent cases in point:

1. The western var. hirtella of L. spicata did not come from the var.
originalis of the eastern states, as has been assumed, but from a plant
like L. spicata var. leptosiachys, in a central position (c/. Fig. 19).

2. The Appalachian phase of L. puberida is replaced in the East
by one derivative, and in the West by a similar but distinct one, both
of which must have come from the first, as neither of the outlying ones,
so far as known, occurs in between (c/. Figs. 12-13).

3. On the Coastal Plain of Florida and Georgia, west about to the
Apalachicola River and the eastern edge of Alabama, occur three
species; L. glandulosa, L. paludosa, and L. flaccidifolia. West of this
line, their places are taken quite abruptly by L. bremfolia, L.floridana,
and L. Halei, respectively: taken in each case by a closely related
species. Wherry, in a geographical study of the southern Sarracenias
(Mss.), finds somewhat the same situation in that genus, and assumes
that these related forms have come separately from a common
ancestor, along one of the many more or less parallel streams leading

V

,4

\

\

1

% f

I

out of the Appalachian region to the Coastal Plain (Figs. 10 and 11.
of. also Figs. 24, 25, 26).

In the foregoing the writer has attempted to show that the North
American Lobelias form a distinct unit, clearly separable from
geographically neighboring ones. It is true that the conclusions
reached here are largely theoretical, but in reaching them every
effort has been made to stay within the bounds of evidence actually
at hand, and those of logic.

In the following pages is given a conspectus of the North American
species, with detailed data of the plants themselves, and their ranges.
No attempt has been made to give the complete synonymy for all
species; only the most important references are included.

In citing herbarium specimens, one record only is given for each
county, except in special cases such as large counties or districts,
little-known species, or areas near the limits of ranges. For the wide-
ranging and well-known species L. Cardinalis L., L. siphilitica L.,
L. inflata L., L. Kalmii L., and L. Dortmanna L. a few citations only
or dots on the maps are given for each state or province. For all other
species and varieties, at least one record is given for each county or
district where the plant is known to have grown. Near the edges of the
ranges of the above five species, all known county records from certain
states have been given; in such cases the citations from that state
are followed by (All).

All other things being equal, specimens with collection-numbers
have been cited ; likewise, those which are represented by duplicates in
several herbaria. Except where noted, the ranges given at the ends
of the descriptions of species have been compiled only from material
actually seen.

Conspectus of the North American Species

Lobelia [Plumier] Linnaeus, Gen. PI. 897. Ed. V. 401. 1754.

Type Species: L. Dortmanna L., Sp. PI. II: 929. 1753. This is
chosen as the type because it was the species best known to Linnaeus
in Sweden, and was mentioned in the "Flora Lapponica."

Not Lobelia Plumier, Gen. 21. 1703; plate 31. { = Scaevola L.).

Dortmanna [Rudbeck] Linnaeus, Syst. Ed. I. 1735 (fide Index
Kew.).

Lobelia Linnaeus, Gen. PI. Ed. I. 267. 1737.

Rapuntium Tournefort, Inst. R. H. 163. 1700. Presl, Prodr. Mon.
Lob. (1836).

Our species annual or perennial herbs, with acrid milky juice con-

r

- A

258

Rhodora

[July

tainin, .0. o. less ^^^^^^^^^^^ tb^"scf fv^ril

pedicels usually. P"^es<^"V J* f main axis, and branches, if
racemose or paniculate; "="^¥ '?™ pio^ers perfect, 5-merous,
any, subordinate and devd^mgater^^* ^ ^„ he

rJ, purplish or blue to ^^'^e Hf^h the stamens just where the
ovary; corolla irregular "?' ^vary " the lobes of the corolla mostly
calyx becomes free from the ovary t ^^ ^^^ ^^^ ^^^^^^

vaivate or induphcate •» *e bud, the tu ^^^ inflorescence;

two of them (in &<^^^^\'XZt the revLal of position is brought
actually the two "^'^^ *e bract je ^.^^ rj^^ tals next to

about by the twistmg of the ped"^' '» j^^ t^be, from below

the cleft often ^^'f K ^InSte Limb bilabiately irre^lar, the
upward, making the tube «"«^*''t„ J; "° ore or less reflexed, usually
three (apparently) lower l°bes spreading^ »^^^^^^^^ ^^^^^^

broad; the two (aPP'''*"'^^!"^^!' X Lower Up hairy at base m
and shorter than those of *« >°^«i,;P^ Pedicel usually with two
some groups, oft«\^"^^^"itve it Stamens as many as the obes
bracteoles near the base "'^/'^y.^j, ' V ^ syngenesious and partially
of the corolla "^"d alternate with ^em sy g^ .^^^ ^ . ^^^

n^onadelphous; »*»>«" ^-^^^H'lj^'fXof white hairs at the tip; the
(apparently) lower smaller, '^'^J jm ^^ ^^^^^ ^ ^

ihVie upper larger, smo.rthpubes^ent^ ^^ ^^^ the'r length

Filaments flat, united above from .^ j^^^ Limb of the

usually hairy below; P";'?'^^*"*' '"'^^^^^ inferior or sometimes

calvx divided down to the ovary, wmcti IS w ,1^ j jt^ normally
Sy free; calyx4obes entire or tooAed^^speci^^^^^ ^^^^^
entire calyx-lobes may ^ave 'ndi™ w^^^^ ^_^^^^^^^ 1

often with appendages at the base. . ^ • 2-lobed, with a
Slcentae. loculicidally 2j^%d;„iSmer^ . anatropous. Embryo
bf stSKtfc:&arendosperm. S.ds small, rough-

ened.'foveolate-reticulate.

27 eastern North American species and vane^-J- . ^^^^.^^^ ^

In addition, the foUowmg ^P«"-J°;,*°™4 United States; they
such) occur in ^^;.:;^Z^^,^X:^et^.e^.r,.,.r^oH^^^
are not --d-^** '" *^P7American. None except the last belongs
ranges are South or ^ent^al Am ^^^rican":

to the group here designated . North^ ^ ^^ ^^^

,.^CSta"y,'l.^— . L, L. .r^in. Cav.. L. X^^
HBK., L. splendens Willd.

*^^

f-/^

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•i

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<>i>

t

fi*^

A^

19361 McVaugh— The Taxonomy and Distribution of Lobelia 259

ANALYTICAL KEY TO SPECIES

1 Flowers larjre, straightened out 18-45 mm. long, including
cSyx. Corolla noVally fenestrate. Terrestrial or swamp
plants with leafy stems. Seeds rough-tuberculate (2).
2 Flower crimson (white forms occurrmg as sports), dU-40
mm. long, including calyx (3).
3. Anther-tube 4.0-6.5 mm. long. Filament-tube 24r-33
(usually 28-30) mm. long. Leaves lanceolate to
ovate-lanceolate, three times as long as wide or less,
1 5-6 0 X 6.0-18.5 cm. Smooth or sparsely hirsute-
pubescent. Plants of the eastern half of the con^ Cardinalis.

3. A^the?-tube '3.5^!5 "mm. long.' Fiiament^tube i9-23
(26) mm. long. Leaves lanceolate to bnear-knceolate.
about seven times as long as wide, 0.4-3.0 X o-^-^^-"
cm. Smooth or sparsely pubescent. Plants of south-

western United States and Mexico ^ • • • • (1) ^- splendens.

(The related rough-pubescent L f^ens WiUd., of
Mexico, has not been seen from the United States).
2. Flowers blue or violet (white forms occurring as sports),

18-33 mm. long (4). ^ i i u „,;+u

4. Filament-tube 12-15 mm. long^ Calyx-lobes with
broad, leafy auricles at their bases (auricles ovate-
obtuse to -acute, 2-3 mm. long, not glandular-dentate).
Pedicels with a pair of conspicuous bracteoles just
below th^ calyx or H-'A the length of the pedicel

a!" Whole pknt more or less hairy. Calyx and its lobes
hirsute. Inflorescence usually long a^d dense
Leaves broad-ovate or -lanceolate, 2-6 X 6-l»
'°^-)^''l^tem Plants oft^ntaU (75-100^ ^^^^^.^^^

aa. PlSnearly^sm^thiusuaily 30-60 cm. high Calyx
and its lobes sometimes sparsely hirsute. Inflores-
cence shorter (6-20 flowers). Leaves smooth,
lanceolate, about 1.5 X 6.0 cm., shallowly toothed
or subentire. Plant of mid-western United States.

(2) L. si-phihiica, var.

4. Filamentltube'e^ii mm.' long'. Pedicel with a Pair of
bracteoles at or near the base. Auricles of the calyx

present or absent (5). . ., i j i... j««*of^

6. Salyx-lobes (usually) prominently glandular-dentate

or pectinate, never hirsute. Flo>yers few, 3-20 (27 ,

in loose, secund racemes. Pedicel stout, m fruit

usually stiffly upright (b). u^^„j

b Calyx-lobes pectinately toothed. Auricles broad,

leafy, round, fimbnate, nearly covering the

hemispheric calyx. Leaves numerous, to 200,

small, narrow, obtuse to 0.5 X |0 cm^^^Pro^"

nently denticulate. Flowers &-20, 18-20 mm.

long, hairy-strigose outside. Lower lip of

cor&la smooth or nearly so. Filament-tube

about 7 mm. lo°g ••; • ■ •,• • j' ; ' ' ;';.;i;^ «i.
bb. Calyx-lobes prominently glandular-toothed, or
nearly entire. Auricles none or very small, tri-
angular. Flowers large, 20-33 mm. long, smooth
outside, the corolla-lobes about equahng the
tube in length (c).

ludoviciana.

I

260

Rhodora

[July

« Plflnt« usually weak; leaves 5-10, long-linear, to

c. ^^^^^^'^^^ ^^ally prominently denticula e.

Flowers 1-10 (15), 20-33 mm long Lower hp

of corolla hirsute at base Filanaent-tube 8-10 ^

mm long. Calyx often chaffy-hirsute • ■ • • • • 6. L. glaruiulosa.

cc pS slender or erect, smooth throughout^

rX ^^?am^iTtur7-M^^^ .^an.^^/-

^re or less secund racemes. Pedicels m fruit

curved to one side (6). narrowly

fi Plants smooth or nearly so. Calyx-lobes narrowiy

hnear-^nceolate, about 1 mm. broad, smooth.

Sles none or'very smaU.. Calyx campanulate

in flower, becoming globose m fruit (d).

A Leaves narrowly lanceolate, to 1.5 X 10.0 cm.,

FUament-tube 5-7 mm^ ong ^Anther-tube
2.5-3.5 mm. long. Calyx-lobes 5-11 mm-
long Species of mountains and Piedmont

southeastern United btates . , • * ;

6. Plants more or less fhort-Pubescent t^^ghou^^
nalvx-lobes lanceolate or broader, i-4 mm. wiae,
^irmm long, ciliate-pubescent, usually more or
kiaSSulSt the base. Calyx flat or turbinate
in flower, becoming conic-hemispheric m frmt (e).
e Flower-bracts usually leafy, broad-ovate, to 1£
X 20 cm. Calyx usually densely hirsute-
chaffy. Calyx-lobes broad at the base, ovate-
trianLlar, to 4 X 12 mm., the edges much
roUed back, especially in f "lit, formmg large
rounded auriclS. Leaves obovate-obtuse
below, coarsely toothed, ovate above. Plant
of Cokstal Plain and adjoinmg territory, Ga.
?o NX In the region from Texas to Ark. and
Mo"^ a similar plant with smoother calyx and

strongly dentate leaves ■, • * ' : ' iul P^^^

ee Flowe?-f)racts lanceolate, 1-2 cm long m the
^' "^ lower flowers. Calyx usually pubescent^^^^^^^
times glabrate rarely hirsute). Calyx-lobes
lanceolate, to 2 X 12 mm., httle roUed at the
S even in fruit. Auricles none, or small,
Sgulir Leaves 10-20 oblong acute, or
obtuse below, mostly sharply dentic^ate.
Plant of uplands, Ohio to Ga. and westward. ^ ^^^^

2. Hants aquatic; leaves fleshy, Unear, hoUow, forming a

•- ♦

^ ,-<*>*

t^

!<
V

1936] McVaugh,— The Taxonomy and Distribution of Lobelia 261

basal rosette. Scape nearly naked, few-flowered.
Anthers all densely tufted at tip. Capsules long-stalked,

pendent. Occurring north of the moraine 22. L. Dortmanna.

2. Plants aquatic or terrestrial, leaves flat. Stems leafy or
sometimes leaves nearly all radical (3).
3. Plants with slender, more or less delicate stems and
narrow leaves, seldom over 50-60 cm. high. Base of
lower Hp of corolla smooth (4). , j

4. Flower 10-13 mm. long, calyx elongate, capsule ovoid.
Pedicel with a pair of sub-opposite bracteoles about
the middle. Plants of calcareous bogs and rocks, ^ , ..

north of moraine -21. L- Kalmiu

4. Flower 7-10 mm. long. Calyx flat or conic. Bracteoles
of the pedicel at its base (5).
5. Plants 20-60 (75) cm. high, erect. Leaves lanceo-
late, the basal spatulate. Calyx flat; capsule

hemispheric, half inferior, often bristly 19. L. NuttalU,

5. Plants 10-30 cm. high, weak, decumbent. Leaves
sub-orbicular and petiolate below. Capsule tur-
binate, acute at base, % or more inferior, smooth.

Peninsular Florida 20. L. Feayana.

3. Base of lower lip of corolla densely haio or rarely nearly
smooth. Plants not delicate, often tall (75-125 cm.);
usually not diffusely branched. Leaves broad or some-
times linear (6). i n >i
6. Leaves linear-lanceolate or filiform, cauline rarely 0.4
cm. wide. Inflorescence loose, mostly branched (7).
7. Pedicels and calyx smooth. Bracteoles of pedicel
none. Usually much branched above; aquatic,
with leaves often deciduous, flowering May-June. ^
12. L. Boykimt.

7. Pedicels and calyx scabrous. Bracteoles at base of

pedicel. Simple or somewhat branched, leafy;

not aquatic, but living in swamps. Flower ^ _ _ , .

August-October ... 11. L. Canbyt.

6. Leaves broad, seldom less than 1 cm. wide. Inflores-
cence not diffusely branched (except in L. inflata)

(8).

8. Caiyx ovoid; capsules developing early, much in-

flated, ovoid, inferior. Usually much branched,
especially in &ge. Stem usually long-hirsute. . .10. L. inflata,
8. Plants never diffusely branched (sometimes with
few subordinate side branches); inflorescence a
terminal spike or raceme. Stem never long-
hirsute. (Japsules various, never much inflated

(9).

9. Leaves strap-shaped or oblanceolate, mostly

basal. Bracteoles of the pedicel inconspicuous

or none. Semi-aquatic or swamp plants of the

Southern Coastal Plain (10).

10. Plants tall, 80-100 cm. Filaments 7-9 mm.

long, deflexed. Corolla-tube not fenestrate,

but often with a thin place on each side of

the wall. Pedicels with inconspicuous

bracteoles 17. L. floridana.

10. Plants 50-60 cm. tall. Filament-tube about
3.5 mm. long. Corolla-tube fenestrate. No

bracteoles visible on the pedicel 18. L. palvdoaa,

9. Leaves mostly cauline, or, if radical, broad-ovate,

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11.

Rhodora i^^^

petiolate. Terrestrial plants of wet or dry

places (11). 1 X 1

11. Pedicel with two conspicuous green bracteoles
about half way up. Auricles of calyx de-
flexed, round, small. Plants of the southern
Coastal Plain (12).

12. Plant nearly unbranched, with thick oblan-

ceolate or lanceolate acute leaves. Flower

19-20 mm. long, pubescent. Filament- , „ , .

tube 6-8 mm. long • • • • lo- ^- ^«*«*-

12 Plant simple or branched, with thin oblong
usually obtuse leaves. Flower 15-16 mm.
long, nearly smooth; corolla-tube some-
times fenestrate; filament-tube 5-6 nun.

long 16- ^' flacctdyoha-

Pedicels with bracteoles at base. Auricles
various (13).

13. Stem-leaves thin, sessile with a broad base,
short-ovate, neariy smooth. Basal leaves
small or none. Stem neariy smooth, even
at base. Raceme more or less plainly
secund (14).

14. Calyx-lobes smooth; auricles none; cen-

tral Tennessee -14. L- GaUtngen.

14. Calyx-lobes strongly glandular-ciliate.
Auricles glandular-cihate, very smaU or
larger, foliose, scarious-tipped . . . 13. L. appendtculata.
13. Stem-leaves ovate or oblong to lanceolate,
somewhat pubescent and narrowed at
base. Stem densely short-pubescent be-
low. Inflorescence a terminal unbranched
spike, not plainly secund (15).
16. Basal leaves large, roundish, conspicuous;
the cauline 1-5, very small, bract-hke.
Raceme loose, nearly half the height of
the plant. Auricles of calyx evident,
but not long-filiform. . . .9e. L. ajricata, var. acapoaa,
15. Leaves mostly cauline; if basal, rarely
roundish and usually with cauline
leaves also present (16).
16. Plants strongly rough-pubescent, in-
cluding stem, bracts, and the long
calyx-lobes. Lower bracts leafy.
Plants often short (20-50 cm.), with
leaves low on the stem. Auricles

small or none 9c. L. spicatat

16 Plants smooth or pubescent, leafy,
often 50-100 cm. high (17).
17. Auricles long-filiform, deflexed, often
as long as the calyx-tube. Inflo-
rescence usually a dense narrow
spike. Leaves oblong, more or less
appressed. Plants sometimes cil-

iate 9a. L. spicata, var. lepioatachya.

Auricles very short or none. Plants
usually smooth. Inflorescence a
terminal raceme or spike, usually
much less than half the height of
the plant (18).

var. hirteUa.

17.

1936] Eighth Report of Committee on Plant Distribution 263

18. Anthers blue; calyx in anthesis
flattish. Flower light blue.
Raceme dense, many-flowered.

Capsules short-hemispheric

9b. L. spicata, var. origincdis.

18. Anthers white. Calyx in anthesis
roundish. Flowers dark pur-
plish-blue. Raceme few (10-30)
-flowered. Capsules globose, of-
ten somewhat inflated

9d. L. spiccUttf var. campanvJata,

(To he contimied)

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STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES OF

LOBELIA

Rogers McVaugh

{Continued from page 263)

1. L. Cardinalis Linnaeus, Spec. PI. II: 930. 1753. Type

Locality: "Habitat in Virginia." Type Specimen: In Linnaean

herbarium in London; seen by Linnaeus before 1753. Photograph

seen.— Trachelium Americanum flore ruberrimo, Parkinson "Para-

disusTerrestris";356. 1629. i^/o* Cflrc?ma/w, Rivinus, " Introductio

Crenerahs m Rem Herbarium," with plate, 1690. Cardinalis Rimni,

i

Fig. 3. Range of Lobelia Cardinalis.

Rupp, Flora Jenensis"; 242. 1718. Rapuntium maximum, coccineo
spicaio flore, Tournefort, " Institutiones Rei Herbariae"; 163. 1719.
Plate 51.— Stem erect, unbranched, coarse (sometimes 1.5 cm in
diameter at the base), green, usually dark purplish-red below, some-
times purple-flecked or purplish throughout, 40-180 cm. high, smooth
or short chaffy-pubescent. Cauline leaves 10-30, spreading, thin
or papery, smooth or short bristly-pubescent, sub-entire in outline
but very irregulariy coarsely or finely dentate, the teeth callose-tipped-

i

1936] McVaugh,— North American Species of Lobelia 277

size 1.5-4.0 (6.0) X 8-12 (18) cm., often three times as long as broad
or longer; lanceolate or lance-ovate to oblong, less often ovate; usually
acute at the tip, narrowed at the base, the lower short-petiolate.
Perennial by offsets. Roots fibrous. Inflorescence a terminal raceme*
unbranched, few-50 cm. long, not noticeably secund, densely (or
loosely) few-100 flowered. Pedicels more or less upright, slender,
4-14 mm. long in fruit, short bristly-pubescent, each with a pair of
bracteoles at or near the base. Flower-bracts linear or the lower
lanceolate, leafy; smooth or neariy so, with prominent callose teeth,
1-5 cm. long. Calyx in anthesis conic or short-campanulate, smooth
or somewhat pubescent, becoming cup-shaped or hemispheric in
fruit, strongly ribbed, usually broader than high, 8-11 mm. across.
Capsule about half inferior. Calyx-lobes linear-subulate, with a
short-deltoid base, smooth or ciliate at the tip, 8-16 mm. long.
Auricles none, or minute, triangular. Flower 30-45 mm. long, includ-
ing calyx. Corolla deep crimson (pink or albino forms occur rarely),
somewhat puberulent, the lip smooth. Corclla-tube fenestrate; lobes
of the lower lip spreading, deflexed, ovate, acute, narrowed at the base,
nearly equalling the tube, 3-5 X 13-20 mm.; the two upper lobes
erect, linear, 1-2 X 13-20 mm. Filament-tube 24-33 mm. long
(ave. 28-30 mm.), much exceeding the corolla-tube, red, pubescent
below, connate above more than half its length. Anther-tube 4.0-5.5
mm. long, bluish-gray, the two smaller anthers white-tufted, the three
larger smooth or lightly pubescent.— Coastal swamps, river banks,
borders of lakes; sometimes in open swampy places; a plant of neutral
soil, penetrating acid-soil and dry regions only along river systems.
New Brunswick and Ontario to Minnesota, south to Texas and
Florida; west of the Mississippi only along rivers (reported from
Nebraska by Petersen); throughout the range, but local or absent
from large areas such as the Pine Barrens of New Jersey, where con-
ditions are unfavorable. — Flower: late July-early Sept. Fruit:
mid-Aug.-Oct. The species is so definite and the range shown in such
detail in the map (fig. 3) that the citation of specimens is unnecessary.

From Kansas and Texas westward the closely related L. splendens
Willd. is to be distmguished by the (usually) narrower leaves (some-
times narrowly linear), and by the smaller flowers (filaments 20-24
mm., rarely longer, little exceeding the corolla-tube; anther-tube
3-4 mm. long).

The following material has been seen of the closely related plant
which is apparently L. splendens Willdenow:

Missouri: jackson: Courtney, Btish 294 (Mo). Nebraska:
HITCHCOCK: Culbertson, Wagner, Aug. 1911 (W). Kansas: Mcpher-
son: "Linsborg," Bodin, Jy. 1887 (M). riley: Hitchcock 816a
(G, Mo, NB, R). Oklahoma: blaine: Watonga, Stratum 481 (Mo).
CLEVELAND: Normau, Bruner, Sep. 1924 (W). osage: Pawhuska,
SUtens 1993 (G). payne: Stillwater, Waugh 259 (Mo), woods:

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Alva, B^ietens 28S6 (G, M). Texas: "between Ft. Leavenworth and
El Paso," Diffenderfer, ann. 1871 (ANS). bell: Holland, Mackensen
£35 (Mo). BEXAR: San Antonio, Bush 1266 (Mo), comal: New
Braunfels, Lindheimer, Sep. 1850 (ANS, UP), culberson: Guadalupe
Mts., Moore and Steyermark 3613 (ANS). Gillespie: Jermy herb. 761
(Mo). OLDHAM: "1 mi. N. Canadian R., on Amarillo-Dalhart rd.",
Ferris and Duncan 3481 (Mo, NB). wilson: Sutherland Springs,
Palmer 2090 (ANS). Colorado: yuma: Wray, Shantz, Sep. 1907

Fig. 4. Northern Extension of Lobelia splendens.

(NB). Utah: Washington: Springdale, Jones 6077 (Del, R); Zion
Natl. Park, Pilshry, Aug. 1925 (ANS). New Mexico: bernalillo:
Albuquerque, W. Harward (Mo), chaves: Roswell, Earle 497 (M).
DONA ana: Organ Mts., WooUm 10644 (NB); "Donana," Parry
et al. (Mex. Bound. Surv. 694) (ANS). uncoln: White Mts., WooUm
202 (M, R). sierra: Kingston, Meicalf, Aug. 1904 (W). Arizona:
COCHISE: Huachuca Mts., LemvKm herb. 2806 (ANS, CM); Paradise
Falls, Blumer 1731 (W). California: san Bernardino: San Ber-
nardino Mts., Abrams 2937 (ANS). Chihuahua: Sierra Madre,
Pnngle 2287 (ANS); Cumbre, Palmer 368 (ANS); Chuichupa,
Tovmsend and Barber 427 (R).

2. L. siPHiuTicA Linnaeus, Spec. PI. II: 931. 1753. Type Local-
ity: "Habitat in Virginia." Type Specimen: in Linnean herbarium
in London; seen by Linnaeus before 1753. Photograph seen. Rapunt-
ium Americanum, /lore dilute caeruleo, Dodart, "Memoires" 105.
1676 (ace. to Toumefort, Inst. R. H. 163. 1719). Rapunculus
galeaius, Virginianus, flore violaceo, majore, Morison, "Plantanim
historiae"II:466. 1680. (This is i>ossih\y L. piiberulaMx.). Lobelia
cavle erecto, foliis ovato4anceolatis crenatis, floribus kUeralibus, Linn-

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1936] McVaugh,— North American Species of Lobelia 279

aeus, Hort. Cliff. 426. 1737. The name " siphilitica" may have been
suggested by Kalm (1750) (32). L. antisiphilitica Hayne, Arzn.
Gewachse. XIII: plate 9. 1837.— Stem erect, unbranched, rather
coarse, 20-130 cm. high, light green, quite smooth or sparsely chaffy-
hirsute, especially on the angles formed by the decurrent leaf-bases.
Cauline leaves few-25, usually loosely spreading, very thin and papery,
the lower narrowed into margined petioles; in shape obovate, oblong,
ovate or ovate-lanceolate, usually short-acute at the tip, 2.0-4.0 (6.0)
X 6-12 (18) cm., nearly smooth beneath, sparingly strigose above;
sub-entire in outline, or more or less coarsely serrate, the teeth callose-
tipped; upper leaves usually merging gradually into the bracts of the
inflorescence. Perennial by offsets. Roots fibrous, rootstock thick.
Inflorescence a terminal raceme 10-30 (50) cm. long, usually densely
6-75-flowered, not secund. Pedicels loosely upright, 5-10 mm. long in
fruit, more or less flattened, smoothish or chaffy-hirsute, each with a
pair of conspicuous bracteoles just below the calyx or 1.5-4 mm.
below. Flower-bracts smooth or somewhat ciliate-fringed, the lower
(sometimes all) leafy, the upper usually smaller, lanceolate, 1-2 cm.
long. Calyx in anthesis flattish-hemispheric, usually more or less
chaffy-hirsute, becoming hemispheric in fruit, somewhat flattened
(broader than high), 8-10 mm. in diameter. Capsule }^^ inferior.
Calyx-lobes foliaceous, broad-lanceolate or ovate, acute or acuminate,
often 5-6 mm. wide by 8-11 (14) mm. long, the margins usually much
folded back, ciliate and serrate, undulate or crisped. Auricles folia-
ceous, flat, small or covering the entire calyx, obtuse or acute, some-
times connate, 2-5 mm. long. Flower 23-26 (33) mm. long, including
calyx. Corolla bright blue (albino forms sometimes occur), white-
striped in the throat; base of the lower lip white, with two raised
tubercles; corolla smooth or hirsute on the veins outside. Corolla-
tube fenestrate; lobes of the lower lip narrow-ovate, short-acute,
sharply deflexed at base, about half as long as the tube, connate below
or nearly to the tip: two upper lobes long-acuminate, about as long
as the lower. Filament-tube 12-15 mm. long, pubescent below,
connate above more than half its length, somewhat deflexed. Anther-
tube 4.0-5.5 mm. long, bluish-gray, the two smaller anthers tufted,
the three larger smooth.— Moist woods and swampy places; often in
light shade; less frequently by streams or in open wet places; a plant
of neutral or somewhat calcareous situations. Maine and southern
Ontario to eastern Minnesota, south in the Mississippi Valley to
Tennessee; common in the Appalachian region, south to Alabama
(possibly Mississippi and Louisiana); rare or absent on the south-
eastern Coastal Plain. Not common in New England. Flower
Aug. lO-Sep. 20. Fruit September to mid-October. Representative
material seen: Ontario: grey: Hanover, Hauch, Jy. 1895 (W).
HASTINGS: Madoc, hey, Aug. 1906 (Toronto), huron: Wingham,
Morton, Jy. 1890 (NB, US). Middlesex: London, Millman, herb.
G. S. Can. 15259 (O). welland: Niagara Falls, McCalla 413 (O).

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YORK: Toronto, HoUingsworth 1601 (Toronto). Maine: aroostook:
Fort Fairfield, Furbish, ann. 1881 (NE). Cumberland: Falmouth,
Norton 918 (NE). sagadahoc: Topsham, Hutchins, ann. 18— (NE).
(ALL). Massachusetts: Berkshire: Sheffield, Hoffmann, Sep. 1912
(NE). MIDDLESEX: Cambridge, "Tin Canon," introduced, Femald,
Sep. 1891 (G, NE). (ALL). Connecticut: fairfield: Sherman,
Winton, Aug. 1885 (G, US). Litchfield: Salisbury, Weatherby, Sep.
1914 (US), new haven: Derby, Oakes, Aug. 1828 (NB). (ALL).
New York: monroe: Ontario Beach, Bartram 1797 (ANS). st.
LAWRENCE: Stockholm, Phelps 1752 (G). tioga: Apalachin, Fenno

Fig. 6. Range of Lobelia siphilitica.

261 (NB). WARREN (?): Lake George (no county given), Mrs,
Watrous, ann. 1895 (NB). New Jersey: burlington: Moorestown,
Hollinshead (UP), somerset: Watchung, Moldenke 6391 (NB).
Beside the above, known only from bergen, hunterdon, Sussex,
warren cos. Pennsylvania: Bradford: Sayre, Barbour 966 (R).
clarion: Lawsonham, Bright 731^2 (W). york: McCalls Ferry,
Heller 1280 (ANS, G, NB, US). Delaware: Newcastle: Centre-
ville. Commons, Sep. 1878 (ANS). (ALL). Maryland: cecil:
Fairhill, Benner 6321 (ANS). garrett: Grantsville, Stone, Aug. 1911
(ANS). MONTGOMERY: Great Falls, Holm, Sep. 1915 (G). District
of Columbia: Washington, Mohr, Sep. 1882 (US). Virginia:

1936] McVaugh,— North American Species of Lobelia 281

FAIRFAX : Great Falls, ^Fzmer 432 (Duke, UP), james city: Williams-
burg Grimes 4580 (M). paoe: Luray to Stony Man, Tidestrom 6709
(Ub). West Virginia: barbour: Tygart Jet., Moore 258A (G)
monroe: Sweet Springs, Steele 21,5 (G, NB, US). North Carolina*:
HAYWOOD: WaynesyiUe, J527<more herb. 627a (G, NB, US), rocking-
^'' i?n^^Vx?t^^^'^''^ (^^)* Alabama: choctaw: Cocoa, SchucheH,
/ix tJon ^^^^' JEFFERSON: Birmingham, Schuchert, Oct. 1896
m, US). ST. clair: AshviUe, Mohr, Sep. 1899 (US). Talladega:
Talladega Creek, Mohr, Sep. 1892 (US). (ALL). Louisiana: In
the AJNb collection is a nearly smooth specimen perhaps from this state:
It IS majked Tamtuner'' (perhaps an error). Kentucky: bell:
CuEciberland River, Kearney 462 (M, NB). Harrison: Lair (Dono-
van) Pike, Singer 458 (CCD). nelson: Chaplin, Pennell 13670 (ANS)
Tennessee: cheatham: Craggie Hope, Svenson 314 (G). cocke-
Lemons Gap, Kearney 807 (M, NB, US), shelby: Memphis,'
Fendler, Sep. 1853 (G). The last is smaller and smoother than usual
and was identified by Asa Gray as var. ludoviciana, but is closer to the
m^^Jfr, N • ^^^^- HAMILTON: Cincinnati, Stephenson, Oct. 1930
(K, UGa). ^ licking: Granville, Jones 1352 (R). Ottawa: Bay Point.
Eames and MacDaniels 273 (UP). Indiana: blackford: Mollie
Deam 334 (US), daviess: Washington, Beam 53259 (CCD) st'
JOSEPH: South Bend, Deam 55578 (CCD). Illinois: cook: nr.*
/Ax?5P' ^'^^^^^''^ ^599 (G). PULASKI : Mounds, Palmer 1656 A
(ANS). stark: Wady Petra, Chase 178 (ANS). vermilion: Catlin,
i?^?f^T^^{.^^^' US, W). Michigan: bay: Kawkawlin, Dreisbach
5476 {ANS, {JF). J)ELTfL:Esc8inah8i, Henry 206 (VP). washten/iw
.rii.^^li?x^' ^''^'^''^^^ ^^^ (US). Wisconsin: grant: Kieler, Fassett

iofJ^S^?- ^^^'''' ^^''^''^ ^^^^ ^^^ (W)- Lincoln: Merrill, Cheney
2842 (W). polk: St. Croix Falls, Fassett 8195 (W). Minnesota-
naoka: Ham Lake, Costing 2987 (approaching the var.) (Duke)
GOODHUE: Zumbrota, BcUlard, Aug. 1892 (NB, US, W). Houston'
Jefferson, Lyon 310 (M). wabasha: Kellogg, FasseU 3319 (W)
WINONA: moist meadows, Holzinger, Sep. 1886 (M). (ALL). Iowa-
benton: Vinton, Dams (W). decatur: Anderscm, Sep. 1904 (R)
JOHNSON: Iowa City, Somes 3597 (US). Poweshiek: Grinnell, Jones,
ann. 187- (R). (ALL). Missouri: jackson: moist ground. Bush
334 (G, M, NB, US), marion: Hannibal, Davis 1220 (G, NB)
RALLs: nr. Oakwood, Davis, Sep. 1916 (M). st. louis: "E^idroits
hMmides;' Riehl 315 {NB). stoddard: Dexter, ^i/^A ^P^7 (NB US)
stone: Galena, Palmer 4614 (R). (ALL). South Dakota: Minnehaha!
feioux Falls, Thornber, Aug. 1892 (G); this plant has smooth leaves,
but otherwise resembles closely the eastern form. (ALL).

West of the Mississippi River this species runs into a variety,
passing as var. ludoviciana A.DC, distinguished as follows:

Var. LUDOVICIANA A. DeCandolle, Prodr. Syst. Veg. VII: 377 1839
Type locauty: "in Louisiana (Tainturier)." Type Specimen:

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[August

authentic material not seen. DeCandoUe's description is as follows:
"caule glabro, foliis lanceolatis subintegris glabris, calyce glabrius-
culo." The plant now passing as var. Ivdomdana A.DC. is a western
one of prairie and mountain regions, perfectly distinct, and fitting
DeCandoUe's description. However, Tainturier is known to have
collected largely on the Coastal Plain of Louisiana (ace. to Pennell),
and there is a specimen of Lobelia siphilitica in the Academy of
Natural Sciences of Philadelphia, marked "Louisiana, Tainturier,"
which resembles rather material from central Alabama than that from
the Northwest. Furthermore, Louisiana a century ago was a much
more extensive territory than today. Any positive settlement of the
identity of this variety must await examination of authentic material.
—L. Bollii Wimmer, Fedde Rep. Spec. Nov. XXVI: 3. 1929.— Stem
often shorter than in the typical form, 30-60 cm. high (rarely 90 cm.
or more), smooth (rarely short-hirsute). Leaves smooth, sub-entire
or shallowly toothed, usually oblong-lanceolate, acute at both ends,
averaging about 1.5-2.0 X 6.0 cm. Inflorescence often fewer-flowered
than in the typical form. Flower-measurements about as in the
typical form.

Differs mainly by the shorter average size, smooth leaves and stem,
smaller and definitely narrow leaves, smoothish calyx, and the nar-
rower, acute, and often connate auricles. Many intermediate plants
appear in the Mississippi Valley.

Low places in prairies, sandy or gravelly margins of ponds and
streams, wet meadows, sometimes on limestone cliffs; Wisconsin and
Minnesota to Manitoba, south and west to Colorado, Oklahoma and
Texas; prairies and mountains. Flowering period about as in the
typical form. Representative material seen: (The citations marked
with an (*) are those of plants somewhat intermediate between the
variety and the type). Illinois: cook: Des Plaines, Strahler, Sep.
1908 (part) (W*). (ALL). Wisconsin: burnett: Hertel, Austin L.,
Fassett 8181 (W). dane: Madison, Sumner, Aug. 1895 (W). polk:
Star Prairie, McLaughlin 1210 (W). ST. CROix: Hudson, McLaughlin
1211 (W). WASHBURN: Spooner, McLaughlin 1207 (W*). (ALL).
Minnesota: becker: DeSoto Lake, Grant 3069 (ANS, G, M, US).
GOODHUE: wet meadows, Sandberg, Jy. 1886 (M*). hennepin:
Minneapolis, Sheldon 1662 (M). Fort Snelling, Meams, Aug. 1888
(NB*, US*). LINCOLN: Lake Benton, Sheldon 1322 (M). ramsey:
Snail Lake, Jackson, Sep. 1926 (UP*), st. louis: Tower, Lugger,
Jy. 1891 (M*). WINONA: Winona, Holzinger, Aug. 1888 (R*). Iowa:
emmet: Armstrong, CraUy, Aug. 1900 (US*), fayette: Fayette,
Fink, Jy. 1894 (M*) johnson: Somes A3017 (US*). Plymouth:
Akron, Bredall, Sep. 1909 (US*). (ALL). Missouri: greene:
Springfield, Standley 8464 (US). (ALL). Arkansas: Baxter:
Baxter, Palmer 4741 (R). benton: Plank, ann. 1899 (CM*, NB*).
garland: "close to the Hot Springs, Ark.", Engelmann, Sep. 1835 (G).
(ALL). Oklahoma: ottawa: Ottawa, Stevens 24I6 (G). (ALL).

1936] McVaugh,— North American Species of Lobelia 283

Texas: Dallas: Dallas, Reverchon, Sep. 1875 (G). (ALL). North
DAKOTA:RANSOM:Lisbon,Ae/c^*<ac;, Aug. 1898 (R). (ALL). South
Dakota: Cascade Falls, limestone cliffs, Mcintosh 81 4 (R). custer-
Woodplam Battle Creek, Over 13746 (R). Pennington: Rapid City
Williams, Aug. 1892 (US*). Roberts: Big Stone Lake, Over 14298
(US). Nebraska: holt: Beaver Creek, Clements 2857 (Del, G, M

Fig. 6. Range of Lobelia siphilitica, var. ludoviciana.

NB, US). LINCOLN: North Platte, Shear 474O (US), scotts bluff:
Scotts Bluff, Hildreth 620 (R). Kansas: douglas: Lawrence, Stevens
(US*). KINGMAN: Calista, Carleton 550 (US), riley: wet places
A^orfon 5/7 (G, NB, R, US). (ALL). Colorado: custer: Wet Mts.,
Brandegee 813 (ANS, NB). weld: New Windsor, Osterhout, Aug. 22,
1902 (ANS, G, R, W). Manitoba: souris: "Turtle Mt., N.W.T.",
T.J.W.B. 139, Jy. 26, 1874 (Toronto). (ALL).

3. L. AMOENAMichaux, Fl. Bor. Am. 11:153. 1803. Type Local-
ity: presumably in Carolina. There is no material of this species in
the Michaux herbarium in Paris; the original description, which is
given wholly without additional data as to locality or habitat, may
apply to the species here included under the name; it applies equally
well to L. elongata Small. The description follows: "L. majuscula,
erecta, glaberrima; foliis lato-lanceolatis, serratis: spica multiflora,

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secunda: calycis laciniis integerrimis : corollae coenilae laciniis inferior-
ibus ovahbus, acutis."— Z. siphilitica ?, Walter, Fl. Car. 218 1788
referred by Gray to this species, was probably L. elongata Small!
as was the L. puberula var. glabella, Elliott, Sk. Bot. S. C. & Ga. 1: 267.
1821. Not L. amaena A. DeCandolle, Prodr. Syst. Veg. vfl- 377
which is a form of L. puberula Mx. L. glandulosa var. glabra A De-
Candolle, 1. c. 378 (ace. to Gray).— Stem upright, unbranched, rather
coarse, 30-120 cm. high, light green (sometimes reddish at base)
smooth or somewhat hirsute-pubescent below. Leaves cauline, rather
widely spaced, 6-15, very thin in texture, acute at the base; the lower
sub-petiolate; smooth, or pubescent mostly on the veins beneath
and strigose above. Lower leaves sub-entire or somewhat serrate
oblong or ovate, usually obtuse, from 1.5 X 4 to 4.5 X 12-18 cm'
Upper leaves narrower, sometimes lanceolate, often prominently
denticulate. Bracts (sometimes with the exception of the lowest)
not leafy, giving the inflorescence a naked appearance. Inflorescence a
terminal raceme, 10-40 cm. long, usually strongly secund. rather
loosely few-40 flowered. Pedicels rather slender, curved, 3-5 mm
long m fruit, short prickly-ciliate or smoothish, each with a pair of
rather prominent bracteoles (sometimes 1 mm. long) near the base.
J^ lower-bracts smooth, callose-denticulate, linear, 1-2 cm. long- the
lowest often leafy lanceolate or broader, 2-4 cm. long; sometimes all
bracts leafy. Calyx m an thesis hemispheric, smooth or with a few
hairs, becoming globose or somewhat flattened in fruit, widest about
the middle or below, 5-8 mm. across by 4-7 mm. high. Capsule
mostly mfenor Calyx-lobes smooth, entire, 5-12 mm. long (ave.
K^^ nim.}, flat, linear, less than 1 mm. wide, or the base broad, short-
deltoid, the upper portion linear. Auricles none or very small
flower 18-24 mm. long, including calyx (ave. 20-22 mm.). Corolla
bright blue, with a light eye, smooth. Corolla-tube fenestrate; lobes
of the lower lip broad-ovate, obtuse, sometimes even spatulate, often
apiculate, sometimes as long as the tube; upper lobes lanceolate
Jiilament-tube 5-7 mm. long, pubescent below, connate only about a
third of Its length above. Anther-tube 2.5-3.5 mm. long, light bluish-
gray, the two smaller anthers tufted, the three larger smooth or
pubescent on the backs.

This species is readily distinguished from L. puberula and its forms
by the general smoothness, the round calyx, narrow calyx-lobes
broad entire leaves, and the somewhat larger corolla, which seems
even larger in proportion because of the small calyx-lobes and bracts
However, it is very possible that hybridization occurs where the two
grow together; plants are found with intermediate leaf-characters
calyx-characters, and pubescence.— Swamps, moist rich woods or
wet rocks, western North Carolina and eastern Tennessee south to
northern Georgia and Alabama. Mountains and Piedmont. Flower
August-October. Representative Material seen: Tennessee: knox-
KnoxviUe, Ruth, Sep. 1895 (Mo); this plant is on a sheet with two

1936] McVaugh,— North American Species of Lobelia 285

other species, indicating a possibility of confusion. Alabama: clay-

7'![S^Z^^^'^' }^^^ ^^^- ^^^'- Auburn, Earle and Baker, Oct. 1897
(M NB). Georgia: clarke: Oconee Heights, Reade, Oct. 1928
<UGa). COBB: Marietta, Hamlin, Sep. 1928 (UGa). dekalb: Stone
Mountain, Small, Sep. 1894
(ANS,NB). FANNIN : Blue
Ridge Mts., H. H. Smith

m28 (W). HABERSHAM:

Toccoa Falls, Small, ann.
1894 (NB). RABUN:Tallu-
lah Falls, Small, Sep. 1894
<Mo, NB); RICHMOND:
Augusta, Cuthbert, Sep.
1899 (NB). union: Young-
cane, Pennell 14053 (ANS).
South Carolina: Green-
ville: Saluda Falls, J. D.
Smith, Aug. 1881 (ANS, G,
NYS). LEXINGTON: Bates-
burg, McGregor 227 (US).

Fig. 7. Range of Lobelia amoena.

North Carolina: buncombe: Biltmore, Biltmore herb. 622b (ANS,
G, M, Mo, NB, W). HENDERSON: Muddy Creek, J. D. Smith, Aug!
1881 (G). MACON : Highlands, Biltmore herb. 622c (ANS, G, Mo, NB,
UP), polk: Tryon, Wherry, Sep. 1934 (UP). Transylvania: Pink
Beds, Biltmore herb. 622a (G, NB).

4. L. ELONGATA Small, Fl. S.E.U.S. 1144. 1903. Type Locality:
Northwest, Norfolk Co., Va. Type Specimen: Heller 1246, in the
herbarium of the New York Botanical Garden. Stem upright,
unbranched, slender or rather heavy at the base, 30-150 cm. high
(120 cm. according to Small), green or with a purplish tinge, darker
near the base; smooth. Leaves cauline, few-20, more or less upright-
appressed, narrowly lanceolate, sharply dentate or sub-entire, long-
acute at both ends, with prominent veins, smooth or strigose; some-
times papillose beneath; average size about 1.3 X 8.5 cm. (0.5-2.5 X
5-12 cm.). Upper leaves gradually smaller, but definitely larger than
the bracts of the inflorescence. Inflorescence a terminal raceme
10-30 cm. long (ave. 15-20 cm.), strongly secund, rather densely
few-50 (ave. about 20) -flowered. Pedicels rough, 4-7 mm. long in
fruit, each with a pair of bracteoles near the base. Flower-bracts
smooth, dentate, lanceolate or linear, 1-2 cm. long, inconspicuous or
the lowest leafy. Calyx in anthesis short-hemispheric, smooth or
with a few hairs, becoming sub-globose in fruit, 6-8 mm. across,
broadest at the middle. Capsule mostly inferior. Calyx-lobes smooth,
linear-subulate or with a short-deltoid base, 6-13 mm. long (ave. about
9 mm.). Auricles none or very small. Flower 20-25 mm. long,
including calyx (ave. 22.5 mm.). Corolla deep blue (ace. to Small),
smooth. Corolla-tube fenestrate; lobes of the lower lip broad-ovate or

i

>

286

Rhodora

[August

4 \

1

i

■/

f

w H^i.. 'l|

''^^*

\

h

ML ^i:,.'^^-;'^ y'^^i.—

,\^

l" "i 'f^-^-'^^''^^^^

\

!x

\XW\

\

1 '

7 r

^A

f'^l^^'^^K V

tiVS ^>sl

Fig. 8. Range of Lobelia elongata.

oblong, shorter than the tube; two upper lobes lance-Hnear. Filament-
tube 8-11.5 mm. long (ave. 9 mm.), pubescent below, connate about
half its length above. Anther-tube 4 mm. long, bluish-gray, the two

smaller anthers tufted, the
three larger pubescent on the
backs or nearly smooth. —
Swamps, low grounds, tidal
marshes; near the coast,
Georgia to southern Delaware
(south to Florida and west to
Louisiana, according to Small).
Flower August-October. Rep-
resentative Material seen:
Georgia: liberty: nr. Sun-
bury, L. LeConte (NB). South
Carolina: Berkeley: San tee
Canal, Ravenel (G). Dorches-
ter: Summerville, Brownfield,
Oct. 1892 (Mo). North Caro-
lina: COLUMBUS: Whiteville,
Schallert 1647 (Duke), new
HANOVER: Wilmington,
Williamson, ann. 1900 (ANS,
NB). Virginia: Norfolk: Northwest, Heller 1246 (ANS, G, M, Mo,
NB, UP). Maryland: somerset: Princess Anne, Canfci/ (ANS, Del,
G, Mo, NB, O, UP). Delaware: Sussex: Millsboro, Commons, Sep.
1877 (ANS, NB).

5. L.glandulifera (Gray) Small, Fl.S.E.U.S. 1144. 1903. Type
Locality: "S. Virginia to Florida and Alabama." Type Specimen:
Small gives as a synonym L. amoena var. glandulifera Gray (Syn. Fl.
4. 1878). Material identified by Gray as this variety, now at the
Gray Herbarium and at New York Botanical Garden, is a mixture of
two things; the first is the plant called L. glandulifera by Small and
later authors, and the second seems to be a hybrid of L. brevifolia
Nutt. The original description of var. glandulifera Gray is so worded
that it fits either the former, which is smooth and lacks auricles of the
calyx-lobes, or the latter, which is hirsute-pubescent and has the
calyx decidedly auriculate. In view of this confusion, I am typifying
L. glandulifera by the element of Gray's material which Small and
later authors have treated as a species. — L. amoena var. obtusata Gray,
Syn. Fl. 4. 1878. — Stem upright, slender, rather weak, unbranched,
30-125 cm. high, green or dark purplish-red near the base, smooth or
rarely short-hirsute. Leaves cauline, widely spaced, 6-20, spreading,
smooth (rarely with a few hairs beneath), thick, with a parchment-
like texture and a characteristic bluish-green or gray-green sheen in
dried material. Leaves short-ovate or elliptic, broadest at or below
the middle, mostly short-acute at both ends, with small sharp regular

la

287

1936] McVaugh,— North American Species of Lobel

I'om^ wtlt^^^ 7T2 ^"^ sub-petiolate averaging about 2 X 5.5 cm.
fhT«^i^ fl K P ''^•^- ,^PP^^ ^^^^^3 distinctly larger than

the small flower-bracts, givmg the inflorescence a naked appearance.
Inflorescence a loose termmal raceme, bearing 1-20 (30) rather widely
fn w\ ^T^''?, "P«" stout erect smooth pedicels (2.5-4 mm. long
smooth^'n? ^^o P^''.^^ br^cteoles near the base. Flower-bracts

i^Z^A yTj ^-2 c'"-.*^"^' inconspicuous, prominently glandular-
toothed or lobed. Calyx m anthesis conic or short-hemispheric, smooth
or rare y with a few hairs, becoming hemispheric or sub-globose in
^Tfl^Tfnf'^^' mature fruit seen only a few times: somewhat
at hP ton r ""?• uT'' )y^^-^ ^^' high, usually broadest
nL^ ^' 1 y^'''^^ ha If mferior or more. Calyx-lobes smooth,
narrow^ nearly hnear or with a broad base, 5-8 mm. long, acuminate
F?ot.^20?9r"'''' g^^'^^ular teeth. Auricles none or very small.
cZZ K?" ^™i^"^' including calyx, averaging about 22.5 mm.
Corolla blue, smooth. Corolla-tube fenestrate; lobes of the lower Hd
ovate or oblong, broadly obtuse or short acute, often as long as the
tube or longer; two upper lobes oblong. Filament-tube 6.5-8.5 mm.
long (aye. about 7.5 mm.), pubescent below, connate less than half
Its length above. Anther-tube 3.0-3.5 mm. long, light bluish-gray,

the bickr ' ^" ^ ' *^^ *^'^^ ^^'^^' "^^'^^y pubescent on

The following field-notes by A. H. Curtiss (accompanying 6938 in
Gray herb.) may be of value: ^ J' e

or,!i^- ^^u^^^^^ ®P'. ^\*^ ^^"^® ^- ^^^ very thick leaves, growing in low
sprlwhng abou wlt^^." f^"^ '"^ ^'^ ^°°«' ^^e lower H bare,Yhe lon^r oneT

Low grounds, meadows, swamps, and moist woods, eastern Tenn-
essee and western North Carolina to northern Florida, north to
southern Virginia. Mountains, Piedmont, and Coastal Plain. Flower
July-November Representative material seen: Tennessee: knox:
Knoxville /Ji/M, Sep. 1895 (Mo), possibly an error. Florida:

^Q?rm 'i^r "a?' a^^xto'^ (^^^^- ^^^^^^ P«"^« d« Leon, Curtiss
6938 (Del, G, M, Mo, NB). jackson: Marianna, Curtiss 1639 (G

m' 11 ;t "^""ifJi?; liberty: Aspalaga, Chapman, Biltmore herb. 6168
(^, M, Mo, NB). Georgia: bibb: Macon, G. N. Green f (ANS)
RANDOLPH: Cuthbert, Harper 1758 (G, Mo, NB). North Carolina!
buncombe: Biltmore, Ashe, Biltmore herb. 622b (Mo), catawba-

r^^tV^' !\^''^' ^^' ^^^^ (U^)- J^urham: Durham, Martin, Oct.'
1916 (Duke). FORSYTH: Winston-Salem, Schallert, Sep. 1921 (Duke
G). LINCOLN: Lincolnton, Curtis (Torrey herb., NB). orange!
Chapel Hi 1, Co^.r, Sep. 1909 (NC). Pasquotank: Elizabeth City,
^o/rfm^e /(?5^(NB; a fragment only), wake: Raleigh, Ashe, Curtiss
6453 (Del, M, Mo, NB). Virginia: hanover: Noel, Bnnton, Oct

I

**!!!r-

i

1'

»

\

\

Fig. 9. Range of Lobelia glandulifera.

A

288 Rhodora [August^

1890 (NB, UP).
JAMES city: Ewell,
Grimes U72 (NB).
In the Academy of
Natural Sciences of
Philadelphia is a
specimen collected
by Pursh in 1806 in
Greensville or South-
ampton County, Va.
This may be a du-
plicate of one in New
York in the Torrey
herbarium, labelled
"Greenville" and
" herb. Barton."

6. L. GLANDULOSA

Walter, Flora Caro-
lin.218. 1788. Type
Locality: "Carolina Meridialis, ad Ripas Fluvii Santee." Type
Specimen: There are in the Gray Herbarium a few fragments of what
is now passing as L. glandulosa, inscribed by Asa Gray Herb Walter I
Gray considered this plant to be the L. glandulosa of Walter.^ ihe
description from the "Flora Caroliniana" is given here: caule
erecto subpiloso, foliis oblongis obtuse sublanceolatis subdentatis
longitudine florum, flor. axillaribus solitariis purpureis pedunculis
brevibus, bracteis 2 glandula terminatis, capsuhs villosis, calyns
laciniis dentatis longis suberectis."— L. crassiuscvla Uicha.ux, M.
Bor. Am. H: 152. 1803. Although there is no material of this species
in the Michaux herbarium, his description leaves little doubt; he
himself, however, indicates doubt that L. glandulosa Walt is a
synonym. L. glandulosa A. DeCandoUe, Prodr. Syst. Nat. VII: 378.
1839 (in part). DeCandoUe was confused by what seems to be hybrid
material of L. bremfolia Nutt. (Torrey herbarium. New York Botan-
ical Garden).— Stem slender, unbranched, weak, erect or ascending,
30-140 cm. long (often tall, 90-100 cm.), smooth, green, or darker
below. Internodes sometimes zigzag. Leaves cauline, few-20, smooth,
thick, narrowly linear to broad-lanceolate, 0.2-1.4 X 3-15 cm. (ave.
about 0.6 X 8 cm.), on the average about 15 times as long as wide,
decurrent, not much narrowed at the base except the lowest; some-
what appressed to the stem, strongly callose-denticulate or sub-entire
in outline. Upper leaves merging into the floral bracts, but the larger
leaves well below the inflorescence. Inflorescence a lax terminal
raceme, usually strongly secund, bearing 1-20 (ave. 8-10) rather
widely separated flowers upon stout, rough-puberulent or hirsute
straight upright pedicels (5-13 mm. long in fruit), each with a pair
of bracteoles near the base. Flower-bracts smooth, linear, rarely

.<j

19361 MlVaugh.-North American Species of Lobelia 289

Cat iraSs trf-ltftheTcltei'ot^^^^^
ong chaffy-hixsute becomingTmi:^^^^^^^^

^i>-S6 mm. long, including calyx (ave 24-2^ mm \r ii iT

cr^r pLr K^- lutr a^^d S'rFi ^''-^'^ '- 'f

^'v^.l^ZrKrr •^"""'' SnuJa:rc:;teHf8 Sn's NB)'
DUVAL: Jacksonville, C«rtw 53^5 (ANS Del M NTR WvS\ iJ ''
UN: Apalachicola, Chaj^an (BilLore S. ae^SbWG L NbT
GADSDEN: Quincy, Chapman (ANS) Hamilton WMt.Q ■ ''

CM G M M^ NB \ipT*''^^= Bradentown, Tracy. Oct.' 1900
Duke m7'nR TTpf' ^- ""S^^f K"^^ ^''^^' Moldenke 357
NR itpT ' ' ^^^4.. ''«*nge: Bithlo, Moldenke m (Duke Mo
n^N ^P/^T^'"'^'- K'ssimmee, Meams 34 (US), pasco- St'^'

SI ^r Is/rrNR^' ^"'- '^'^ ^^^\ "^- •'°=~«= St. Augustbi
vSomif ' n«?' aV^^^- ??""'«"'E: Sanford, Moldenke 6348 (NB)
V0LU8U. Deland, LaForce, Nov. 1920 (NYS). Georgia: Baldwin-

I

I.';.

.1

'♦

I

290

Rhodora

[August

PS

3

J

4

Milledgeville, Boykin (ANS, NB). colquitt: Moultrie, Harper 1663
(G, Mo, NB). GLYNN : Brunswick, Pennell 4822 (NB, UP), liberty:
nr. Sunbury, L. LeConte (Torrey herb., NB). pike: Zebulon, Harper
2242 (G, Mo, NB). RICHMOND: Augusta, J. D. Smith, Sep. 1883 (G).
tatnall: Reidsville, Leeds, Oct. 1933 (ANS). ware: Manor, Mrs.
Lovett, Oct. 1933 (Duke). South Carolina: aiken: Granite ville,
Eggert, Aug. 1898 (Mo), beaufort: Hardeeville, Leeds, Oct. 1933

(ANS). BERKELEY:

San tee C anal,
Ravenel, Sep. (G).

CHARLESTON :

Charleston, Mol-
denke 143a (NB).
COLLETON: Walter-
boro, Leeds, Oct.

1933 (ANS). DAR-
LINGTON: Hartsville,
Norton, Nov. 1921

(NC). DORCHESTER:

Summerville,
Bro'umfield,Oct. 1S92

(M,NYS). JASPER-

Ridgeland, Mohr,
Nov. 1895 (Mo).
RICHLAND: Colum-
bia, J. D. Smith, Sep.
1883 (Mo). North
Carolina: Bruns-
wick: Wilmington,
west of river, Bariram, Oct. 1908 (ANS). columbus: Schallert,
Nov. 1926 (Duke). Johnston: State Forest Nursery, Blomquist 6716
(Duke). NEW HANOVER: Wilmington, McCarthy, Sep. 1888 (NC).
PENDER: Holmes, Sep. 1884 (NC). Virginia: Mr. Bailey, ann. 1841
(NB); Gray cites the species from "s. Virginia" in his Synoptical
Flora, on Bailey's authority. Kearney {33) (1901) cites his own 2378,
collected "in open fresh-water marshes of the Northwest River'*
(Norfolk Co., Va.).

7. L. BREViFOLiA Nuttall, A.DC. Prodr. Syst. Veg. VII: 377.
1839. Type Locality: "in Alabama Americae bor.". Type Speci-
men : material from Nuttall's herbarium is in the Academy of Natural
Sciences of Philadelphia. — L. Ludoviciana Wood, Class Book 476.
1861. — Stem slender, unbranched, rather weak, 30-90 cm. high,
smooth or nearly so, green or tinged with purplish-red near the base.
Cauline leaves 20-100 (in cases of fasciation 200); close together;
linear, oblong, or the lower oblanceolate, short-acute or obtuse,
0.2-0.8 X 0.7-3.0 cm. (ave. 0.5 X 1.5 cm.), strongly pectinately
toothed, the teeth callose- tipped. Leaves smooth or somewhat

FiQ. 10. Range of Lobelia glandulosa.

i-'i

Ik

1936] McVaugh,— North American Species of Lobelia 291

strigose, decurrent, not much narrowed at the base, except the lowest;
the upper more distant and merging gradually into the bracts of the
inflorescence. Inflorescence a loose terminal raceme, strongly secund,
bearing few-30 (ave. about 15) flowers upon stout, upright, rough or
hirsute pedicels (5-10 mm. long in fruit), each with a pair of bracteoles
near the base. Flowers often standing stiffly at right-angles to the
stem. Flower-bracts smooth or somewhat ciliate beneath, 0.5-1.5 cm.
long, inconspicuous, strongly toothed. Calyx in anthesis short-
hemispheric, smoothish or densely long-hirsute, becoming sub-globose
or hemispheric in fruit. Calyx-lobes broad at the base, 4.5-8.0 mm.
long (ave. about 6.0 mm.), long-acute, strongly pectinately toothed,
sometimes fimbriate; auricles at the base of each lobe broad, round,
foliose, usually as long as the calyx-tube and covering it; sometimes
toothed. Flower 15-24 mm. long, including calyx (ave. 18-20 mm.).
Corolla pale blue (ace. to
Chapman, 1897); azure, ace.
to Mohr, 1901; (P, 45); pu-
bescent outside, the lower lip
smooth or puberulent inside.
Corolla- tube fenestrate or
sometimes entire except for
the dorsal fissure; lobes of the
lower lip narrow-ovate, short-
acute, shorter than the tube;
two upper lobes lanceolate.
Filament-fube (5) 6.5-7.0
(8.0) mm. long, pubescent,
connate above. Anther-tube
3.5-4.0 mm. long, bluish-gray,
all the anthers white-tufted

Fig. 11. Range of Lobelia brevifolia.

at the tip, or the three larger merely pubescent on the backs. — Damp
pinelands, usually in sandy soil, western Florida to eastern Louis-
iana. Coastal Plain. Flower late Summer and Fall. Represen-
tative Material seen: Florida: franklin: Apalachicola, Saurman,
ann. 1867 (ANS); Chapman, Biltmore herb. 4166a (NB). holmes:
Westville, Curtiss 6906 (Del, M, NB). liberty: Aspalaga, Chapman,
Biltmore herb. 4166b (NB). Alabama: lee: Auburn, Earle, Oct. 1896
(CM), mobile: Mobile, Mohr (ANS). Mississippi: Harrison: Pass
Christian, Langlois, Oct. 1882 (CM, NYS, UP). Louisiana: Or-
leans: New Orleans, Ingalls (Torrey herb., NB). In the Schweinitz
herbarium at the Academy of Natural Sciences of Philadelphia is a
specimen marked "Louisiana, Tainturier."

Occasionally, plants are met with that show resemblances to the
above species, and may be of hybrid origin; possible parents are
L. brevifolia and L. pvbenda:

L. brevifolia X L. puberula.— Z. glandulosa A. DeCandolle,

V'

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1

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If

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i

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Y

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I

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292

Rhodora

[August

Prodr. Syst. Veg. VII: 378. 1839 (in part). L. amoena var. gland-
uhfera Gray, Syn. FI. 4. 1878 (in part).-Leaves broad, more or less
pubescent, inclined to be dentate. Calyx-lobes more or less strondy
glandular-toothed; auricles round, rather small, but conspicuous
Calyx smooth or hirsute. Filaments sometimes reaching 9.0 mm in
length. Material seen: Florida: "west Florida," Chapman (NB)
Alabama: Gates (G, NB), both identified as L. amoena var. glandi
uhfera by Asa Gray; the one at New York named L. glandulosa by
ZiQOA ^x^N ^i^E- Auburn, Earle, Sep. 28, 1896 (CM); Earle, Oct.
\}^^^ iP^' ^""'^'^ ^^*- ^^' 18^^ (CM, NB). mobile: Graves 1119
(Mo). Mississippi: Kashtaw, Tra^ mo (NB). Harrison: Biloxi,

S89^(CM)' ^' ^'''''^^''^'' ^^^^^ SP^'^gS' ^^'•^^^ Oct. 27^

8. L. PUBERULA Michaux. Fl. Bor. Am. II: 152. 1803. Type
Locality: Carolina." Type Specimen: There is no material of this
species in the Michaux herbarium in Paris. The original description,
which follows, seems to fit no other species than the one to which it is
now given:

rZh^^^^^'x s^°^P^i9issiina, pubescens: foliis oblongo-ovalibus, obtusis,
Sbt^^atcibi^ P^^-^^^^^ ^^^^ P--' ^^ternis, sub!

The nomenclature of this species and its forms has been in con-
siderable confusion. DeCandolle, in the "Prodromus," seems to
confuse several plants. In the absence of type material, and in view
of the variability of the forms, it seems impossible at present to deter-
mine the exact identity of L. jmberula of Michaux.

L. vvberula var. glabella Elliott, Sk. Bot. S. C. & Ga. I: 267. 1821
IS probably L. elongata Small.

L. puberula var. glahella Hooker, Bot. Mag. LXI: plate 3292 1834
IS probably the smoothish form found on the Gulf Coastal Plain

btem strict, usually unbranched, 30-160 cm. high, often 100 cm
green or sometimes with a purplish tinge throughout, darker below!
densely short-hirsute throughout, or sometimes glabrate. Cauline
leaves few-40, hairy beneath and more or less strigose above, espe-
cially near the margins; thin but fairly firm in texture; shape varying
from lanceolate-acute sub-entire in outline, with prominent callose-
denticulate teeth, to broadly obovate, obtuse, with coarse irregular
serrations and inconspicuous callose teeth. Upper leaves merging
gradually into the bracts of the inflorescence, becoming more finely
toothed above. Basal leaves none. Inflorescence a terminal un-
branched raceme 4-50 cm. long (ave. 15-30 cm.), densely flowered or
often somewhat interrupted; often distinctly secund, bearing few-70
flowers upon short stout puberulent or hirsute pedicels (3-5 mm. long
in fruit) each with a pair of bracteoles at the base or somewhat above
It. Pedicels not stiffly erect. Flower-bracts various. Calyx in
anthesis flattish or short-hemispheric, more or less pubescent, or
hirsute, becoming hemispheric in fruit, widest at the top, usually with

^

v

- m

1936]

McVaugh, — North American Species of Lobelia

293

a flaring rim, prominent ribs and a rough angular appearance; 5-9
mm. across by 4-7 mm. high. Capsule about half inferior. Calyx-
lobes lanceolate or broader, plainly broader near the base than near
the tip, more or less straight-sided, without definite subulate tips,
5-12 mm. long, usually ciliate at least near the tip. Auricles very
small and triangular, or rounded, short, formed of the rolled edges and
lobes of the cordate calyx-lobes. Flower 15-24 mm. long, including
calyx (ave. 18-20 mm.). Corolla blue, with a white eye, ciliate at
least on the veins outside, the lip smooth. Corolla- tube fenestrate;
lobes of the lower lip oblong or ovate, usually somewhat shorter than
the tube, acute or obtuse ; two upper lobes lanceolate, erect. Filament-
tube 6-7 mm. long (rarely 9 mm.), pubescent below, connate about
a third of its length above. Anther-tube 3.0-3.5 mm. long, light
bluish-gray, the two smaller anthers tufted at the tip, the three larger
usually pubescent on the backs. — Wet woods, low grounds, thickets,
in various soils; Coastal Plain and upland provinces, Tennessee and
western North Carolina north to West Virginia, southern Indiana and
Illinois; northeast along the Coastal Plain from South Carolina to
southern New Jersey; south to Florida, Alabama and Mississippi;
west and south to Missouri, Arkansas, Oklahoma, Texas and Louisi-
ana. Flower August 1 (rarely earlier) through the Fall.

A wide-ranging species showing several pronounced geographic
forms which may or may not be worthy of varietal names; the most
conspicuous are as follows:

a) A form with what may be called an Alleghanian range; rarely
found on the Coastal Plain or in glaciated country; native from West
Virginia south to mid-Georgia, west to Illinois and western Tennessee.
In its most characteristic form it may be identified by the rather
sparsely pubescent calyx-tube and lobes, small (1-1.5 cm. long)
bracts, which are lanceolate or nearly linear, the narrow calyx-lobes
(1.5-2.0 mm. wide by 5-12 mm. long), which are usually flat or slightly
rolled on the margins to form small auricles; the leaves are often
spreading rather than appressed, usually prominently callose-denticu-
late, even the lowest sometimes acute, tending to be three times as
long as wide, or longer. This plant may be the closest, now living, to a
hypothetical ancestor living in approximately the same area. Several
lines of divergence from this type may be seen, leading to at least
three others. Representative Material seen: Virginia: Bedford:
Curiiss, Sep. 1873 (Mo, NYS). Washington: Damascus, Core 3883
(NB, WVa). wythe: Wytheville, Shriver, Sep. 1878 (ANS, G).
West Virginia: marion: Winfield Twp., Sharp, Sep. 1929 (WVa).
MONONGALIA: Rumsey, Sep. 1897 (WVa). monroe: Peters Mt.,
Steele 172 (G, Mo, NB). ritchie: Auburn, Randolph 1393 (G).
UPSHUR: Buckhannon, Pollock, Sep. 1895 (Mo), wirt: Elizabeth,
Bartholomew 304 (WVa). North Carolina: ashe: Ashe, Aug. 1891
(NC). buncombe: Asheville, Redfield 5636 and 6636 (Mo), burke:
Morgan ton, Moses, ann. 1914 (NC). Durham: Duke Forest, Oosting

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[August

555^^ (Duke), forsyth: Winston-Salem, Schallert, Aug. 1925 (Duke).
HAYWOOD: Lake Junaluska, Blomquist 5019 (ANS, Duke). Hender-
son: " Chimney Rock to Henderson ville," Small and Huger, Oct. 1901
(NB). JOHNSTON: Parkers Pond, B. E. Smith, Sep. 1932 (NC)
orange: Chapel Hill, Coker, Sep. 1914 (NC). Perquimans: Glasson
(Duke): person: Roxboro, Wherry, Sep. 1934 (UP), swain: Great
Smoky Mts., Beardslee and Kofoid, ann. 1891 (G, M, Mo, NB, WVa).
South Carolina: Anderson: Anderson, Dams 8424 (Mo), green-
wood: Greenwood, Bartram 3313 (ANS). Lexington: Batesburg,
McGregor 66 (US), oconee: Clemson College, HoTise 2892 (Mo).
Georgia : clarke : Athens, Wiegand and Manning 3089 (G) . dekalb *:
Stone Mountain, Small, Sep. 1894 (Mo, NB). floyd: Rome, Canby,

Oct. 1898 (Del, Mo).
jasper: Monticello,
Porter, ann. 1846

(ANS). LAMAR:

Barnesville, Hamlin,
Aug. 1928 (UGa).
MUSCOGEE: Colum-
bus, Boy kin (NB).
RABUN : Tallulah
Falls, Small, Sep.

1894 (Mo). RAN-
DOLPH: Cuthbert,
Harper 1734 {G,Mo,

NB). RICHMOND:

Augusta, Cuthbert,
Sep. 1898 (NB).
Alabama: Chero-
kee: Center, Leeds,
Oct. 1934 (ANS).
7? ^ o ,0^-, /^,, , CULLMAN: Cullman,

^^''h^o^?\lf^J ^^^' ^""^ ^^)- JEFFERSON: Birmingham, Vasey,
arm 1878 (ANS). lee: Auburn, Earle and Baker, Sep. 1897 (M, Mo
NB) mobile: Graves 1194 (Mo), tuscaloosa: Tuscaloosa, Johnso^
iiv,^^.' ^'Vnherula var. glabella, fid. A. Gray. Mississippi: Harrison-

? I'oJ/^St f ^ ^^^^' ^^)- Jackson: Ocean Springs, Skehan,
bep. 1895 (Mo). Lauderdale: Meridian, Schuchert, Oct. 1896 (NB)
I'Lorida: DUVAL : Jacksonville, Curtiss 6565 (Del, G, M Mo NYS)
manatee: Terra Ceia Island, Simpscm 411 (G, NB). ' Louisiana-
bossier: Alden Bridge, Trelease,OQt. 1898 (Mo). Kentucky: bell-
Pine Mountam, Kearney, Sep. 1893 (G, M, Mo, NB). carter: Olive
Hill Svaison 4409 (ANS, CCD, G). edmonson: Mammoth Cave,
Leeds, Oct. 1934 (ANS). fayette: Lexington, C. IV. Short (W)
^^wi^- y^^t^^^S' ^^'^^^^ ^36^7 (ANS). laurel: London, McFarland
^I (Mo). LINCOLN: Kings Mountain, Pennell 13746 (ANS) lyon-
Kuttawa, Epgleston 5195 (NB). mc creary: Parkers Uke, Pennell

Fig. 12. Range of Lobelia puberula, Form a.

1936] McVaugh,— North American Species of Lobelia

295

13801 (ANS). vvLASKiiFloyd, Pennell 13758 (AISIS). warren: Bowl-
ing Green, Price, Jy. 1894 (Mo). Tennessee: blount: Chilhowee Mt.,
Curtiss 1636 (ANS, CM, G, M, Mo, NB). carroll: Hollow Rock Jet.,
Svenson 453 (ANS, G). carter: Roan Mt. Sta., Rydberg 8180 (NB).
COCKE: "Paint Rock to Del Rio,'' Kearney 806 (M, Mo, NB). dick-
son: White Bluffs, Eggert, Aug. 1897 (Mo), franklin: Sewanee,
Eggert, Sep. 1898 (Mo). Hamilton: Chattanooga, Lippincott, Sep.
1895 (ANS). KNOX: Knoxville, Ruth, Sep. 1895 (M, Mo, W). madi-
son: Jackson, Bain 313 (NB). polk: Reliance, Pennell I4OO4 (ANS).
PUTNAM: Cookeville, Hudson 105 (R). sevier: "Great Smoky Mts.,"
Schallert, Sep. 1933 (NB). shelby: Memphis, Fendler, Sep. 1853
(G). TIPTON: Covington, Rhoades, Jy. 1927 (W). wayne: Waynes-
boro, Svenson 4304 (ANS, G). Ohio: hocking: Queer Creek, Griggs,
Aug. 1910 (G). Indiana: clark: State Reserve, Beam 5440 (CCD).
CRAWFORD: Leavenworth, Beam 18566 (CCD). dearborn: Man-
chester, Beam 30126 (CCD). Harrison: New Middletown, Beam
18725 (CCD). JEFFERSON: Kent, Beam 35293 (ANS). perry:
Cannelton, Beam 33211 (CCD). Vanderburgh: Evansville, Beam
33115 (CCD). Illinois: henry: Galva, "^. Br (Horner herb.) (G).
JACKSON: Makanda, Vasey, Aug. 1862 (G). pulaski: Karnak,
Palmer 16554 (ANS, Mo), union: Cobden, Earle, Sep. 1878 (CM).

b) Common on the southeastern Coastal Plain and adjacent
Piedmont, New Jersey and Pennsylvania south to Georgia, is a plant
distinguished by a densely long-hirsute calyx, with broad calyx-lobes
(2.5-5.0 mm. wide by 5-12 mm. long), which are often undulate or
crisped on the margins, with rounded auricles formed by the reflexed
margins and the lobes of the cordate base. Bracts usually leafy,
broad at the base, 1-2 cm. long; leaves more or less closely appressed
to the stem, acute above, but usually obtuse and obovate below,
often entire in outline or irregularly coarsely serrate, but not con-
spicuously callose-denticulate. Representative Material seen:
Georgia: Chatham: Savannah, Nuttall, herb. J. Gay (G). liberty:
nr. Sunbury, L. LeConte (NB). walton: Logan ville. Small, Sep. 1894
(NB). South Carolina: aiken: Aiken, Ravenel, Sep. 1866 (Mo).
CHARLESTON: Charleston, Backman (ANS). North Carolina:
HALIFAX: Weldon, Bartram, Oct. 1908 (ANS, NB). Pasquotank:
Elizabeth City, Moldenke 108 (Duke, Mo, UP), rowan: Salisbury,
Heller, Aug. 1890 (ANS, M, Mo, NB). Virginia: accomac: Franklin
City, Brown, Sep. 1907 (ANS). Arlington: Fort Myer, Mearns 134
(NB). FAIRFAX: Great Falls, Wismer 433 (UP), james city: Wil-
liamsburg, Grimes 4434 (M); 4388 (G, NB). Norfolk: Northwest,
Heller 744 (Mo), prince george: New Bohemia, Pennell 14426
(ANS). PRINCESS anne: Virginia Beach, Heller 1335 (ANS, NB).
District of Columbia: M. S. Bebb, ann. ca. 1863 (ANS, G); Takoma
Park, House 1532 (Mo); 323 (NB). Maryland: anne arundel:
Leon, Shull 259 (Mo, NB). Baltimore: Catons ville. Foreman, ann.
1873 (NB). CECIL: North East, J. P. Otis f P479 (herb. R.R.T.).

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KENT: Golts, Brown, Sep. 1907 (ANS). prince georges: Ammendale,
Bro. Hyacinth 1177 (Mo), queen annes: Centerville, Norton, Jy.
1903 (Mo). WORCESTER: Snow Hill, Moldenke 6618 (NB). Dela-
ware: KENT: Little Creek, Larsen 311 (UP). Newcastle: Newark,
Commons, Sep. 1869 (ANS). Sussex: Rehoboth, Churchill, Sep. 1908
(G). Pennsylvania: Chester: Nottingham, Bartram 1771 (ANS).

Fig. 13. Combined Ranges of Forms of Lobelia puberula, exceptine

Form a. ^ *

DELAWARE: Bethel-Concord line, Pennell, Oct. 1908 (ANS). Lan-
caster: Pleasant Grove, Small and Carter, Sep. 1908 (ANS NB)
YORK: Castle Fin, Crawford, Aug. 1895 (ANS). New Jersey*
ATLANTIC: Leeds Pointy (?ray, Sep. 1833 (G). Burlington: Hartford,'

mf^r<^n J-J^^^' ^^)- ^^^^ ^^^- ^^P^ May, Mackenzie UU
(LCD G Mo) CUMBERLAND: Bayside, Fogg 7445 (ANS). ocean:
Manahawkm, Lmg, Sep. 1909 (ANS).

c) In the southern states, Alabama to Louisiana, especially on the
Coastal Plam, form (a) above becomes practically smooth. This has
been called var. glabella Hooker, by American authors. Representa-
tive Material seen: Alabama: mobile: Theodore, Pennell 4486 (UP)
Mississippi: clarke: Shubuta, Schuchert, Oct. 1896 (NB). scott-
Jorest, Cook, Aug. 1925 (US). Wilkinson: Phares, Sep. 1868 (Miss)*
Louisiana: Orleans: New Orleans, Torrey (G). Texas: victoria*:

Missions valley near the Guadeloupe River above Victoria," Schott,
Uct. 1851 (JNB); this plant is perhaps only a depauperate specimen of
some other form of this species.

d) From Missouri and Arkansas southward to eastern Oklahoma
and lexas, and eastward to Alabama and southern Mississippi

1936] McVaugh,— North American Species of Lobelia 297

occurs a plant resembling that of the Atlantic Coastal Plain, but with
large leafy lanceolate bracts (usually not broad at the base), the calyx
often smoothish or merely strigose. Auricles well-developed and
sepals broad. Leaves often hairy, more or less appressed to the stem or
loose, often with conspicuous sharp small teeth, each callose-tipped.—
L. puberula var. mineolana Wimmer in Fedde Rep. Spec. Nov. XXVI:
4. 1929.— Representative Material seen : Alabama : Dallas : Trelease,
ann. 1879 (Mo). Washington: Fruitdale, A.GJ., Oct. 1903 (Mo).
Mississippi: smith: Taylorsville, Tracy, Aug. 1903 (NB, US). Louisi-
ana: CADDO : Shreveport, Gregg, Sep. 1847 (Mo). Calcasieu : Lake
Charles, Mackenzie 628 (Mo, NB). Natchitoches: Natchitoches,
Palmer 8714 (Mo, NB). rapides: Alexandria, Hale (ANS). Mis-
souri: DUNKLIN: Bush, Sep. 1893 (Del, G, Mo). Arkansas: hemp-
stead: Columbus, Palmer 6838 (Mo), hot spring: Malvern, Palmer
8462 (Mo, NB). HOWARD: Baker Springs, Kellogg, Oct. 1899 (Mo).
miller: Texarkana, Palmer 14634 (Mo), polk: Rich Mountain,
Trelease, Oct. 1898 (Mo), pulaski: Little Rock, Demaree 8163
(CCD, G, Mo, NB); Little Rock, Hasse, ann. 1886 (M, NB, NYS);
all material seen from Pulaski Co., seems to have rather large flowers.
saline: Benton, Greenman 4302 (Mo). Oklahoma: creek: Sapulpa,
Pennell 6395 (NB, UP), haskell: Sans Bois Mts., Sheldcm 310
(Del, NYS). LATIMER: Wilburton, Stratum 603 (Mo), mc curtain:
Broken Bow, Stratton 674 (Mo); this plant has oversize flowers.
PITTSBURG: McAlester, Sheldon 310 (Mo). Texas: Drummond (NB);
Lindheimer, Sinn. 1843 (G). anderson: Palmer 10721 (Mo). Austin
"S. Felipe de Austin," Drummond (G); this is the L. amoena of A.DC,
according to Asa Gray, bowie: Texarkana, Heller 4166 (ANS, G,
Mo, NB). BRAZORIA: Columbia, Bush 1630 (Mo, NB). Cherokee:
Jacksonville, Palmer 8593 (Mo, NB). gregg: Long view, Eggert,
Aug. 1898 (Mo). HARRIS: Houston, Lindheimer, Sep. 1842 (Mo).
HARRISON: Marshall, Bush 782 (Mo). Montgomery: Willis, Warner
(Mo). PANOLA: Beckville, Reverchon 3204 (G, Mo, NB). rusk:
"auf Prarien an Waldrandern," Vinzent, tex. Pfl. 60 (Mo), smith:
Tyler, Reverchon 2086 (Mo), upshur: Big Sandy, Reverchcm 3206
(G, Mo, NB). wood: Mineola, Reverchon, Aug. (Mo); presumably
the type of var. mineolana.

In addition to the above, a few specimens have been seen of the
plant designated as

L. PUBERULA, var. PAUCiFLORA Bush, Ann. Rep. Mo. Bot. Garden
17: 122. 1906.— Type Locality: Swan, Smith Co., Texas. Type
Specimen: Reverchon 3206, Sep. 17, 1902; seen in Missouri Botanical
Garden. Stems slender, densely white-pubescent. Leaves thin,
oblong, hirsute. Calyx and pedicel densely long-hirsute. Flowers
3-15 in number, larger than in L. puberula. Filament-tube 8-11 mm.
long. This plant seems quite distinct, and may deserve specific rank,
but as so little material has been seen, it seems best to leave its status
unchanged. Material seen : Louisiana : Natchitoches : Natchitoches,

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[August

Palmer 8905 (NB). rapides: Alexandria, J. Hale (G, Mo NB)
lEXAs: smith: Swan, i2e»crcAo7i5^0(? (Mo). '

(T'o 6e continued)

1

1936]

McVaugh, — North American Species of Lobelia

305

STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES

OF LOBELIA

Rogers McVaugh

{Continited from page 298)

9. L. SPICATA Lamarck, Diet. Bot. Ill: 587. 1789. This lis a
species with at least five well-defined phases, which may be dis-
tinguished as follows:

a) Var. leptostachys (A. DeCandolle) Mackenzie & Bush, Fl.
Jackson County, Mo. 183. 1902.— Type Locality: "in Carolina
merid." Type Specimen: The plant described by DeCandolle as
L, leptostachys was seen by him in the herbarium of Asa Gray, col-
lected by Fraser. This has not been seen, but there is in the New York
Botanical Garden a
specimen collected at
Lincolnton, N. C, by
the Rev. M. A. Curtis,
which was verified by
Asa Gray, and is also
marked " Lob. lepto-
stachys A.DC. Geneve
1839." — L. leptostachys
A. DeCandolle, Prodr.
Syst. Veg. VII: 376.
1839. L. bracteata
Small, Fl. S.E.U.S.
1146. 1903.— Stem
strict, unbranched, 30-
120 cm. high, dark pur-
plish-red and densely
short-pubescent near
the base, becoming
smooth and light green
above; often pubescent
on the angles formed
below the decurrent leaf -bases. Cauline leaves 10-40, usually quite
close together and somewhat appressed to the stem, thus giving them
an imbricated appearance in dried material; firm or leathery in
texture; sessile, or the lowest narrowed to short petioles; the lower
and middle ones obtuse, long-oblong or oblanceolate, to 2.5 X 12 cm.,
appearing sub-entire, but beset with callose-glandular teeth along the
margins. Upper leaves gradually smaller, becoming definitely bract-
like (lance-acute), and merging into the bracts of the inflorescence.
Basal leaves usually few or none; if present, oblanceolate, obtuse.
All leaves strigose-pubescent above and below, especially the lower

Fio. 14. Range of Lobelia spicata, var.

LEPTOSTACHYS.

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[August

19361 McVaugh,— North American Species of Lobelia

305

Palmer 8905 (NB). rapides: Alexandria, J. Hale (G, Mo, NB).
Texas: smith: Swan, Reverchon 3206 (Mo).

{To he continued)

't

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STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES

OF LOBELIA

Rogers McVaugh

{Continued from page 298)

9. L. SPICATA Lamarck, Diet. Bot. Ill: 587. 1789. This lis a
species with at least five well-defined phases, which may be dis-
tinguished as follows:

a) Var. leptostachys (A. DeCandoUe) Mackenzie & Bush, Fl.
Jackson County, Mo. 183. 1902. — Type Locality: "in Carolina
merid." Type Specimen: The plant described by DeCandolle as
L. leptostachys was seen by him in the herbarium of Asa Gray, col-
lected by Fraser. This has not been seen, but there is in the New York
Botanical Garden a
specimen collected at
Lincolnton, N. C, by
the Rev. M. A. Curtis,
which was verified by
Asa Gray, and is also
marked " Loh. lepto-
stachys A.DC. Geneve
1839." — L. leptostachys
A. DeCandolle, Prodr.
Syst. Veg. VII: 376.
1839. L. bracteata
Small, Fl. S.E.U.S.
1146. 1903.— Stem
strict, unbranched, 30-
120 cm. high, dark pur-
plish-red and densely
short-pubescent near
the base, becoming
smooth and light green
above; often pubescent
on the angles formed
below the decurrent leaf-bases. Cauline leaves 10-40, usually quite
close together and somewhat appressed to the stem, thus giving them
an imbricated appearance in dried material; firm or leathery in
texture; sessile, or the lowest narrowed to short petioles; the lower
and middle ones obtuse, long-oblong or oblanceolate, to 2.5 X 12 cm.,
appearing sub-entire, but beset with callose-glandular teeth along the
margins. Upper leaves gradually smaller, becoming definitely bract-
like (lance-acute), and merging into the bracts of the inflorescence.
Basal leaves usually few or none; if present, oblanceolate, obtuse.
All leaves strigose-pubescent above and below, especially the lower

Fia. 14. Range of Lobelia bpicata, var.
leptostachys.

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[September

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leaves and near the margins. Plants, when dried, often with a
characteristic brownish-green color. Inflorescence a terminal virgate
spike 20-30 (50) cm. long, densely flowered, not noticeably secund,
bearing 20-200 (ave. 30-60) flowers upon very short (in fruit 2-4 mm.)
rough-pubenilent curved pedicels, each with a pair of inconspicuous
bracteoles near the base. Flower-bracts ciliate-pubescent or some-
times smooth, lanceolate, acute, sometimes linear-lanceolate, usually
conspicuous, 1-4 cm. long. Calyx in anthesis flattish or conic, smooth
or pubescent, becoming hemispheric in fruit, strongly ribbed, about
3.5 mm. in diameter. Capsule }4-% inferior. Calyx-lobes subulate
or Imear-lanceolate, (2) 3-6 (7) mm. long, bristly-ciliate especially
near the tips, or smooth; auricles at the base of each lobe filiform,
1-3 (5) mm. long, deflexed. Flower 9-12 mm. long, including calyx.
Corolla light blue, smooth or pubescent outside, the lower lip pubes-
cent at the base inside. Corolla-tube entire, except for the dorsal
fissure; lobes of the lower lip ovate, not sharply reflexed, slightly
shorter than the tube; two upper lobes lanceolate, curved upward.
Filament-tube 3.0-3.5 mm. long, pubescent below, connate above
about half its length. Anther-tube 1.8-2.0 mm. long, light bluish-
gray, the two smaller anthers each with a tuft of white hairs at the tip,
the three larger usually pubescent on the backs.

While this plant in its most characteristic state seems wholly dis-
tinct from var. originals and var. hirtella, it is separated from them
by no constant characters, and where the ranges of any of the three
overlap, puzzling intermediates are frequently encountered. "Typ-
ical var. leptostackys differs from var. originalis by the longer auricles
shorter pedicels (and consequently narrower spikes), somewhat longer!
denser inflorescences, longer bracts, leaves thicker, longer, more
numerous, more neariy entire, more appressed; somewhat heavier
pubescence. It differs by similar characters from the pubescent
var. hirtella. However, many specimens from Illinois, Missouri,
Iowa, can be referred to no one of the three; all the distinguishing
characters break down. In consequence, it seems best to consider all
three as varieties of the same species (See Fig. 19).

Dry sandy woods and hillsides, northern West Virginia to central
Alabama, west to northern Illinois, eastern Kansas and western
Arkansas. Rare or absent on the eastern Coastal Plain; occasional
on the Gulf Coastal Plain in Alabama, Mississippi, Arkansas and
western Tennessee. Flower June 20-August 1. Representative
Material seen: Virginia: Montgomery: Leidy, Jy. 1867 (ANS UP)
SMYTH: Chilhowie, Small, Aug. 1892 (NB). West Virginia: green-
IT^'n T^^^'^L fu"^ f^CCM). MONONGALIA: "Morgantown,
wV^?R• ^"^'^f^^'* ^^'^' (^)- ^^^^- Leachtown, Millspaugh
l^i^Jii^^^K^^^r^S^^^^^^^ buncombe: Biltmore, Biltmore

P >ff /£' J!^{ ^^' ^^' U^' US> W)- CALDWELL: Hudson,
Randolph llSl (G). cherokee: Andrews, Huger, Sep. 1900 (NB)
DURHAM: Durham, Blomquiat 6023 (Duke), forsyth: Winston-

1936] McVaugh,— North American Species of Lobelia 307

Salem, Schallert, Jy. 1911 (Duke), haywood: Ashe, Sep. 1893 (NC).
IREDELL: Statesville, Hyams (M). Lincoln: Lincolnton, Curtis
(ANS, NB). orange: Hillsborough, M. A. Curtis f (G). polk:
Columbus, Tovmsend, Oct. 1897 (US), rowan: Salisbury, Heller 108,
Jun. 28, 1890 (ANS, Mo, NB). swain: Great Smoky Mts., Heming-
way, Aug. 1891 (WVa). South Carolina: pickens: nr. Clemson
College, House 2957 (US). Georgia: Baldwin: "Georgia, Dr.
Boykin," Torrey herb. (NB). clarke: Princeton, Miller and Maguire
1489 (CU). CLAYTON: dry woods, Harper 230 (G, Mo, US), cobb:
Wilson S3 (G, Mo, US), dekalb: Stone Mt., Eggert, Jy. 1897 (Mo)
FANNIN: Blue Ridge Mts., H. H, Smith 2633 (W). floyd: W. Rome,
Pennell 4079 (UP, NB). gwinnett: McGuire's Mill, Small, Jy. 1893
(Mo, US). HABERSHAM: Clarkcsville, /. D. Smith, Sep. 1883 (G, US).
walker: Lookout Mt. (no Co. given), Ruth, Jy. 1898 (Mo, NB,
US). ALABAMA:AUTAUGA:Prattville, 3/oAr, Jy. 1880(US). blount:
Mohr, Jun. 1883 (US), clay: Millerville, Pollard and Maxmi 169
(NB). JACKSON: Scottsboro, Earle, Jun. 1899 (NB). Talladega:
Riddell's Mill, Mohr (US), tuscaloosa: Tuscaloosa, Mohr, Oct. 1879
(US). Mississippi: lowndes: Mayhew, D(mald, Jun. 1927 (W).
OKTIBBEHA: Starkvillc, Phares, ann. 1883 (Miss). Kentucky:
FAYETTE: Lcxington, C. W. Short, ann. 1836 (NB). pulaski: Bum-
side, Pennell 13796 (ANS). warren: Bowling Green, Price, Jun. 1897
(Mo). Tennessee: bradley: Cleveland, Leeds, Jun. 1929 (ANS).
CHESTER: Henderson, Bain 214 (NB). coffee: Tullahoma, Gattinger
Jy. 1880 (Del). Cumberland: Daysville, Anderson 14OI (G). hamil^
ton: Lookout Mt., Vasey, ann. 1878 (ANS, US), knox: Knoxville,
Ruth, Jy. 1895 (ANS, M. W). Ohio: clark: Springfield, Williams
(Mo). HAMILTON: Cincinnati, Lea (ANS, NB). Montgomery:
Dayton, Morgan, Jy. 1879 (NB, US). Indiana: benton: Barce,
Beam 11867 (CCD). brown: Belmont, Deam 43469A (CCD). cass:
Cicott, Deam 26899 (ANS). fountain: Fountain, Deam 26819
(CCD). HARRISON: Elisabeth, Deam 20629 (CCD). henry: Spring-
port, Deam 46344 (CCD). kosciusko: Winona Lake, Deam 422
(CCD). LAWRENCE: Mitchcl, Deam 17263 (CCD). Marshall:
Culver, Deam 9017 (CCD). newton: Thayer, Deam 60607 (CCD).
noble: Albion, Deam 20761 (NB). perry: Derby, Deam 11634
(CCD). tipton: Kempton to Goldsmith, Deam 13630 (CCD).
WASHINGTON : Campbcllsburg, Deam 37164 (CCD). white : Burnetts-
ville, Deam 39340 (ANS). Illinois: cass: Beardstown, Geyer, Jy.
1842 (ANS, G, Mo, NB, W). champaign: Raymond, Jy. 1869 (W).
christian: Taylorville, Andrews, Aug. 1898 (CU). Hancock:
Augusta, Mead, Jy. 1847 (NB). logan: Lincoln, Mills, Jy. 1899 (G).
MC lean: Hendrix, Robinstm, Aug. 1904 (G). mason: Havana,
Gleason, Aug. 1903 (G). menard: Athens, Hall, ann. 1861 (G, US).
PEORIA: Peoria, Brendel (NYS). ST. clair: French Village, Eggert,
Jy. 1886 (Mo), shelby: Windsor, Gleascm 748 (G). union: Cobden,
F.S.E., Jun. 1879 (CM). Missouri: "upper Missouri," Geyer, ann.

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[September

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Enaelmanv Ancr isaa /tv/t \ v-^"^";- Ji^*FERS0N; Aimmswick,

non: Mo'n&r "^S S (Zf ' f ^^'™T' AJ(Mo S

Marshfield Part„<i,.. jy. jg^g (w);t'KfcH'T: MaLffi piTS
{mo). Arkansas: Baxter- Cott<^r P^/^.,* f:o^/ nt^r \ ^^^

Plank, ann. 1899 (in part) Wsf clERot^ Fnr b ^^° " "^o''T'"'=

Mammoth Spring, Demaree 5^75 (US) jeffe Jo^ T.ff^i' r"°~^
i'mneH 10668 (ANS) prLAmri • T Ittt ^ u^^n "If «f ^^on Spnngs,
tian: Fort SmV ILJJ^nn iS^Jl mc^ ^"^"^ ^^ ^^^ • sebas-
ville, Harva, 4(ANS CM m' Mo^^^^" ^^^^'^^^n: Fayette-

aucun cil, iZniette? nulles r,7"''/T'''^*"'"^"* glabres, sans
la veiUe de I'anthfee dressi n,.;?"^ ^galant moitig du bouton k
delafleur." (PlSe 436 -C »"" ^J^ ^*?'f.> ''^P^nouissement
the "Flora VirS " a^ ,te^^ if "^"^'""!.-'?/'" '^^°^ ^^ of
This is probabfy i U?r J^Mx C ^'^"J^'^^H ^K"^'' V": 374).
plant (Clayton n. 66^bZS'LuZZT I'^f ^ '***^' *''«* t^is

states that this is a synonvm for ri»,,V - i °^- ^ichaux also
L. goodenioides WiUde^w l^rt R Yi"" ? P'""* o^nt'oned above.

Muhlenberg, Cat. rS &pt 22 ,8,?'''*t|''M 'l^' ^- ^'^^'^
name as a synonym for T oW^Vfv «r.?J Muhlenberg gives this
Muhlenberg herCium ar/l!^^^^'"'^-'>x' ^^^ «^t«™' '" the
Philadelphil isin sich 'n^r t a^^^'H'^ ."^ Natural Sciences of

mine the ident^^; of i pS'l*^'*!.:" «"« '* '^ ''"P°^'"« *" ^eter!
I: 15. 1820. i Wa tt' f ■ T" ^fi"««q"e. Annals of Nature

Stem strict, unbranXdL3;rr'^7"i^™y' S^"- F'- 6- 1878.-

\

t

/

1936J McVaugh,— North American Species of Lobelia 309

3-20, usually not appressed to the stem, thin, sessile, or the lower
narrowed mto short margined petioles; the lower obtuse, oblanceolate,
oblong or obovate, as large as 2.5 X 10 cm, shallowly coarse-dentate
or sub-entire. Upper leaves gradually smaller, acute-lanceolate,
often becommg more conspicuously denticulate above, and sometimes
mergmg imperceptibly into the bracts of the inflorescence. All leaves
strigose-pubescent above and below, especially near the margins and
near the base of the plant. Basal leaves, if present, obovate, obtuse,
pubescent, 1-12, narrowed into well-defined petioles. Inflorescence a

Fig. 15. Range of Lobelia spicata, var. originalis.

terminal virgate spike, 20-30 (60) cm. long, usually less than half the
height of the plant, interrupted, not noticeably secund, bearing few-
100 (in cases of fasciation 200 or more: the average is 30-60) flowers
upon short (in fruit 5-8 mm.), slender, rough-puberulent pedicels,
each with a pair of inconspicuous bracteoles near the base. Flower-
bracts smooth (rarely ciliate-pubescent), narrowly lance-linear, about
equalling the pedicels, or larger below, lanceolate, to 2.5 cm. long.
Calyx in anthesis flattish or broad-conic, smooth or sparingly pubes-
cent, becoming hemispheric in fruit, strongly ribbed, about 3.5 mm.
in diameter. Capsule }/^% inferior. Calyx-lobes subulate to deltoid,
usually flat, 2.0-7.5 mm. long, smooth or somewhat bristly-ciliate;
auricles usually present at the base of each lobe, distinctly short-
triangular, or longer; in extreme cases filiform, as long as 1.0 mm.;

310

Rhodora

[September

1.^

i

/

wMtf tTn t "^- .•^v.'ir' ^^2 '"'°- '""S- '"«'"ding calyx. Corolla
wh te to dark purpIish-blue, smooth outside, the lower Kp pubescen?

fLt^ fT r'\ Corolla-tube entire, except for the dorsal fissure

^Ja 'Y '°'''' ''P.T'I' '''«^*'y '^'"^' *''«" the tube, not sha"pK;
reflexed two upper lobes lanceolate, curved upward. Filament-tube

l^^A^'uTT^ ?-^" "'"'•) """• '""g. Pubescent belo" connate
about half Its length above. Anther-tube 1.7-2.0 mm. long light
buish-gray the two smaller anthers each with a tuft of wh"te hairs
a the tip; the three larger smooth or merely pubescent on the back?
Style included in anther-tube. Meadows and thickets preferring
moist, rich soil (frequently in pastures and hayfields • becomC raZf
weedy); New Brunswick to Pennsylvania, and^south'inThe mountls
to Georgia; west to northern Michigan, eastern Minnesota and extern
Missouri; southern North Dakota; north-western Arkansas Inter
grading freely in the western part of its range with the vl^' Airtel"

^fl:S^"^*n- ?r ''""''' P'''"*^ '" the Eastern Stetrwkich are
referred to v^t hrtella seem rather to be more or less ciliate variant, of
typical var. onginalis. Flower June 15-August?. appaS a fe^^

G scan ilfi^roT ri «'=«™o»=I'A';?: Moncton, BriUain, herb,
v^b. Can. 15266 (O). Quebec: hull: Hull, Harrington, Jv 14 1905
(Toronto), rouville: Abbotsford, Knowlin, Aug 1923 (GMof
Ontario: bbuce: Bg (=Dorcas) Bav ICrntlrn^ 'yev^ /rr' <"
=i f-dwich, ^iouWgroS ^^tf (h fb. &Ca.^-
Wil^ l^vf "^"T"-^" ^*°""'«' Jy- 1896 (Toronto). MroDLESEx-

S lugl fexoT- f7^ i?"°"'"*°>- '"''''^'"^■- P^rt Sydney,"
^wy Aug. 7, 1907 (Toronto). Maine: Androscoggin: South Poland

^«rfc«A, ann. 1893, 1896, 1897 (NE). Cumberland- Cumberland

Ckamberla^n Jy. 1903 (US, W). franklin: Farmrngto„"lX,l'

NB NF^hi w^^ ^- "sc^TAQure: Dover, Femald 311 (G. Mo

rNE)'^\o'i'K:''iennTbunr^S "f^f^^' l^"^' 'y' ^^

rr'-M Wo'fboro^" /IS g. 926''(W) ""c^'e:
Jaffrey, Rohmon 2S1 (G, NE). coos GoA^T Pease 16706 mfv
perhaps nearer var. hirteUa. grapton: Hanove;, ^Xm/ Jv lIlO
(G). hilisborough: Peterboro, BatcheUkr Tv iQici ^rra ^"
mack: Hooksett, Batchelder Jv 1021 nwS ^" ^ ^^- '"^'""■

Batchelder.Sy 1913 (NB Np/^Vl!^ ^^^- "O^kinoham: Derry,
Dm, 19.1 rr TTo\ ^ ' *'■'• Vermont: bennington: Manchester
i^ayl^l ((j, US). CALEDONIA: Lyndon Jv 21 1S7^ CMR^ """l"*'*'^^"^'
Fairlee Dev^lmn T„ iooq ,^t„v"'J'' •^'> ^o'o (J^H). orange:

74^3 (NBl^' ^i^?,^ ^^?^ '"'Ti'*N'>: Clarendon, EgglesUm
'■^o u>D, us;. WINDHAM: Vemon. Robinson A„„ ibqc r<^\

WINDSOR: Queechee Gulf, Kennedy Jv l^^T' i^ ^' ^^^•

BERKSHIRE: Stockbridge u2naknl^T 1^2 ,^tr^«"SETTS:
Swansea, Sanford WSsI (,.n)%"Sy:i>tZoS F^^i l^T^

4i

t

1936] McVaugh,-North American Species of Lobelia 3n

(NE). DUKES: Chilmark, Harri'< Tv isos rKrt^\ « ,

(G, Mo, US), franklin: BucUand.7oV6i Li f^08 mv^' ^"^^
den: Granville Sevmmir to^ '"• •^''™««. Aug. 1908 (NE). hamp-

et al.. Jy. 1931 (NeT Middlesex- F^rW^T"^ ^ Prescott.Goodale

(CU, UP). NORFOLK GrantXL«jTfcMtr^ '^^ '^

fylmi^K^'^'""'"'' ''■ i"o« w:'LS^LereTr;

Warwick, Fernald, Jun 1910 (G WV^^ ^ ^^^)' ^^^t:

haven: New Haven SaiTord 2P<? ?;',' y'^*'^"*"" 1*^3 (Mo), new
Woorfjoard Jv 2 1906 ^^1!! \7^^- ""^"^ mndon: Franklin,
lRJ.ni (ch\ ' ^^r . ^^ Yo»K= albant: Elsmere Housl-

IS4OI (CU). BROOME: Binghamton, MiUsvauah Jv iswrVr^
CATTARAUGUS: Quaker Bridgt, ^tea^d^ aC 1926 rMV«> ^™^-

27T^f m^"?' ^'^ ^"^"^ I^B)- ™NGof-B risLS. 5un"
27, 1887 (US). COLUMBIA: Kinderhook, McVauoh ^^7? rr^T Mvc'
UP). DELAWARE: Bo Vina. ^01/ Jv i!iQ9 (rrn^t ^^^Ar*^^'
Mt., McVaugh mz (CU NYs' UP) Essix M' '"'^'^^^^■- Stissing
(NYS). GREENE: Platte ChltwilliarZ^jJfS^r,''^:^^'' '^'^^
ton: Back Log Camp, M /ie^ /S?1m«; ^ ^^^^S^- ^'^^^

^x:; At 'r^hon''^' ^^^^ s^-' ^b) t^Lo^fui oi-

7t^V%\ ^' 1^2 (CU). queens: Richmond Hill Bicknell /?/■??

1099 Amo\ ■'• t"^^^"- ^l^e Mmnewaska, Sowdm .Tun -Iv

1922 (ANS). Washington: Shushan, DobHn 166(N\^\ I' ^'
CHESTER: nr. Croton, PenneU 767'i rAM^r m t '^^^^f- ^^st-
Westwood PenneU 9L^ZV<\ ^ ^'- New Jersey: bergen:
r»MnZ T- J ij o (^NS). burungton: Burlington fANS^

gZ Aue IsT^M^f^ ^^''^ ^^NS)- '^''"BERLANDrVineland

fiSsbufy, II T^ ISa^rp';-- «'""'''- (NB). HUNTERDON:

m (CCD) OCEAN- Iffiirhnffn/ r^^- """""v Chatham, Mackenzie
an error h^locahty) PAfsfic PomrTl"' "^^>'^,^ ^^^^ P^^aps

somerset: y^p^u^TiLliTir^tS'^^: f/ ^^|^

McDowell tno lAKo^ -*'"«'^ Jy- ISf9 (CM), bucks: Neshaminv
^cuowetl S09 (ANS). bvtler: Saxonhurg, Shafer 65 (CM CfT P.

jLn^ig^fmpV ^'''«''*°" i^NS^- ^^'-^''^ Juniato Jet ' ^w'
Jun. 1934 (UP). CHESTER: W. Chester, Wm. Darlington, sLrtZA.

» I

k*

312

Rhodora

[September

(ANS). CRAWFORD: Hartstown, Jennings, Jun. 1922 (CM), dauphin:
Harrisburg, Small, Jun. 1888 (G). Delaware: Newtown Twp.,
MacElwee 603 (ANS, G, Mo), franklin: Mercersburg, T. Green,
May 1845 (ANS). Lackawanna: Dalton, Twining, Jy. 1907 (CM).
LANCASTER: Reinholds, Small, Jun. 1890 (US). Lancaster, Porter,
Jun. 1857 (G; type of var. parviflora Gray), lehigh: Allentown,
Pretz 6716 (ANS). luzerne: Beech Haven, Heller, Jun. 1889 (ANS,
G). mercer: Sharon, Aschman, Jy. 1886 (CM), mifflin: Lewistown,
Jennings, Jy. 1908 (CM, Pa), monroe: Pocono Lake, Harshberger,
Jy. 1904 (ANS, Mo). Montgomery: Tylersport, Long 24840 (ANS).
NORTHAMPTON: Bethlehem, Moser, Jy. 1832 (US). Philadelphia:
Falls of Schuylkill, Jeanes, Jy. 1828 (ANS). somerset: Stoyestown,
Patterson, Jun. 1880 (CM), venango: East Sandy, Garber, Aug. 1869
(ANS). Washington: Charleroi, Jennings, Jun. 1904 (CM). West-
moreland: Ligonier to Donegal, Jennings, Jun. 1904 (CM). Dela-
ware: NEWCASTLE: Stantou, Randolph 101 (CU, G). Sussex:
Frankford, Commons, Jun. 1875 (ANS). Maryland: garrett:
Oakland, Shreve 651 (US), harford: Flintville, Adams et al. 929
(ANS). Virginia: Bedford: Curtiss, Jy. 1871 (G). Campbell:
Lawyers Road, Heller, Jy. 1893 (ANS). West Virginia: Green-
brier: Cranberry Summit, G, Guttenberg (CM). Pocahontas: Rimel,
Core 3488 (WVa). North Carolina: buncombe: Biltmore, Biltmore
herb. 626 (CU, M, US); Asheville, Kram, Jun. 1925 (W). haywood-
True Love Mt., Blomquist 6031 (Duke), jackson: Cullowhee,
Thaxter, ann. 1887 (G). swain; Great Smoky Mts., Beardslee and
Kofoid, Aug. 1891 (Mo; near var. leptostachys). Georgia: floyd:
Rome, Ravenel (Mo). Alabama: Talladega: Riddell's Mill, Mohr
(US; near var. leptostachys). Tennessee: knox: Dead Horse Lake
Sharp, Jun. 1930 (G). Ohio: clark: Springfield, Williams (Mo)
CUYAHOGA: Berea, Ashcroft, Jy. 1897 (NB). erie: Castalia, Jennings,
Aug. 1911 (CM). FAIRFIELD: Sugar Grove, Werner 98 (G) frank-
lin: Columbus, W.S.S., Torrey herb., ann. 1840 (NB). mahoning-
Perkms, Moseley, Jy. 1894 (Mo), summit: Akron, Ashcroft, Jy 1896
(Mo), wood: Bowling Green, Moseley, Jy. 1919 (Mo). Indiana-
LAGRANGE: Mongo, Beam 39084 (ANS). lake: Pine, Beam 43306
(CCD). noble: Kendallville, Beam 36624 (ANS). porter: Crisman
Beam 31664 (part) (CCD). steuben: Lake Gage, Beam, Jun 1903*

/^SRx ^' ^^' ^^' ^^' ^)* warren: Rainsville, Beam '26874
(CCD). Illinois: champaign: Rantoul, Gleason, Jy 1907 (G)
DUPAGE: Naperville, Wellner, Jun. 1896 (W). jo daviess: Portage
Lansing 4091 (ANS). lake: Beach, Umbach 3712 (W). lasalle:
Ottawa, Huett (G). mc lean: Bloomington, Robinson, Aug 1886 (G)
Michigan: bay: Bay City, Breisbach 6968 (ANS). Cheboygan*
Indian River, Ehlers 2968 (W). chippewa: Sault Ste. Marie Mc-
Mullen {G). crawford: Grayling, Mell 214 (US); Piper, Jy.' 1922
(US; G, m part), ingham: Haslett, Yuncker 4I6 (US) ionia-
Hubbardston, E. F. Smith, ann. 1877 (G). mason: Ludington'

-<•

1936] McVaugh, — North American Species of Lobelia

313

Chaney 12 (US). Menominee: Menominee, Schuette, Jun. 1891
(US, W). midland: Midland, Breisbach 102 (ANS). st clair: Port
Huron, Glatfelter, Jy. 1888 (Mo), washtenaw: Whitmore Lake,
Ehlers 1488 (G). wayne: Detroit, Lugger, Jy. 1891 (M). Wisconsin:
ADAMS: Friendship, Rhoades, Jy. 1927 (CU). brown: DePere,
Kellogg, Jy. 1888 (US), buffalo: nr. Fountain City, Finkelnburg
(CM). CRAWFORD: Prairie du Chien, H. H. Smith 7636 (W). dane:
Madison, Sprague 1200 (W). dunn: Menomonie, Bachman, Jy. 1928
(W). EAU CLAIRE: Fall Creek, Kunz 220 (W). 10 wa: Dodge ville,
Breakey, Jun. 1929 (W). jackson: Black River Falls, H. H. Smith
6796 (W). JUNEAU: Camp Douglas, M earns 629 (NB). Lafayette:
Fayette, Cheney, Jy. 1888 (W). marathon: Mosinee, Cheney 3270
(W). MARINETTE: Athclstanc, Bavis, Jy. 1915 (W). Milwaukee:
Milwaukee, Hasse (NB). walworth: E. Troy, Almon, Aug. 1926
(W). WAVPACA'.Re&d&eld, A. Smith 110 (yV). waushara: Richford,
Taylor, Jy. 1932 (W). Minnesota: cass: Bridgeman, Sheldon 3289
(part) (M). mille lacs: Princeton, Sheldm 3116 (M). olmsted:
Rochester, Ainslee, Jy. 4, 1902 (M). scott: Savage, Rosendahl,
Sep. 1924 (part) (M). wabasha: Lake City, Manning, Jy. 1892 (M).
WASECA: Janes ville, Taylor 650 (M). Missouri: butler: Poplar
Bluff, Eby, Jy. 1893 (Mo), iron: Pilot Knob, Glatfelter, Aug. 1895
(Mo). ST. LOUIS: Engelmann, May 1836 (ANS); Eggert, Jun. 1879
(Mo). Arkansas: Washington: Fayetteville, Watts, Jun. 1925 (W);
this plant is perhaps related to L. appendicidata. North Dakota:
KIDDER: Dawson, Metcalf 274 (CU).

c) Var. hirtella Gray, Syn. Fl. 6. 1878. Type Locality:
"chiefly towards and beyond the Mississippi." Type Specimen:
authentic material seen in the Gray Herbarium. — L. hirtella Greene,
Pittonia III: 349. Sep. 27, 1898. — Differs from the var. originalis in
being bristly-pubescent nearly throughout; the lower stem is bristly,
especially on the angles. Flower-bracts densely bristly, especially
near the margins; the lower usually lanceolate, leafy, to 2.5 cm. long,
often exceeding the flowers. Calyx densely bristly, especially on the
conspicuous ribs. Calyx-lobes usually narrowly lance-linear (some-
times deltoid), 3.5-7.0 mm. long, bristly; often with a conspicuous
raised midrib. Calyx in fruit hemispheric or varying to ovoid, 3.5-6.0
mm. long and 3.0-3.5 mm. across; capsule 3^^ inferior. Auricles of
the calyx-lobes often conspicuous. Plants as a rule smaller than var.
leptostachys or var. originalis; mostly 15-60 cm. high. Inflorescence
usually 30-60 flowered. Leaves in many cases clustered on the lower
half of the stem. — Low open meadows and prairies; sometimes in dry
soil, north-western Indiana to eastern Kansas, northwest to western
Nebraska, South Dakota, northern Minnesota, Saskatchewan (north
to the 52d parallel), and Alberta. In nearly or quite typical form
passing eastward through Michigan, northern New York and New
England to the Gasp^ Peninsula; apparently common on Long Island.
Flower June 1 -August 1 ; somewhat earlier in the southern part of its

I f^

V

vl

if

314

Rhodora

[September

Kro^orrhi (A r.r^- ^'S'*'«o- BRUCE: Big (= Dorcas) Bay
j\roiKO^ /HSd (U). DURHAM: Hampton AUin Tv TQ97 rv fK'
ESSEX: Leamington, Macoun herh K S'rf' L,lr ]?fJ (Toronto).

Belleville. Maclun in5(GyMlun!^ FCn^ f ^^^^ "^'"^^^
Sydney, /rev Jv 31 ^Qna/'^Z^^rJ•°'^°''^°^■ "uskoka: Port
f^at^J^'imnlroniJ, mIZ st Timagam, Forest Reserve:
S89 (NE) cuMBERLAvn R ''""ROScoggin: E. Auburn, MerriU

fra^ J:- cSrWlirCAlrZf'jft^l TnE) '''' ^^^>-
Swan Island. ^.^ ..,. (NE). oxr^of^NeiS! SL,T?^^

Fio. 16. Range of Lobelia spicata, var. hirtella.

(G). New Hampshire: coos: BandolTAZlshrl^^f^ (??'*)
Jv 19^ nVTnT mI^' V ^^^^- RUTLAND: Brandon, Z)i/«on

if27TurLfrN^rmr='^^^/r^'^^^^^^^

Hempstead, GmAot/ 595 rPTT^ o^ r (^, NYS). Nassau:

(G, NB). 'su^roTf :^Bab5o^^- cZieZ^lcm ^''°"' '^^^'^ ^^^
Clarke. Umbach 3867 (W) lake 0«rv l/^^' ,^'">'*n*- ci^kk:
newton: Morocco. Aeam S rcrhf °^' "^""- ^P (^"''«)-

f

!^

4

t.;

A.

<

.A

4':r

«•»'

i

I

i

1936] McVaugh, — North American Species of Lobelia

315

Augusta, S, B. Mead, Jun. 1859 (ANS). Henderson: Oquawka,
Patterson (ANS, US), lake: Beach, Johnson 1821 (NB). mc henry:
Ringwood, Vasey ann. 1862 (G). mason: Kilbourne, Gleason, Aug.
1903 (G). MENARD: Athens, Hall, ann. 1861 (G, Mo), ogle: Hol-
comb, Beck, Jy. 1907 (W). peoria: Peoria, McDonald, Jun. 1903
(NB); Jun. 1904 (G). rock island: Joslin, Harper, Jy. 1886 (W).
stark: Wady Petra, Chase 692 (ANS, Mo), will: Romeo, Powell,
Jun. 1898 (W). WINNEBAGO: prairies, Behh, Jy. 1858 (G). Michigan:
CHEBOYGAN: Burt Lake, Ehlers 4634 (CU). keweenaw: Keweenaw
Point, Wood 1642 (NC, US); sterile hills, Farwell, ann. 1889 (G; fid.
A. Gray), monroe: Raisinville, Atkinson, Jun. 1883 (CU, US).
Wisconsin: buffalo: nr. Fountain City, Finkelnburg (M). dane:
Primrose, Fassett 3459 (G, NB, W). lacrosse: LaCrosse, Trelease
(probably), Jy. 1887 (Mo), racine: Racine, Davis, Jun. 1881 (W).
rock: Janesville, Skavlem, Jun. 1889 (W). sauk: Baraboo, True
( W) . VERNON : Viroqua, H. H. Smith 721 8 (W) . wal worth : Dela van,
Hollister 25 (US), winnebago: Winnebago, James, Jy. 1894 (W).
Minnesota: blue earth: Leiherg, ann. 1883 (M). brown: Spring-
field, Sheldon, Jy. 1891 (M). chippewa: Montevideo, Moyer, Jy.

1891 (M). CHISAGO: Sandberg, Jun. 1886 (M). clay: Glyndon,
Solheim 165 (R). douglas: Lake Christina, Sheldon 3466 (M).
HENNEPIN: meadows, Sandberg, Jy. 1890 (M, W). Houston: Spring
Grove, Rosendahl 477 (M); 501 (G). jackson: Heron Lake, Skinner,
ann. 1902 (M). Kandiyohi: Spicer, Frost, Jy. 1892 (G, M, W).
lake: Echo Lake, Barber 12 (G). nicollet: Nicollet, Ballard, Jy.

1892 (M, R, US). OLMSTED: Rochester, Ainslee, Jun. 19, 1902 (part)
(M). OTTER tail: Fergus Falls, Sheldon 3699 (M). pipestone:
Pipestone, Sheldon 1475 (M). polk: Crookston, MacMillan and
Skinner 57, 297 (M). pope: Glenwood, Taylor, Jy. 1891 (M). Ram-
sey: New Brighton, Moyer, Jun. 1897 (M). st. louis: Duluth,
Johnson 1263 (part) (NB). scott: Savage, Rosendahl, Sep. 1924
(part) (M). STEARNS: Pleasant Lake, Campbell Cl65 (M). waton-
wan: St. James, Fowler 25037 (W). winona: Winona, Holzinger,
Aug. 1890 (US). Iowa: benton: Vinton, Davis (W). black hawk:
low prairie, Burk 388 (Mo), decatur: J. P. Anderson, Jun. 1902
(Mo, R). emmet: Armstrong, Cratty, Aug. 1898 (US), fayette:
Fayette, Parker (M). greene: Grand Junction, Wiegand et at. 2401
(CU). JOHNSON: Iowa City, Somes 3202, 3204 (US), kossuth:
Wesley, Breithaupt, Jun. 1898 (W). story: Ames, Ball and Combs
500 (G, Mo, R, US). VAN buren: Bentonsport, Graves 1894 (Mo).
Missouri: chariton: Young, Jun. 1925 (CM), jefferson: "Crystall
City," Eggert,Aug. 1886 (CM), marion: Hannibal, Davis 1190 (Mo).
NODAWAY: Marysville, Palmer 25434 (Mo). North Dakota: "upper
James River," (no state given), Geyer, Nicollet's N. W. Exp., Jy. 13,
1839 (US). BENSON: Leeds, Lunell, Jy. 6, 1901 (ANS, M, R, W).
CASS: Buffalo, Westergaard, Aug. 1897 (R). grand forks: Grand
Forks, Brannon 240 (Mo), nelson: Tolna, Jy. 25, 1911 (R). rich-

V.

1-1

^ *

4^

/

316

Hhodora

[September

Jy. 21, 1893 (ANS, M, Mo), custer: Custer, Rydberg, 850 (US)
day: Waubay, Moore 65 (M). deuel: Toronto, Mole 885 (M
T^''^^kl^f^°''\'Ir^• "^"°- 1895 (M). moody: Flandreau. Griffith,',

1858 (Mo.), cedar: Aten, Clements 2656 (CU Del G M TI«?1
cherry: sand-hills, SmUk and Pound 37 (Mo), custer- 'Call'awrv
Bate, mSJfl). orakt: Whitman, Rydberg 1818 (G US) keaS
Mmden, Hapeman. Jy. 1897 (W). Pi.ArrE: Columbus, PenneuTsOU
(ANS). SC0TT8 bluff: Platte Bottom, RyMerg 2n (US) thoma«-
Halsey, War, Jy. 1907 (UP). Ka/sa8: "LErvENwoRTr-Ft"

Jy. 1883 (US). Manitoba: "N. W. Territory," NicoUet 4H (ANS)

Can. 14146 (O, US), soueis: Turtle Mt., T.J.W.B. 139, 3y 1874
(Toronto). Winnipeg: Winnipeg, Scott, Jy. 1904 (Toronto/' Sas
katchewan: "20 m. W. Yorkton? along Grand Trunk ScR R^

Can^4^?rn ^''^ ^"l** Saskatchewan," ^ao<mn, herb. G. S

iTo^;^ /^N ^°^- ^siniboia: Souris Plain, Macmn, herb G. S Can
15254 (O). humbou)T: Wadena, itfcC^jare (Torokto); Touchwrd
Hills, Macoun and Herriot, herb. G. S. Can. 78480 (0 AlC^
craigmyle: prairie, "SE 28^2-16 W 4," Brinkman, Jy 1923 (UsT

Note: Wh. e the above are all referred to the var AtVfe«a, it should
be observed that the variety reaches its most character stic form

Prl'w*-''' ''^'°" ^"'* "i '^' Mississippi, and occasionally eastwar™
From Wisconsin eastward, most plants seen lack the short stature of
the prairie plant, and are placed in var. hirtella because of the combi-
nation of general bristly pubescence, combined with leafy Zer-

^ulJ"' «?"P«»^»t*. var. nov.; caule paucifloro. calycis tubo
oblongo vel hemisphaenco, antheris albis, capsulis subinflatis Type

widely spaced flowers upon short (in fruit 4-5 mmT rather stout
rough-puberulent pedicels, each with a pair of inconspicuous brS
oles near the base. Flower-bracts smooth, linear, somewhat lone^
than the pedicel (5-9 mm. long), callose-d;nticulate "x in bud

» •■

«^.

"j

^i

I

^

^«

1936] McVaugh,— North American Species of Lobelia

317

and in anthesis short-campanulate, smooth or slightly puberulent,
becoming sub-globose in fruit, strongly ribbed, 3.5-5.0 mm. across
by 4.5-6.0 mm. high. Capsule % inferior or more. Calyx-lobes
short-lanceolate, flat, 3-4 mm. long, smooth, entire; auricles lacking.
Flower 7-9 mm. long, including the campanulate calyx. Corolla
distinctly dark-purplish, smooth, except for the slightly pubescent
base of the lower lip. Corolla-tube entire, except for the dorsal
fissure; lobes of the lower lip ovate-oblong, slightly shorter than the

Fig. 17. Range of Lobelia spicata, var. campanulata.

tube, not sharply reflexed; two upper lobes lanceolate. Filament-
tube 2.0-2.5 (3.0) mm. long, slightly pubescent below, connate about
half its length above. Anther-tube 1.0-1.5 mm. long, white, the two
smaller anthers tufted, the three larger smooth or pubescent on the
backs. Style exserted, recurved. Seeds apparently somewhat larger
than in var. originalis.

This plant differs from var. originalis by the campanulate calyx,
the globose, larger capsules, white anthers, smaller flowers in every
way, fewer flowers, earlier flowering period (7-10 days in eastern
New York). It probably does not deserve specific rank, as plants
with white anthers and small flowers are sometimes found in the other
varieties of L. spicata. This phenomenon may be one of arrested
development or some other abnormality, as similar conditions are
seen occasionally in other parts of the genus (e.g. L. puberuld). How-
ever, when the earlier flowering period is considered, as well as the
definite changes in calyx, capsule and inflorescence, it seems desirable
to separate this plant from var. originalis.

Everywhere with var. originalis, but apparently rare except on the
Eastern Seaboard from southern Maine to Pennsylvania; from
Pennsylvania to Virginia it intergrades freely with the var. scaposa.

ki

\

II

318

Rhodora

[September

1936J McVaugh,— North American Species of Lobelia

319

Flowers a few days earlier than var. originalis; from June 1. Material
seen: Maine: Hancock: Mt. Desert, fields nr. Long Pond, Redfield,
Jy. 1892 (part) (NE). york: N. Berwick, Parlin, Jy. 1892 (1 sheet)
(G). New Hampshire: coos: Dalton, Pease 12160 (part) (NE).
Vermont: Addison: Ferrisburgh, Horsford, ann. 1882 (part) (G).
Massachusetts: Berkshire: Tyringham, Vail, Jun. 1897 (NB).
BRISTOL: Swansea, Sanford 10336 (part) (NE); Dartmouth, Fernald
1068 (part) (NE). essex: Montserrat, Hubbard, Jy. 1913 (NE); S.
Georgetown, Williams, Aug. 1899 (NE). Middlesex: Concord, Deane,
Jy. 1886 (NE). S. Acton, Deane, Jy. 1886 (part) (US); Chelmsford,
Fames, Sep. 1914 (CU). Norfolk: Needham, Fuller, Jy. 1883 (part)
(NE); Norfolk, Ware 2485 (part) (NE). Connecticut: Chas.
Wnght (part) (CM). Hartford: Southington, Bissell, Jy. 1897
(part) (Mo), new London: Franklin, Woodward, Jy. 11, 1906 (G,
NE). New York: Columbia: Kinderhook, McVaugh 2673A (CU,*
NYS, UP). DUTCHESS: Madalin, McVaugh 267 4 A (UP), st. law-
rence: Lisbon, Phelps 914 (CU). New Jersey: burungton: Mt.
Holly, Meredith, Jun. 1922 (ANS). Pennsylvania: berks: Lobachs-
yille. Long 12632 (ANS). bucks: Upper Black Eddy, True, Jy. 1925
(part) (UP). CHESTER: Paoli, Pennell 3905 (part) (UP) ; West Chester,
Pennell 1224 (ANS). Delaware: Haverford, Burk (part) (UP);
Lenni, Redfield 4516 (Mo); Wawa, Pennell 21 (part) (ANS); William-
son, Pennell 194 (part) (ANS). Lancaster: Lancaster, Stevens, Jy.
1874 (part) (Mo). Montgomery: Penllyn, MacElwee, Jun. 1903 (1
plant, ANS). Northampton: Easton, Porter, Jun. 1896 (ANS).
Delaware: Newcastle: Centreville, Commons, Jun. 1866 (part)
(ANS). Maryland: garrett: dry open woods, J. D. Smith, Jy. 24,
1883 (US). Ohio: stark: Canton, Steele 14 (US). Indiana: clark:
Deam 6897 (part) (CCD). Lagrange: Mongo, Beam 20706 (CCD)
lake: Hammond, Deam 1456 (part) (CCD). noble: Eagle Lake!
Dernn, 31252 (part) (CCD). porter: Crisman, Deam 31564 (part)
(CCD). Illinois: Vasey, herb. Olney (G). cook: Chicago, Bab-
cock, Jun. 1874 (part) (US), lake: Waukegan, Umbach 5840 (W)
WHITESIDE: Fulton, Harper, Jun. 1892 (W). Michigan: Crawford*:
Grayling Piper, Jy. 1922 (part) (G). wayne: Detroit, grassy fields,
tarwell, Jun. 1900 (part) (G). Wisconsin: dane: Madison, Trelease,
Jun. 1889 (part) (Mo). Minnesota: anoka: Centreville, Sandherg,
I'r3J^^^ ^P^"**) (^^^)' "Northern A. Co.," Butters and Rosendahl
5056 m). CASS: Bridgeman, Sheldxm 3289 (part) (M) ; this also shows
Its affinity to var. hirtella. olmsted: Rochester, Ainslee, Jun. 1902
(part) (M). Iowa: decatur: Fitzpatrick, May 1896 (part) (Mo).

e) Var. scaposa, var. nov., foliis radicalibus ovatis obtusis petiolatis,
caulims raris parvis, racemo elongato laxo, calycis appendicibus non
^^^\r N Locality: "Danville to Fall Creek" (Pittsylvania

Co., Va.). Type Specimen: Small and Heller 108, Jun. 3 1891 in
the University of Penna.— Possibly L. pallida Muhlenberg, Cat.'pi
Am. Sep. 22. 1813. Probably L. pallida Elliott, Sk. Bot. S. C. & Ga.

I: 265, 1821 (in part). Probably L. paniculata Rafinesque, N. Fl. N.
Am. 11:18. 1836. i. «i)ica<a Hitchcock and Standley, Fl. Dist. Col.
263. 1919.— Stem strict, unbranched, 40-110 cm. high (ave. about
65 cm.), green, or with a slight reddish tinge near the base; smooth,
or pubescent, especially on the angles, below. Cauline leaves incon-
spicuous, thin, 1-6, usually only about 3 in number, all below the
middle of the Ltem, all acute, bract-like, 1-2 cm. long, lanceolate,
sharp-denticulate, or the lowest one oblanceolate or oblong, obtuse,
about 1.5 X 5.0 cm. (in extreme cases 2.5 X 9.0 cm.). Basal leaves
1-12, conspicuous, obovate, ovate or round, narrowed into margined
petioles, 1.5-5.5 X 3-10 cm. (ave. about 3.5 X 5). All leaves more
or less strigose-pubescent, especially near the margins and near the
base of the plant. Inflorescence a loose terminal raceme, 10-60 cm.
long (ave. about 30 cm.), usually about half the height of the plant,
appearing naked, because of the disproportionately large basal leaves;
somewhat interrupted, sometimes sub-secund, bearing 10-90 (ave.
about 40) widely spaced flowers upon short (in flower 4-6 mm.)
slender pedicels, each with a pair of inconspicuous bracteoles near
the base. Flower-bracts smooth, lance-linear or sometimes narrowly
lanceolate, about equalling the pedicels, 4-10 mm. long. Calyx in
anthesis broad-conic, smooth, becoming hemispheric to sub-globose in
fruit, about 4 mm. in diameter. Capsule (where seen) % inferior.
Calyx-lobes lanceolate to linear, 2-6 mm. long, smooth, entire;
auricles usually conspicuous at the base of each lobe, triangular or
short-filiform. Flower 7.0-10.5 mm. long, including calyx (ave.
9.0 mm.). Corolla pure white to light blue, the lower lip pubescent
at base inside, otherwise smooth. Corolla-tube entire, except for the
dorsal fissure; lobes of the lower lip ovate, shorter than the tube,
sometimes sharply reflexed in dried material; the two upper lobes
lanceolate, recurved or erect. Filament-tube 3.0-3.5 mm. long,
pubescent below, connate about half its length above, often deflexed,
so that the anther-tube and the upper part of the filament-tube are not
contained in the corolla, but project above it, through the dorsal
fissure. Anther-tube 1.7-2.0 mm. long, light bluish-gray, the two
smaller anthers tufted at the tip, the three larger smooth or pubescent
on the backs.

Differs from var. originalis by the larger basal leaves, fewer stem-
leaves, the naked inflorescence, which is longer in proportion, the
more conspicuous auricles, slightly smaller flowers, broader calyx,
earlier flowering period. Is sometimes confused with var. lepto-
stachys because of the auricles, which may be conspicuous, but the
almost naked stem should prevent such a mistake. — Low open woods
and hillsides, in moist or dry situations, southern Pennsylvania to
central North Carolina, central Mississippi, and Louisiana. Appar-
ently mostly confined to the Piedmont. Flower May 20-June 20.
Material seen: Pennsylvania: Rev. J.H.B. (Porter herb.), Jun. 10,
1845 (ANS). franklin: Mercersburg, DetmlUr herb., Jun. 2, 1845

\

i

3. 320

Rhodora

[SEPTEBfBEB

cl^S' /^o wp'^*' """• ^T ^^°)- '^^^«''-- Big Cove Tannery

and ^. Z). Snitk, Jy. Igk (US); cITmbXdtTXJ-" Jri894
(NB. BALTIMORE: J. D. Smith, Jun. 8 1880 (VS) ''nJliJ/' t

SrflJImsV^" '^ri' (^,^F- "ONTGOMEEY Glen Ecfo
maimer ISU (Mo); Kensington, House 1000 (Mo)- "J nrv
at Mass. Avenue," Bo^en f,e (UP). wZi^^i': Harp^n Fe^"!

Fig. 18. Range of Lobeua spicata. var. scaposa.

Kl^nSuiTsvK'A. ^Tr ,- C-™-- Washington,
•;nr. WashinVn^-- mSJdS ^m&tfi^"^ '""^ ^^U^''
ARLINGTON- Ft Mv»rW T '^ f°'^^^. TideHrom 6366 (US)

May 1934 (UP) Fl,RV^x^.>h^f„'=?JT''^'f.' , ^interpock. Wherry,
Great Falls, MoJremf(c\ a"^<^^' ^'^''^' •^""- ^^^ (NYS)

MONTGOMKRv: Bra;ksS,^^;.^rjr"i8^"^^^^ '"' (cy- «^'

Fall Creek, Sm^W "^ ll ^;^|'i!f , ^sV^A\1^'^=.^''"^'»« *»

PHINCE «^luam: BucklSilrMay'fgyS^'^r'^^' T"
Pennell 13326 (ANS) spotsyt vamt? ^ ^i • i J^^^-'' Gainesville,

May 1917 (CU) STAFFORn fJ^ i «^^^^^^ ^">^^ ^< «^.

\y^v). STAFFORD. Falmouth, W-z.^an(i arwi Manmng 3092

1936]

McVaugh, — North American Species of Lobelia

321

(CU, G). West Virginia: Hampshire: Hanging Rock, Frye, Jun. 7,
1933 (WVa); Hanging Rock, Koenig, Aug. 1887 (WVa); Hietts Run,
W.V.U. Bot. Exp., Jy. 2, 1926 (WVa). North Carolina: Chatham:
12 m. S. Chapel Hill, Totten, Jun. 1931 (NC). orange: Chapel Hill,
Tottm, Jun. 1915 (NC). person: Blomquist 5030 (Duke). South
Carolina: oconee: Newry, House 3J^77 (NB). Mississippi: choc-
taw: Ackerman, Jensen, Jun. 1905 (Mo). Louisiana: Orleans:
"nr. New Orleans, Dr. Ingalls," Torrey herb. (NB).

The following list is made up of specimens which seem not to belong

Fio. 19. Ranges of Varieties of Lobelia spicata: Var. leptostachys
(vertical lines); var. originalis (within heavy line); var. hirtella (withm
stippled line). Range of L. appendiculata (plain, within light line). Dots
represent plants not surely referred to any of the named forms; that is, inter-
mediates. Records from the Atlantic states and southern New England are
merely of occasional hairy individuals of the smooth var. originalis.

to any of the named varieties of L. spicata, but to possess features
peculiar to two or more of the varieties; that is, they seem to be inter-
mediates of various sorts. The initials in parenthesis after each
citation indicate the varieties which the specimen seems to resemble
most closely. The letters "app" stand for L. appendiculata. Vir-
ginia: STAFFORD: Falmouth, Wiegand and Manning 3093 (CU)

322

Rhodora

[September

/

t' ;

(sc-ca) Kentuckt: christian: "barrens," C. W. Short fANS^
(leFMJr). Indiana: allen: Ft. Wayne, Deam 1337 (CCD) (leD-^r)

WINNEBAGO: ^^66 Jv IS^iS rAMC5\ n«J N tIt ^ ^ Uep-Hir;.
Virgin (?) Lake, Hoffmann, Aug. 1918 (Mo) (hir-ca^ Mtmm^u^^;'

LOUISA: Columbus, Pa/mer ^055/ (NBWhir-^r) M>™, * ^'^^
Broadhpfjtl Mav ifi7i /a/t ^ /u • \ ^ \,nir-or;. Missouri: bates:
nromneaa May 1871 (Mo) (hir-or). Crawford: Woodsm 595 CMn^
(hir-IepK>r) greene: Springfield, Standley m^lv^ (^^^^^
iron: eastern Iron Co.." Treleasp RRQ (\/[^\ n n ^ ^^*
Waldo Park Bn^h A7Q(1^\ ttgwi u- 1^7 Gep-or). jackson:
/r AT xmN n 4/S' (Mo, US) (lep-hir-or); Martin City Bush SOHR

USWll^ ^^'^'^''^- '^f?«^ ^^^^ City, PdmerfoofTG Mo
US) (lep-hir). LACLEDE: Lebanon, Pmn.// ;7^i.? (paVt) rANS^'
(lep-or). OREGON: Alton to Thayer Trelease Tv ISQ-^ r?? wi^ V
polk: "Stockton Road," TreleZTm Mn\ (L \ ^ ""^ ^^^'^'^'

Z^^Cr^/rMTn ^^ ^ *''''^'')- «^- FRANCOIS: Bismarck,
x^tt^ar^ 7^ (Mo) (lep-or). st. louis: St. Louis, Monell Sen IsTs
Mo) (hir-or); St. Louis Eggert, Jy. 1875 (ANS CM L NB R
US) (lep-or). TEXAS: Plato, Emig 161 (Mo) (Cr^T\ I'S ' '
BENTON: Plank, ann. 1899 (part) (U) (lep-a^^^^^^^
Creek, Demaree 3137 (ANS) (^ - ano) ot^^. JL ^^^^^^^N-' ^^g

Idabel,//or.^Aton5^55(G,Mo NB) (aoo - T^^Z^' ""^ ''''^''^'^•*
A^orfoT. and Clothier 318 (G Mo R ^TN^ P* ^^''T' ^^^^^nsee:
Rosedale, 3f oc W>, jj. \ 896%^ ^ Wyandotte:

" H«h;.n/'''''^tr'' Lij^naeus, Spec. PL II: 931. 1753. Type Locality-
Habitat in Virginia, Canada." Type Specimen- There are fntY.*

r^^/I^^Wt'^'^^^^ V ^nia: i. -^

L. chffortiana Michaux, Fl. Bor.-Am II- 152 Ism P u x,^ I '
Sent II- 44S isi^ r ai- i '. ^r * ^^^'^- ^ursh, Fl. Am.
76 IsVs f 7 ^' ^*^^«^* Nuttall, Gen. N. Am PI IT.

76. 1818. L, caule erecto hrachiaio, foHis ovato-lanceola^ obsj^te

1936] McVaugh, — North American Species of Lobelia

323

incisis, capsulis inflatis, Linnaeus, Hort. Cliff. 500. 1737. This plant
was sent to Linnaeus by Gronovius, who had it from Clayton in
Virginia; Linnaeus remarks on its similarity to L. Cliffortiana, but
makes it a distinct species. — Stem erect, usually with many racemose
axillary branches in age, sometimes becoming sub-corymbiform
through elongation of the lower branches, 15-100 cm. high (ave.
30-60). (In sterile soil dwarf plants 10-20 cm. high are often seen;
usually unbranched, bearing a few flowers and apparently maturing
seed). Lower part of the stem usually purplish (sometimes green),

Fig. 20. Range of Lobelia inflata.

upper part light green. Whole plant loosely long-hirsute (rarely
nearly smooth), or the upper stem and upper sides of the leaves
smooth or strigose. Hairs flat, chaffy, most numerous near the angles
formed on the stem by the decurrent leaf-bases. Cauline leaves 10-25,
obtuse and obovate or broad-ovate below; above ovate-oblong or
ovate-lanceolate, short-acute; sessile or sub-petiolate below; irregu-
larly rough-serrate or dentate; usually 1.5-2.5 X 4.5-8.0 cm. (some-
times larger). Upper leaves often passing gradually into the broad-
leafy lower flower-bracts. Inflorescence consisting of loose racemes at
the ends of the branches, the central one the largest, to 30 cm. long
(usually 10-15 cm.), not secund, bearing 1-30 flowers upon slender,
more or less erect, finely prickly-puberulent pedicels (5-10 mm. long

• ^

\u

i

324

Rhodora

[September

Ij

m fruit), each with a pair of inconspicuous bracteoles at the base
1^ lower-bracts leafy-ovate below, lanceolate or linear above, little
longer than the pedicel, finely callose-denticulate, smooth or ciliate
Calyx in anthesis campanulate, smooth, becoming much inflated in
fruit, oval to sub-globose, 3.5-8.0 X 7.0-11.5 mm. Capsule inferior.
Calyx-lobes subulate or linear, 3.5-5.0 (8.0) mm. long, smooth or
rarely slightly ciliate; auricles none. Flower inconspicuous, 8-10 mm
long, including calyx. (In this species the corolla is quite short in
proportion to the calyx, being only about 7 mm. long.) Corolla
violet-blue to nearly white, sometimes with a suggestion of pink which
shows plainly m dried material; base of the lower lip pubescent-
corolla otherwise smooth. Corolla-tube entire, except for the dorsal
lissure; lobes of the lower lip oblong, shorter than the tube; the two
upper lanceolate. Filament-tube 2.5-3.0 mm. long, slightly pubescent
below, united % of its length above. Anther-tube 1.5-1.7 mm. long
bluish-gray, the two smaller anthers tufted at the tip, the three larger
merely pubescent on the backs. Annual.

This species is readily identified in flower by the campanulate calyx
rather long calyx-lobes, and the short corolla, which seems even
shorter in proportion to the large calyx. Flowering specimens are
sometimes mistaken for L. spicata, as the branches often do not
develop until frmt has appeared on the main axis. Fruit matures
quickly, so that a single plant is in flower during a long period and
still has only a few flowers open at a time, instead of the long flower
spike of L. spicata.— Dry woods, fields, roadsides and waste places-
an aggressive, weedy species. Prince Edward Island to Hudson Bay
(Hooker, Fl. Bor. Am.) and Saskatchewan (Hooker, 1. c), south to
Georgia; west to the Mississippi Valley (Nebraska, ace. to Petersen).
.S . M* «S5'''^^i'^^ dry, rather open woods; now spreading
rapidly to old fields. Rare or absent on the Coastal Plain. Flower
beginning about July 1 ; flower and fruit through the summer The
speaes is so readily identified that the specimens examined are not

Ih L. Canbyi Gray, Man. Bot. Ed. 5.: 284. 1867. Type Local-
ity: pme barrens of New Jersey, especially at Quaker Bridge . .
(also South Carolina)." Type Specimen : Material collected by W M
Canby and by C E. Smith at Quaker Bridge, as well as M. A. Curtis's
collections from South Carolina, seen in the Gray Herbarium —L Nut-
taUn, m part, of early American authors.— Stem erect, tall and slender,
unbranched or with few-several short racemose branches (sometimes
much racemosely branched), 30-100 cm. high (ave. 60-70 cm)
smooth or sparsely pubescent and reddish below, becoming smooth
and deep green above, leafy. Leaves cauline, 20^0, linear or nar-
rowly lanceolate, 005-0.4 X O9-5.0 cm., often closely appressed
giving the plant a very slender appearance; rather thin, nearly smooth'
obscurely callose-denticulate, but sub-entire in outline; the uppe;
often merging gradually into the bracts of the raceme. Roots fibrous-

T936] McVaugh, — North American Species of Lobelia

325

annual, ace. to Canby, in a letter to Dr. Britton, now in the N. Y.
Botanical Garden. Inflorescence a loose terminal raceme 10-30 cm.
long, never prominently secund, (10) 15-20 (30)-flowered. Branches,
if present, bearing 2-10 flowers each, rarely as many as the central
raceme. Pedicels somewhat angular, 7-11 mm. long in fruit, more or
less upright, usually distinctly upwardly barbed-ciliate, each with
a pair of inconspicuous bracteoles near the base. Flower-bracts
linear, :.bout as long as the pedicels, or longer, to 10-20 mm.; smooth
or ciliate, callose-denticulate. Calyx in anthesis long-campanulate,
rough-puberulent, becoming oval or oblong-oval in fruit, 2.5-4.0 X
4.0-7.0 mm. Capsule more or less upright, about J^ inferior. Calyx-

FiG. 21. Range of Lobelia Canbyi.

lobes narrowly lance-linear, acute, 2.5-6.0 mm. long, obscurely callose-
denticulate, smooth or somewhat ciliate. Auricles none. Flower
9-14 mm. long, including calyx (ave. 11-12 mm.). Corolla blue,
smooth, except for the pubescent base of the lower lip. Corolla-tube
entire except for the dorsal fissure; lobes of the lower lip ovate, slightly
shorter than the tube; two upper lobes lanceolate, about the same
length. Filament-tube about 3.5 mm. long (3-4 mm.), nearly smooth,
connate more than half its length above. Anther-tube 1.9-2.1 mm.
long, light bluish-gray, the two smaller anthers tufted, the three
larger nearly smooth or pubescent on the backs.— Open grassy or
sandy swamps, or pineland swamps; Tennessee and Georgia to the
Pine Barrens of New Jersey; Appalachian Provinces or Coastal Plain,
but in its north-eastern extension confined to the Pine Barrens.
Flower August-September (sometimes in July). Representative
material seen: Tennessee: coffee: Tullahoma, Svenson 4^45 (ANS).

V

326

Rhodora

[September

f

Georgia: Chatham: "Savanna, Aug. 17, 1908" (NC). South
Carolina: Orangeburg: Eutawville, Eggleston 6000 (CM, M, NB
US) North Carolina: Brunswick: Southport, Oosting 33705
(Duke). COLUMBUS: Whiteville, Schallert (Duke); harnett- Erwin
Oosting 33613a (Duke), iredell: Statesville, Hyams (M). john-
^7J^^'' ^}^y^^^' Blomquist 5026 (Duke), new hanover: Wilmington
J^cCarthy^ ann. 1885 (US), pender: Burgaw, Hyams, Aug. 1879
(Ub). Delaware: Sussex: Ellendale, Canhy (ANS NB US)
New Jersey: Atlantic: Mays Landing, Pennell 8113 (ANS) ' Bur-
lington: Speedwell, Stone 7U0 (ANS). camden: Cedar Brook
^o^Ehvee, Aug. 1893 (ANS). cape may: Belle Plain, Stone 3A8A
(AJNbj. ocean: Lakehurst, Long, Aug. 1908 (ANS)
12. L. BoYKiNii Torrey & Gray, A.DC. Prodr.'Syst. Veg. VII-

374. 1839. TypeLo^
cality: "in paludibus
Georgiae (Boykin) et
Floridae (Chapman)."
Type Specimen: au-
thentic material, col-
lected by Boykin, in the
Torrey herbarium in the
New York Botanical
Garden. Aquatic, the
lower stem immersed.
Stem erect, slender,
simple or with spread-
ing racemose branches
above, smooth, green,
fistulose, 50-85 cm.
high. Leaves cauline,
filiform, smooth, 0.52-
2.5 cm. long, few-50 or
more, often deciduous,

Fig. 22. Range of Lobelia Botkinii.

or^+.v^ ^« ^k 1 11 1 . . more, oiten deciduous,

fntothphr^.'"'?fu'-"fl^^^ *^" "PP^^ '"^^gi^g graduali;

^f n^l.« T fl ^ ""^ *^^ inflorescence. Spreading by thick creeping root-
stocks. Inflorescence a ax terminal raceme, more or less secund, 7-20
n^diip?«f;^iT"^ 10-25 oosely spreading flowers upon slender smooth
pedicels 6-17 mm. long (ave. 10-11 mm.). Bracteoles of pedicel none
Flower-bract^ smooth, filiform, much shorter than the pedicels, 2-8 mm'
long Calyx in anthesis very small, round, or appearing flattish because

about' fo"^"^- '^^r-^^^^^'' ^r^*^' *^^^^"^^"g henlspheric inTuTt!
about 3 0 mm. in diameter. Capsule about half inferior, somewhat
longer than broad Ca yx-lobes spreading, filiform, 3.(M.5 mm.Tong
entire, smooth. Auricles none. Flower 10-13 mm. long, including
calyx. Corolla blue, with a white eye, smooth, or ciliate nsTde Sf
lower hp smooth or ciliate. Corolla-tube entire, except for thVdoi^al
fissure; lobes of the lower lip oblong, short-acute, shorter than The

1936] McVaugh, — North American Species of Lobelia 327

tube; two upper lobes long-linear, nearly as long as the tube or shorter,
erect. Filament-tube 3.5-5.0 mm. long, smooth, connate above about
% of its length; deflexed. Anther-tube 1.5-1.8 mm. long, bluish-gray,
the two smaller anthers sparsely white-tufted at the tips, the three
larger pubescent on the backs.— Pineland swamps or cypress ponds,
often partially immersed; northern Florida, Georgia, South Carolina,
southern Delaware; Coastal Plain. Flower May 1-June 15. Repre-
sentative material seen: Florida: gadsden: Quincy, Chapman (NB).
Georgia: Baldwin: Milledgeville, Boykin (ANS, NB). berrien:
Alapaha, Curtiss 6819 (Del, M, NB). coffee: Douglas, Harper 2199
(NB US), lee: Leesburg, Earle (Tracy 9118) (NB). telfair:
Lumber City, Biltmore herb. 4167a (M, US). South Carolina:
SUMTER : Cane Savanna, Stone 4^0 (ANS). Delaware: Sussex:
Ellendale, Long and Bartram 1636 (ANS).

13 L. appendiculata A. DeCandoUe, Prodr. Syst. Veg. Vll: 376.
1839 Type Locality: "Texas." Type Specimen: collected by
Drummond; seen by DeCandoUe in Bentham's herbarium. Speci-
mens collected by Drummond in Texas, now in the Gray Herbarium,
are from Hooker, and are probably authentic— In appearance and
vegetative characters not to be distinguished from L. Gattmgen Gray
except for the larger size (25-90 cm. high; ave. about 50 cm ); more
numerous and larger leaves (4-15 in number, ave. 8-9) ; size 1-3 X 2-7
cm Inflorescence a terminal raceme, usually distinctly secund,
about 20-30 cm. long, bearing 15-70 (ave. about 30) flowers, upon
short curved pedicels (4-11 mm. long in fruit), which are rough-
puberulent, each with a pair of inconspicuous bracteoles near the
base. Flower-bracts ciliate or nearly smooth, callose-denticulate,
linear or narrowly lanceolate, 4-10 (18) mm. long. Calyx in anthesis
short-campanulate or flatter, nearly smooth or puberulent, becoming
long-campanulate in fruit, with a smooth inflated appearance, 4-6 nam.
in diameter. Capsule Vs-H or more inferior, horizontal or somewhat
pendent at maturity. Calyx-lobes narrowly ance-inear, 4-8 mm.
long (ave. about 5.0 mm.), densely bristly-ciliate (sometimes only
near the tip) ; auricles conspicuous (rarely small or lacking), lanceolate-
acute or broad-foliaceous, 1-3 mm. long, often white-scarious tipped;
often connate; usually bristly-ciliate on the margins Flower 10-15
mm. long (ave. 12 mm.), including calyx. Corolla light violet-blue
or lilac, smooth outside, pubescent inside at the ba^e of the lower lip.
Corolla-tube entire, except for the dorsal fissure; lobes of the lower lip
broad-ovate, about as long as the tube; two upper lobes shorter,
lanceolate. Filament-tube 3-4 mm. long, pubescent below, connate
about half its length above, somewhat deflexed. Anther-tube about
2.0 mm. long, bluish-gray, the two smaller anthers tufted at the tips,
the three larger pubescent on the backs.

This plant cannot surely be separated, except by geographical
range, from L. Gattmgen Gray. Apparent differences disappear when
various series of measurements are averaged; L. appendiculata is a

?'■

328

Rhodora

[Septbmbeb

' <

/ 4

latter, and th/fact that'Xivt^, few inte™^^^^^^ It^e

seen, it seems best for the present tom^inf^lfu'''*"*^ ^"''^ ''««"

Fig. 23. Range of Lobelia appendiculata and (insert) of L. Gaitinoeri.

Alabama: Dallas: Marion Jet., Mohr Mav l«Qq /^A1^Jc^
Uniontown, Mohr Jun ISOn rM^( t ^ "^ (ANS). perry:
Pmn^Z/ 10m7mv\ L¥^V Louisiana: acadia: Crowley

HARRiq W^i.of^r, r- ^71 • "^f^^N. J^leteher, Palmer 957 A (Mo)

Hx:oJf°aaft't?r;v^s:.7^or^^ '"""' '"" (^« •

Augustine, Pa/,«er OSnmo) Ziv.^^Uf^'^A ^^^^T'^^V^""
US). UPSHUR: Big Sandy Ti.reK5i^rM'r' ^'""^ ^^^ ^^*''
Lake, Reverchcm lo«/(Mo) Sk^hom! -^^.l ''^t'^ '^''''^^ S''^^"-
Butler, May 1877 (ANs! ni USh'tXr^A^J/T ^"^(0^^^:

19361 Femald, — Rydberg's Flora of the Roeky Mountains 329

(probably) "Fort Cobb to Fort Arbuekle," Palmer, ann. 1868 (NB,
US). CHOCTAW: (probably) "Fort Towson, Ark./* Dr. Edwards (G).
COMANCHE: Caehe, Stevens 1316 (G, M); Fort Sill, Mrs. Clemens 11803
(Mo, R). LEFLORE: Page, B/oA:% 1405 (G, M, Mo, US), pittsburg:
McAlester, Pennell 10601 (UP). Arkansas: benton: Plank, ann.
1899 (CM, NB). miller: Texarkana, Eggert, Jun. 1898 (Mo).
NEVADA: Prescott, Hollister 20 (US), ouachita: Camden, Fendler,
Jun. 1850 (G). pope: Russell ville, Pennell 10623 (UP), prairie:
Hazen, Palmer 26054 (Mo), pulaski: Little Rock, Engelmann 4^3
(Mo). SEBASTIAN: Fort Smith, Bigelow, ann. 1853-4 (US). Wash-
ington: Fayette ville, Palmer 8186 (Mo, NB). Kansas: Cherokee:
Melrose, Rydberg and Imler 2j2 (NB). miami: Paola, Oyster, Jy. 1883
(US); this specimen is on a mixed sheet: the locality may be doubtful.
Missouri: jasper: Asbury, Palmer 34664 (ANS); this specimen is
hardly typical ; the characters are close to those of L. spicata. Illinois :
HANCOCK: Augusta, S. B. Mead, Jun. 1859 (Short herb., ANS); this
is a mixed sheet; two plants are surely L. appendiculata, while the
third is close to L. spicata var. hirtella.

{To he continued)

f«^

346

Rhodora

(October

i ,

' X

f

I *

STUDIES IN THE TAXONOMY AND DISTRIBUTION OF
THE EASTERN NORTH AMERICAN SPECIES

OF LOBELIA

Rogers McVaugh

(Continued from page 329)

l««9' V^^V'NGERi Gray, Proc. Am. Acad. Arts Sci XVII • 991

Gray Herbarium. Stem erecriZ-M cm Lw ^^ '^^'"^'"^M; in
unbranched, or often wif hi Ififiiv high (ave. about 30 cm.),

each bearing rfewflowI«ei;fn*iV:vh"''K«''* '^'^''^'^ ''^''"'^''^^
the base; newly sIootHxce^?'! W t '^T"'?^""^^ *'"Se near

clasp^gf otat: tnirn/ wi^;"^ -ooth sessile or even somewhat
upper fhort-tute, irSkMrr^lttnT. fi ' f^^\'^^^?- »' the
2.0 X 4.5 cm., the upCtolfsSaLr i^h^ ^ ^^^"^^ ? ^'^^ *°
the bracts of the inflo?escen«. nZl I ' ' v ^" ^''"'P' ''''*"6« *<>
obtuse, nearly smooth narr^edaTfl^T"' '' T'"",* ^-^' "'"'^"te-
(1901). Inflorescence 'a teSl rLeme T27^""T'' T ^"^ ^^'^^
15 cm.), usually distincdv ,lr.„n? t '• L'^™- '°"8 (*^^- about

flowers upon shoVcurvedLSwA'*""^^^^^^ i"^^- ""'""t 30)
nearly smooth eac^wkh .Tn.^ V^ °'°?- '°"6 >n fruit . which are
base. Plower-bracto Tmooth^ .• ° '"«'"«P«=r"= bracteoles near the

the pedicel S'mm.'rnfor ^7' clri^Tnth"''^*^' '°"«^' J''""
smooth, becoming long-campLulate tt^J li!^'" '=»"P«''"l«e,
appearance, 3.5-4 5 mm wrot hi ^ J ' ^'^^^ """"tb, inflated

inferior, horizontel ^r somewhat^ntfn ?' Y^' ^?P^"'^ ^ "^ ■"<>'«
narrowly lance-line^ ah™T ^ J 1*°* ** maturity. Calyx-lobes

near the tip; auricles Aone or in ^f-^^' ^"'^*''' °' '"'"'^ ""''^^
Flower 10-13 mm long ?a;e 12 JZ "^ ^•'*'"'=V ^^ort-triangular.
violet-blue or lilac smooth o.-lSr'''' r'"*^'"*^- '^''-'y^- Corolla light
the lower lip C^^^rtMh- .^ ' P^^escent mside at the base of

lobes of the WerViXa'd-^vafTr f'?' '°' *•>« ''°"''' fi««"re

lobes shorter, lanc^r'^rafmSib" 2^3%*'^ *"*?= *"° "^^^^
smooth, united about half its lemrth «hT.r '"""long, neariy
Anther-tube 1.9-2 5 mm lonihM?! I' «°'ne'^hat deflexed.

slightly tufted, the thr^ larL u« ^f.r^' K ''" *^° ^^^''^^ ^nth^"
Springy places of calcar^.., wfff i^ pubescent on the backs.-

from thrL «es ?n X Kn " ''"'""f ^"^ ''"'"^" ""ly
Flower May-June Material ^^T." '^^°" °^ '^"*™' Tennessee

1936]

McVaugh, — North American Species of Lobelia

347

is presumably from LaVergne, but is labelled merely " middle Tenn-
essee" (seen in ANS, G, M). wilson: Lebanon, Biltmore herb., Aug.
1899 (US); Lebanon, Pennell 11384 (ANS); 11391 (ANS, UP).

15. L. FLACCiDiFOLiA Small, Bull. Torr. Bot. Club XXIV: 338.
1897. Type Locality: Ochlockonee River Swamp, Thomas ville,
Thomas Co., Ga. Type Specimen: Small, Jy. 12-22, 1895; in New
York Botanical Garden. Stem slender, erect, simple or with several
filiform branches bearing a few flowers each, 30-90 cm. high, smooth
or sparsely hirsute, green, or reddish below. Leaves cauline, thin,
few-15, smooth, sub-entire, with numerous obscure shallow (often
crenate) callose teeth; in size 0.5-1.5 X 5-11 cm., lanceolate or long-
oblong, short-acute at the tip, mostly rather abruptly narrowed at
base, the lower short-petiolate. Floral bracts definitely smaller than
the leaves; larger leaves well below the inflorescence. Inflorescence a
loose terminal raceme, sometimes plainly secund, few-30 cm. long,
bearing 3-20 flowers (when branched, the branches developing later
than the main inflorescence, with 1-8 flowers each). Pedicels rough,
slender, curved, 4-1 1 mm. long in fruit, each with a pair of conspicuous
green smooth or ciliate bracteoles near the middle or below. Flower-
bracts linear, smooth, denticulate, about equalling the pedicels.
Calyx in anthesis short-campanulate, somewhat rough-puberulent,
becoming hemispheric in fruit, strongly ribbed. Capsule more than
half inferior, longer than wide, 4-6 mm. in diameter. Calyx-lobes
narrowly sagittate, acute or acuminate, ciliate, usually glandular-
toothed, 3-5 (7) mm. long; auricles reflexed, round, small, but con-
spicuous, 1 mm. or less long. Flower 14-19 mm. long (ave. 15-16
mm.), including calyx. Corolla blue, pubescent outside or smooth,
the lower lip somewhat pubescent at the base, with two tubercles.
Corolla-tube entire, except for the dorsal fissure, or fenestrate; lobes
of the lower lip oblong or narrow-ovate, nearly as long as the tube;
two upper lobes lanceolate, erect. Filament-tube 5-6 mm. long,
pubescent below, connate more than half its length above. Anther-
tube 2.0-2.5 mm. long, bluish-gray, the two smaller anthers tufted
at the tips, the three larger pubescent on the backs. — River swamps,
Coastal Plain, Georgia, northern Florida, Alabama. Flower June-
September. Material seen: Georgia: colquitt: Moultrie, Harper
1676 (Mo, NB). JOHNSON: Wrightsville, Harper 1341 (Mo, NB);
(most of these specimens are not to be distinguished from L. Halei
Small). THOMAS: Thomasville, Small, Jy. 1895 (NB). Florida:
"Ad ripas fluv. Ocklockonne," Rugel, Jy. 1843 (NB). okaloosa:
Milligan, Curtiss 6855 (Del, M, Mo, NB, UP). Alabama: mobile:
Spring Hill, Graves 1273 (Mo).

There is some evidence to show that this species and the next are the
same, and intergrade freely (cf. the plant cited above. Harper 1341)-
However, L. flaccidifolia differs by the thinner, more oblong leaves,
greater tendency to branch, smaller and more numerous flowers, and a
somewhat later flowering period; in view of these differences, it seems

IRREGULAR PAGINATION

348

Rhodora

[October

''■i

I t

I' /

kllen. ^'^^ ^^""^ ""^ ''^^'''*' 'P'"^"'' ^* ^"^^* "^tiJ ^«^e "material

16 L. Halei Small, FI. S.E.U.S. 1145. 1903. Type Locality-

wet prairies western Louisiana; Texas, near Houston^? 'tI^^^^^

Specimen: authentic material, collected by J. Hale at Alexandria

La and Identified by Asa Gray as L. Ldovidala seeT^ the

Xn^m 1^^T'~/'tYT'^''^^ ^^^^' ^^^^- ^^' Acad. Arts Sc"
^n. 60. 1877; not L. Ludomciana Wood, Class Book 476 1861 -^

br^rchra'o^r^^^^^ ^^' ^^"^!..^j: -^^^ ^-^ ^^-^ Sht

leaves few ^T^^^^ ^T' ""' '"^^^'^ '^"^^ *^^ ^^^^^ Pauline

n?J2; Ki ?*^' somewhat appressed to the stem, the lower
petiolate; oblanceolate, short-acute below, about 1.0 X 5-8 cm. 7 be-

coming lance-
acute above;
subentire or
sharply toothed.
Upper leaves
bract-like, dis-
tant; the larger
leaves well be-
low the inflo-
rescence. Inflo-
rescence a termi-
nal raceme to
35 cm. long, not
noticeably se-
cund, loosely 5-
30 flowered.
Pedicels rough,
curved, 5-8 mm.
long in fruit,
each with a pair

1936]

McVaugh, — North American Species of Lobelia

349

/r^'^'j^^'^T^^*® °^ Lobelia Halei (unframed) anH

long as the pedicels. Calyx in anthesis short-campanulate or hpm;
nte' rtT'''™'."="*' '^«'™"« hemispheric ZlvTt stronSv
wWe Ca^Io^viT- '." '^'''^''''- C''P="lehalf inferio™ ongSn

round, conspicuous, less than 1 mm. long. FWct 15-20 V22?l^

mm. long, pubescent below, connate above A^ther-luZ 2 1^^ *

Sp''ttthr"'t^''\*''''f"° ™'"'- -tht densely whttt^T,^
at tip, the three larger heavily pubescent on the backs.-WeJ prS

■^

v-iy

Sk'

"^y

r3^

f

-^

W^

usually in sandy soil. Coastal Plain, Louisiana and eastern Texas;
doubtfully north to Kansas. Flower April-July. Material seen:
Louisiana: "Cigers Point, W. La.," Langlois, May 1886 (NB).
CALCASIEU: Sulphur, Palmer 7722 (NB); DeQuincy, Pennell 10226
(UP). RAPIDES: Alexandria, J. Hale (ANS, G, NB). Texas: F.
Lindheimer, Fasc. I. 116., ann. 1843 (ANS). Harris: Houston,
Lindheimer 41 (Mo), liberty; Tharp 2506 (US), orange: Orange,
Tharp 2733 (US). Kansas: labette: Parsons, Letterman, Jy. 1880
(Mo); this specimen is apparently L. Halei, but there may be' some
confusion as to locality.

17. L. FLORiDANA Chapman, Bot. Gaz. Ill: 9. Feb. 1878. Type
Locality : " Margins of ponds and swamps in the pine forests of West
Florida." Type Specimen: material from Chapman seen in the
New York Botanical Garden and the United States National Herb-
arium.— L. palu-
dosa var. flori-
dana Gray, Syn.
Fl. II. pt. I:
Suppl.394. 1886.
— Stem erect,
slender, or rather

coarse in large
specimens, un-
branched (or
often with several
stout upright
or spreading
branches, de-
veloping later
than the main
axis, and bearing
fewer flowers);
smooth, 50-150

J

Fig. 25. Range of Lobelia floridana.

cm. high (ave. 80-100 cm.), green or stramineous, less often purplish
at base. Leaves basal, 1-10, smooth, strap-shaped, oblanceolate or
lanceolate, acute or obtuse at tip, long drawn-out at base into margined
petioles; to 2.5 X 40.0 cm. (ave. about 1 X 25 cm.); more or less up-
right ; entire or shallowly crenate, or dentate with callose teeth. Stem-
leaves bract-like, 3-4, lanceolate, 2-3 cm. long, acute, distant, callose-
denticulate; the lowest sometimes larger, to 0.5 X 8.0 cm. Inflorescence
a terminal raceme (sometimes with secondary, shorter racemes in
branched individuals), 10-50 cm. long, loosely or rather densely 10-40
flowered (ave. about 25) ; not secund. Pedicels stout, rough, more or
less upright, 3-6 mm. long in fruit, each with a pair of inconspicuous
bracteoles at the base. Flower-bracts smooth, linear, shorter than the
pedicels or equalling them, inconspicuous. Calyx in anthesis flattish or
conic, usually rough under a lens, becoming hemispheric in fruit (some-

i

350

Rhodora

[October

/

t

/

(

«lTt l7-%** •"T-)'."''";'* ''•° """• ■" diameter or larger Capsule

tCdi^tTi ^'^^'' *'"'." ^'°'"^- Calyx-lobes broad^rnceolateor
deltoid 2-6 mm long, smooth, acute, usually callose-denticulate (ottZ

tTlWrnir'-'^rr^^f'*^*."^"'^^- F'owtlllTomilon"
n?,f!M \k I -V T'"'^',''« '^y''- Corolla blue, usually pubescent
outside the lower Ijp densely hirsute-pubescent at base inside Sla
tube entire except for the dorsal fissure (rarely fenestoVtT) lohe,Tllf»

h^rb' Km '^'''''°~-' ^n*"'"y ^'''■)= ^*«P^«. Aug 1^ (xt^;
nerb., rMij;. liberty: Biltmore herb Ana loni WTc^ VAorrey

Milton, /Taroer i5 CO ^TK ifl\ ^' "^ IF^^- ^^^^a rosa:

TAMMANY: Covington, £ro. ^r*ene //S«5 (NB US)

LocAmv-'?nT ^"**t"' ^'"- ^- ^'"- P'- "^ '75- 1818. Type

Warrlto GeoS-n^rs^BaMr'/r .'"^^^^ -""^^ '"
Academy of Natural Sr.Vn/.^ „rSl". j'."^*''*^''*"' material in the

Sk.Bot S.C.&Ga I.2^r,l2. r ^t'P^'^- A ^'^'**' Elliott.

usually oblanceolate, short-acite frarelv v^V^ . . Jf- P '^'"•)'

short petiole). Stem-bracte I 0-2 l^Z^^i , " ?^^^ '''*'^« *nd a
rescence 2-35 cm Wgh ?ave at™,^ ^^ "^^ '"'"^ f" '='"•)■ I""""
(ave. about 15). ^^dUs routh Lt^fS^' '^l "^'^^ ""'""'"'^

, oout d.u mm. long. Auricles none or very small. Flower

1936] McVaugh,— North American Species of Lobelia 351

11-16 mm. long, including calyx (ave. 12-13 mm.). Corolla light
blue or nearly white, practically smooth outside the lower lip densely
hirsute-pubescent at base inside. Corolla-tube fenestrate ; lobes of the
lower lip ovate, Httle shorter than the tube, scarcely reflexed; two
upper lobes lanceolate. Filament-tube 3.0-4.5 mm long (ave. 3.5
mm ) hairy below, connate about half its length above, somewhat
deflexed Anther-tibe 2-3 mm. long, light bluish-gray, the two smaller
anthers tufted at tips, the three larger merely pubescent on the backs
ThTs sS seems wholly distinct from the larger L. flondana. It
m^Tv be distinguished by the shorter filament-tube, general y smal er
r/and sSr corolla; the fenestrate corolla; the absence of bracteo es
orthfpXel -S and low pinelands and ponds, often partially
l^mersS so^^^^^ Georgia and throughout peninsular Florida;
reported (perhaps always
upon the authority of Nut-
tall) from Delaware; the
report may be based upon
L. Boyhinii T. & G., which
occurs there. Flower Feb-
ruary-May, or more or less
throughout the year. Rep-
resentative material seen:
Georgia: camden: St.
Marys, " Bal." Schweinitz
herb. (ANS). charlton:
Traders Hill, Smally Jun.
1895 (NB). Florida: ALA-
CHUA: Gainesville, Craw-
ford, Apr. 1897 (ANS,
Duke). BREVARD : "Indian
River," Palmer, ann. 1874
(US). BROWARD: Ft. Lau-

FiG. 26. Range of Lobelia paludoba.

(US). Broward: rti.au- Hibernia, Canby, Mar.

?r.o''rANrNB' UST IZ^ms^i^frnaTsm NB). duval:
1869 (ANS, JNrJ, Uo;. *^^"*", ^tr^ xTr» xtvc ttq^ wbanklin*
Jacksonville, Curtus 4715 Del, M ^B, N^, US), franklw

Apalachicola, Chapnu^n Biltmore herb 2679MNB ^S • t^^^,
Wewahitchka, Leeds, Apr. 19^^ (,AIN&;. uii^i^o^ivw r

GaXr May 1876 (NB, US). Indian «^^^i,^f^'^^.^"'^Jft,

(NB). P,--E"f'i^7*lffiB''usrSo;y<^^w

Mvers. Miss Standley 4O (AMb, i^m, i>D, ^^J- \ /vrm tfvy-

"in^ter'Tallah^see yt. MarW' i^^^^^^^^^

Bronson, J. D. t^miih, Apr. ^^^V^^. ^* ^^ PraiVip Mrarns Apr.

T,^mf (NB, US). "<;~"ijsart?S' sSito "

352

Rhodora

[October

'/

/

ft
U

I

May \^n (Del NB) (Octobei

VOLUSIA; Lake Helen Mr^^n^'^^' ^^^^^^ota, Leeds, Apr 19Si M vc^

Locality: "KeiT'^'^"^*-'^^«t-W 1«tq t^

the Academy of Natural Sdences of P^^
Rayunhum minimum Rore ««//,T ; ^^^'^^elphia.— Perhaos 1 1^^

species / li , .; ''°'- ^ep. V: 340. ISn-i „,u::u'- i^^^- not
P«"es. i. Kalmu, var. j.<j„fo. Barton FIn/ """r^'^ ^"^'^

r— p -, ' • ^^' ^^- I: plate 34

' ^' ^ ^ iS^l.^Stem

slender, erect,
20-75 cm. high,
sometimes sim-
ple, but usually
with one-several
filiform race-
mose upright
or spreading
branches •
smooth and
green above,
usually dark
purplish-red and
short-puberu-
lent below. Cau-
line leaves few-
20, smooth
rather thin, ob-

^10.27. Range of LoBE^,^ j^^^^^^

olate. „ore orVssU s^/,^" S'"x\ ?'^''' '-'- o^ate! S-
'ong. Inflorescencp a ♦^ ^V'^^'^^ ^-^ X 1.5 cm P*»fmi^ * '^^ pen-

flattish; often ^?rf^''''^™^^'^^tdenticula?rrif- ^"^ '^'^'

iiuwer 8-11 nmj

1936] McVaugh,— North American Species of Lobelia 353

loDK, including calyx. Corolla blue, with a white eye and two greenish
tul^rcles at the base of the lower lip; smooth, or hairy inside the tube.
Corolla-tube entire, except for the dorsal fissure; lobes of the lower
lip narrow-ovate, shorter than the tube; two upper lobes lanceolate,
curved upward. Filament-tube about 3.0 mm. long, pubescent below,
connate more than half its length above. Anther-tube about 15 mm.
lone (1.0-1.8 mm.), bluish-gray, the two smaller anthers tufted at the
tips, the three larger smooth or pubescent on the backs. Open sandy
or CTassy swamps, or woods, sometimes in brackish marshes, or dry
sandy places; usually in acid situations; central Tennessee and
Kentucky to Alabama (Mississippi, ace. to Small), Florida north to
southern Long Island. Appalachian Provinces and Coastal Plain;
Tpennsylvania and New York confined to the Coastal Plain and the
region south of the moraine. Flower July 1-September 1, from N. J.
northward; fruit July 2(H)ctober. Southward flfwermg earl^r,
beeinning May 20. Representative material seen: Kentucky.
Sl" Sandy Swamps, Braun, Oct. 1933 (G);MC creary: Pine Knot,
PenneU 13899 (ANS). Tennessee: "'^^■^"n: Wn^, i.- A.
Smith sm-a 1881 (W). morgan: Huffmans, PCTinc«i;S£>37 (ANb, ur;.
^T^': Wnfield, PenneU 1S926 (ANS). Alabama: Baldwin: Perdido
MoL, Jun. 1890 (US) Cherokee: Round Mt.i.e^. Jun^^9^4
(ANS). CULLMAN : Cullman, Eggert, Sep. 1897 (CM, Ub). etciwah.
BaUplay, Mohr, Jy. 1890 (US), •'ackson: Flat Rock, ^^^^"^ J"»-
1933^ANS, UP). JEFFERSON: DeSoto Falls, ««'*l /^ 1898|NB, O,
US). Florida: gadsden: Qumcy, CAoymon (ANS) Walton.
DeFuniak Springs, Curius 590^ (Del, M, NB.NYS,US)^ Georgia
BALDWIN: "Dr. Boykin. Ga.," Torrey herb. (NB). ^.^^^^J-p^'*/.
mont i/ar»er «/4 (NB, US). IRWIN: Ocilla, fforycr ^/S (NB,IJ!5;.
S^RTSnC^I. 'LecJnte (NB). Richmond: Au^^^-/"'*^^^
Tun 1900 (NB) South Carolina: aiken: Aiken, /tojimet, Aug. 1800
(NB) andeSon: Anderson, Davis 82H (US), darlington: Harts-

%'^^t^^S.J''^^Hoi.%li (US).' -kens: TaUe Mt
^mnll Aufi 1896 (NB). RICHLAND: Columbia, fay/or Jun. 189UM;.
f^MTER Cane Savanna, Stone m (ANS). North Carolina:
SUMTER. v.ane ^"'^»:" ' ^q TANS) Brunswick: Southport,

ANSON : Wadesboro, Leeds, Jy. 19^^ ^ „ ,Vu^.^ Wh^rr^t S^n 1934
Bhmquist6027 (Duke), co^^mbus: HaUsboro T^^err^^^^
rTTPV^ CRAVEN- New Bern, Loomis and Croom (ANS). J^^^^^^^^^'
FavetteX^ito^^^^ 624k (NB). Durham: BlomquiM 5029

fffl H^^^^ ^^9er (NB). Henderson : Hender.

Se, B^re624a (NB, US) HYDE:^ranton ^^^^^^^ Jun^^S^S
(NC). iredell: StatesviUe, Hyairw (M) ^^^ff.^' .^^^^^
Pmnell U178 (ANS). Johnston: Clayton, Blomquist 5026{Du)ii^.
i'mnea i4yo v^^^-^;- ,^^. moore: Pinehurst,

macon: Highlands, Cuthbert, Jy. l»y/ '^ VxrM^:„„t^n Comlle 71
Wicker, Sep. 1931 (NC). new hanover: Wihnington ^^^^^J^^
(US), orange: Chapel Hill, Totten. Jun. 1915 (NC). pender.

i

\f

\

i'i

354

Rhodora

[October

(ANS). TRANsrLv/Nu.:^S« K„tam^^i"»<^ t"^^ Aug. 1891
Virginia: AccoMAC- Franklin r.-tvR' f'^^'^' Sep. 1934 (UP).

BETH ciTV: HamptoSrZ aS S"Vr^- '""' ^^^S^" ^"^^-
Lecmard and KUliv 562 (KKi TTcf ^ °''- Gloucester: Ark

Jun. 1893 (ANS UP T qwF ^" ^"^^''^''■''^•■"Belfield," iyX

^eamej/ 1676 (US). Norfolk r-ff^ ^ t ^- n^nsemond: Suffolk.
GEORGE: New Bohemia? K«S fAM«^^^2 ^N^^' ""'"CE
Virginia Beach, £„•«„„ , oTX 1892^NbTO- ""'"^^^^ ^•'~^=
Magnolia, Carter, Aug 1904 avl? ^ ^- Maryland: harford:
Otis (?) C/S^ff herb RRT) !^;.'"''°'''*'°i Sharptown, ^. P
Jy. 1893 (Del). DElAWA„E7"NEwrA^Tr"=A°'T ^''y- ^»*S'

fV^if- Z^^'^'^y- ^^«^WA W TaNS^ ^^'i" Pennsylvania:
Island, ^. .ff. 5mi<A, Jy. 1864 (UP) NpwT "^^^^^RE-- Tinieum
If nding, PenneU 8117 (ANS »„ivf ^^'^'^y^ Atlantic: Mays

(ANS). CAMDEN: Lindenwol^-i4^S7A^.'t*''°"' ^' ''''
Spring, Stme Ism (ANS) r^Lt, ^^^.^^' "ape may: Cold
^W (ANS). GLOUCESTER ^karZeX?-' ^'/i*"^ ^reek. Long
cer: Bear Swamp, 5„rt,„:^; ^ ml^AN^l^^^-^ ^^^^^- «^«-
J»faoi«„«e .?*0S (US). MONMOUTH F««L;^;i "T'-'^^E'^' Milltown,
ocean: Lakehurst, WS rANsf ^'''''^' ^^ ^'^^'^ (ANS)
Aug. 1927 (herb. R.^) NeVS ^'"''^''^ i"*""™' ^«««4
Jy- 1916 (NYS). queens: Wence St. T\''= Oceanside, House
SUFFOLK: Central Islip, /'.r^o:rjy 1920 (NYlf ^"^^ '^"^ ^^B).
t'p E L^'L^r-E^;^^' ^"''. /■" ^'^- Sci. XII. 60 1877
Material dte7by trarVf) te^'ln'th'^''"" J^"' « "— J
L.aphylla Nuttall, Am. Jou^ Sd V 297 ^';Y,,"^'^?™"'-Not

ioirfra; see Nuttall in Jour. Acad Phila Vri J^B ''^"'^ '^ ""t »
f • ™?f»PMa Rafinesque Ati^jmrr f /i;^^";!!- ^^34. Perhaps

described as follows :"S?em simple smooth . ' ^^^- ^''^ '''"^^^
sessile dentate, flowers termnal Wj' ^*T,** """"*« ^mote ovate

ana. '-Stem ^eak, sfendTdecuX " or^'T''''- /'°"'^'' ""-^ ^o"'^-
simple or diffusely branched Trai? «'*nding, 5-30 cm. long

purph-sh below, smooth? itrefL"iirn''.*'''H'''^^'-^^"- '"^^
Cauline leaves 1-10, smooth, laSate '^ U "^ '~*'".« "* """le^-
denticulate, about 2.5 X 8 0 mm t!. '*"<=«-o^ate above, acute,
orbicular, 8-13 mm. in diameter with aTfi'^^'" broad-ovate o;
2.0 cm. long; entire or crenai" tootled "^^^"'^ P^t'^'e sometimes
ower cauline ones. Roots?^k slender ' *r r "' 'T^' ''"'"''' to the
terminal raceme 2-18 cm ZiVrnf* V ?,"''?«• . Inflorescence a lax
plant) more or less secund, bef rit o 1 ^^^ i""^*'' °' '^' ^"^^^
smooth slender pedicels wh ch are il'^f ™ ''^' ^T"' ^'''^^'^ "Pon
P- Of inconspicuous ^^.0^:^^^^,^^,^,^^^:^'^^^^^

1936]

McVaugh, — North American Species of Lobelia

355

inconspicuous, 1-3 mm. long, acute. Calyx in anthesis conic or
short-campanulate, smooth, becoming turbinate in fruit, usually
acute at the base, averaging 2.5 mm. in diameter, by about 3.5 mm.
high. Capsule % or more inferior. Calyx-lobes narrowly lanceolate,
smooth, entire, 2.0 mm. long. Auricles none. Flower 7-10 mm. long,
including calyx. Corolla blue, with a white eye, and two greenish
tubercles at the base of the lower lip; smooth, or the tube hairy inside.
Corolla-tube entire, except for the dorsal fissure; lobes of the lower
lip narrow-ovate, shorter than the tube; two upper lobes lanceolate,
curved upward. Filament-tube about 3.0 mm. long, nearly smooth,
connate more than half its length above, deflexed. Anther-tube 1.0-
1.5 mm. long, bluish-gray, the two smaller anthers tufted at the tips,
the three larger smooth or pubescent on the backs. Seeds rough-
reticulate, ovate, about 0.5 mm. long.

This species is very close to L. Nuttalli, from which it differs by the
weaker shorter stems, usually roundish
lower leaves, general smoothness, includ-
ing pedicels and calyx (in contrast to L.
Nuttalli, which is often prickly), and the
more elongate calyx. It is apparently not
at all closely related to L. Cliffortiana L.
and its relatives, as has been supposed by
most American authors, but is in a wholly
different section of the genus. It is, how-
ever, superficially like the plant passing
for L. Xalapmsis HBK. (L. Cliffortiana
var. Xdapensis Gray), which may be dis-
tinguished by the smooth shining seeds,
more upright, coarser leafy stems, longer Fig. 28. Range of Lobelia
inflorescence, serrate leaves. — Moist Featana.
places, ditches, seashores, swamps, often

in sandy places and pinelands; peninsular Florida; not seen from
extreme southern Florida. Flower January-June, or more or less
throughout the year. Representative material seen: Florida: bre-
vard: Cape Canaveral, Curtiss 5831 (Del, M, NB, NYS). charlotte:
Punta Gorda, Leeds, Apr. 1931 (ANS). duval: Pablo, Lighthipe 539
(M, NB, W). glades: Palmdale, Leeds, Apr. 1931 (ANS). highlands:
Sebring, Mar. 1935 (UP). Hillsborough: Tampa, Churchill, Mar.
1923 (ANS). INDIAN river :Fellsmere,Sma//^^Pi(NB). lake: Lake-
land, Blanton 6971 (NB). lee: Coconut, Moldenke 695 (Duke, NB,
UP), manatee: Bradentown, Tracy 75/0 (CM, M, UP, W). pinellas:
St. Petersburg, Mrs. Beam 5000 (M). st. lucie: Fort Pierce, Tatnall
864 (ANS). SARASOTA: Osprey, B. //. Smith, Mar. 1904 (ANS, CM).
SEMINOLE: Sanford, S.R., Apr. 1915 (NB). volusia: Mosquito Inlet,

Curtiss 1641* (CM, M, NB, UP).

21. L.KALMII Linnaeus, Spec. PI. 11:930. 1753. Type Locality:
"Habitat in Canada." Type Specimen: in the Linnaean herbanum

"1

356

Rhodora

iv

I'

•'>

(October

1^'- TY^K\otf^r"Zck?M^''™*n'4 F'- Montana 378.-

ME.: Teton RrS; sS S^n fn^NS D^'^ofr
Humcane Hills, Assiniboia, J/ocoan ann ISS^ tk ' ^ ' °^.= *'="
to differ from the typical form h^ f'ht .^^•j.^'?*' ^»"ety is said
base of the capsules Howe ve^ th^it."*! P^^"=*'' ""^ t^e acute

even in the ty^e Zt.Z^il7r^d^;TstZ^.e:'r^''''' ?*'''?'^'
has seen material from Montana o„^ Vu ' ™°'^^^f'^. the present writer

bia which exactly d^cateTeastrrnm»! "^T'r"^' °l ^"t'^'' Colum-
not to recognize the varietv ^*f ^™.'"?t«™'- It >s thus thought best
No. 2: 8. Nov 3 Im -Pifntt "'f "'" i^""^"' ^""- Le«d« Herb,
characters. Stims tall slendir n.!T"°1^^ """I'f^' '° vegetative
high, green or reTdish betw smZh L",'}^^,^'*"'^^ ^^^^ ^^^ '^'"■
base; varying to a diffusein'ranTH f °L ^^k*'J: .P"'i^<=«"t near the

shorter, and^ometimes1o^tSf^m:ittem11^f°"n!'^^^

7 0 .r" m"^ "' r^"' P'""*: ^^^^ broader, Targ^r 0 O^i s' v n 7"
7.0 cm., oblanceo ate or broader pv^n t,. „„ ""^8^^, u.08-0.8 X 0.7-

the lower sometimes MTOwed fnT„ K ! "'T***' "^"^''^ °btuse,

present few, spatulate or obovate „°h '"*'°'*'-- ,^'«^' '^''^^^ '^
pubescent, often purplish abontn^' v ^ f ^' P«V.°'"«' s<"»ewhat
3.5 cm.). RootstockXnL ,Z ?■ ^ l^ """■ (Maximum 0.8 X
a loose terminTraceme soS^ Main inflorescence

flowers upon long slZTro^.Z^TjSs ^8 m'' I ''""', ''K'
each with a nnir r.f ««,. • *^, "*^^'s (^»-i8 mm. long m fruit^

middle. ^tdS IrCust^'iT*'' bracteoles'nearJh^e
racemes shorter than the centmrnn. . If^ appressed. Branch-
bracts linear, smooth, aboretaTne the rH^^r^ Y"' ^'°^«'-
more luxuriant, branched pante) ctlyx iS °' ^""^^ ^'" '^'
panulate, smooth, becomiL ^LJ'^ , i? ^"thesis conic or cam-
fruit, varying with a^r on ^fhi^*'', """'TS' »' sub-globose in
young). Cap'suTmoTtran^TnTeri^^^/i"'"""^ ^"''^^ "''»

&d?n'g1aW;;™nL^r ^^ "^^^^^

white eye, or^someTime^if, wWtl' ~ [" '''"*? '^'*'' "^ conspicuou
the lower lip smooth CoroHa t ,h^°''*> "' '^^ *"'"' hairy inside,
fissure; lobes of the lower l,W»t • "^' ^^^^P* '"' the dorsa

the tube; two up^r ToLs lanceltr' '''^'''"'''""« °' "<*«ding

Fpc looes lanceolate, curved upward. Filament-

/

1936] McVaugh,— North American Species of Lobelia

357

tube 2.5-3.5 mm. long, smooth, connate above more than half its
length. Anther-tube 1.6-1.8 (2.0) mm. long, bluish-gray, the two
smaller anthers tufted, the three larger smooth or pubescent on the
backs. Seeds long-fusiform, acute at both ends, 0.6-0.8 mm. long.

The diversity of form shown by this plant seems to be related to
changes in habitat; it is a plant of calcareous situations, in general,
and the slender, unbranched form is characteristic of grassy or marly
bogs, while the coarser or more luxuriant form is known more from
limy beaches or cliffs. All efforts to separate them by good characters,
independent of habitat, have failed. — Wet meadows and bogs, in
neutral or calcareous situations; shale or limestone beaches or cliffs;

Fio. 29. Range of Lobeua Kalmii.

sometimes in sandy bogs; almost always in wet places. Newfoundland
to western Massachusetts and south to the moraine in Pennsylvania
(also Lancaster Co.); west to Ohio, Minnesota, and Colorado, north
to Hudson Bay, Montana, British Columbia and Great Slave Lake.
Absent from large regions of prevailingly acid soil. Flower July 1-
September 1. Fruit July 20-September 20. So characteristic that
it is unnecessary to cite the abundant material seen, except for the
following outlying stations. Pennsylvania: Lancaster: Dillerville
Swamp, Heller, Sept. 1901 (CCD, CU, W). Ontario: Cochrane:
James Bay, mouth of Albany River, Spreadborough, herb. G. S. Can
62542(0). Mackenzie: sandy muskeag, N. arm. Great Slave Lake,
G. S. Hume, Jy 31, 1920(0). Colorado: larimer: Fort Collins,
Baker, Aug. 1896 (CM).

22. L. DoRTMANNA Linnacus, Spec. PL II: 929. 1753. Type
Locality: "Habitat in Europae frigidissimae lacubus & ripis."

I I

J

i .

358

Rhodora

[October

before rn^P^ V^^T "^^b"*™? i° London; seen by Linnaeus
before 1753. Photograph seen.-Gladiolus lacmtris Dortmanni
CIusius "Curae posteriores," 40. 1611. Leucmum palm^Zre
subcaeruleo, Bauhm "Pinax," 202. 1623. Dortmanna^i2
flonbm,parm pendulis, Rudbeck, Act. upg. 1720 d 97^ 2

FMapo"l27 n'^'^'^A V"''^ bilocuhribv. subuiati; LuinLs,
ri. lapp JJ7. 1737.— Aquatic; smooth throughout. Stem UDriirht
unbranched (rarely with 1-2 branches). 5-100 cm. high Tve 3^35
cm.) usually .mmersed about % of its height (all excep? the Infl^^f
cence); green above water, and green to stramineous below leafless
bearing 0-7 hnear fleshy bracts 1-7 mm. long; stem hollow ' Wes
basal (rarely developing 1-3 cauline leaves 2-t cm. long) ifnea?
bvTo^'n^ '" "V^ber 2-27 (ave. 15-20), 2.O-5.0 (8 0) c'm.ToS
Tnfli ".■ '^"^^' y^^" flattened out; obtuse or short-acute

Inflorescence a lax terminal raceme to 45 cm. long (ave. 1^20 cm )'

Fio. 30. American Range of Lobeua Dobtmanna.

PedTcerinf™it'f,r""'^'/''^.'°°Hy ^'^^ fl°^«'«d (»ve. 5-6).
redicels in fruit 4-13 mm. long (ave. about 7.0 mm ) curved einn

fS \^-7Vf ••^'^'^"S ^"'*^«'' «o that the flower' fs often hori-
zontal, while the fruit is pendent. Bracteoles of pedicel none Flow^
bracts Ob use, fleshy, entire, 2-3 mm. long, with a broad bSe c2;
ITa'^" k°"' • °-' '?n8-tri'"»gular, becoming long^ylindric hand
^^12 mm" ri"" ■' T*"^ "'"^ " lonl-attenuate base, Ke

fr^lt::!:^^^ °'*^" "Pr""« ^'^^ ^^aHest. T^Is tend:; y
IS seen tnroughout the group, but not so striking v as in T DnJ

base. Lobes of the lower li^ long.vatef nl^arl^X^^^^^^^

1936] McVaugh,— North American Species of Lobelia 359

not sharply reflexed; two upper lobes linear, curved upward. Fila-

^n«."i?l^ ] T-u"""^ ^T- ^^^"* ^'^^' pubescent below, connate
most of Its length above. Anther-tube 1.3-1.7 mm. long, dark gray
or black the two smaller anthers heavily tufted, the three larger
densely bearded, especially near the tip. Seeds dark brown, with a
promment square base at one end.-Sandy or gravelly borders of
ponds, usually partly immersed; more rarely in mud or in quiet
streams; Newfoundland and central Ontario to northern Pennsvl-
vania west to northern Minnesota; also in Oregon, Washington and
British Columbia; Slave Lake, Richardson (Hooker, Fl Bor -Am )
Apparently identical with the plant of northwestern Europe. Flower
July 1-September 1. Fruit July 15-September 15. So distinct that
It IS unnecessary to cite the specimens seen, except from the following
outlying localities. West Virginia: A specimen in the Detwiller
herbarium m the Academy of Natural Sciences of Philadelphia is
labelled: nr. Harpers Ferry, Virg. Aug. 14, 1846." The Detwiller
herbarium is mostly from near Mercersburg, Pa., and it is possible
that the above collection is from Pennsylvania or even further north
?A^^^T' ^^^^^^^^^'- Cascade Mts., Dark Lake, Sweetser, Aug. 1926

Explanation op Abbreviations for Herbaria

Mo^^ T«^'''''^'*^'4??^^¥^u^^«?f' ^'^^ ^^bo^' Mich.; ANS = Academy of
Natural Sciences. Philadelphia, Penna.; CCD = Herbarium of Chas. C. Beam,
Bluffton Indiana; CM = Carnegie Museum, Pittsburgh, Penna.; CU =
Cornell University, Ithaca, New York; Del = Delaware Society of Natural
History, Wilmington, Del.; Duke = Duke University, Durham, N. C; G =
Oray Herbarium Harvard University, Cambridge, Mass.; herb. R.R.T. =
Herbarium of R. R. Tatnall, 1100 W. 10th St., Wilmington, Del.; M = Univer-
sity of Minnesota, Minneapolis, Minn.; Miss = Mississippi State College,
xfS^ x?"^^^; ¥^^'' M° = Missouri Botanical Garden, St. Louis, Mo.;
NB = New York Botanical Garden, Bronx Park, N. Y.; NC = University of
North Carolina, Chapel HUl, N. C; NE = New England Botanical Club,
Gray Herbarium, Harvard Univ.; NYS = Herbarium of New York State
Museum^ Albany, N. Y.; O = National Museum of Canada, Ottawa, Ont.;
Pa = Pennsylvania State Herbarium, Harrisburg, Penna.; R = Rocky
Mountain Herbarium, Univ. of Wyoming, Laramie, Wyo.; Toronto =
Toronto University, Toronto, Ont.; UGa = University of Georgia, Athens,
i^*"'- , w University of Pennsylvania, Philadelphia, Penna.; US = U. S.
National Museum, Smithsonian Institution, Washington, D. C, W = Univer-
sity of Wisconsin, Madison, Wis.; WVa = West Virginia University, Morgan-
vOwn, w . V ft.

Explanation of plate 435

Fig. 1, L. Cliffortiana L. (possibly L. xalapensis HBK.); Fio. 2, L.
Feayana Gray; Fig. 3, L. Nuttalli R. & S. (one seed of L. Cliffortiana L.
may be seen near the top of the picture); Fig. 4, L. Gattingeri Gray; Fig. 6,
L. Canbyi Gray: Fig. 6, L. spicata Lam., var. originalis McVaugh; Fig. 7,
L. spicata Lam., var. campanulata McVaugh; Fig. 8, L. puberula Mx •
Fig. 9, L. Kalmii L.; Fig. 10, L. glandulosa Walt.; Fig. U, L. floridana
Chapm.; Fig. 12, L. inflata L.; Fig. 13, L. Dortmanna L; Fig. 14, L.
AMOENA Mx.; Fig. 15, L. siphilitica L.; Fig. 16, L. Cardinaus L.

The seeds photographed were from the following specimens:

*i

'(

)

I

f,

360

Rhodora

[October

Fig. 1, Brooksville, Hernando Co., Fla., Leeds Apr 1931 (ANS); Fig 2
Mrl Dekm 6000 (M); Fig. 3, Wadesboro, Anson Co., ^.C.,Leeds,Jy. 1929
CANS)^ F?G V'ceda; barrens," Jun. 1879, Gatiinger (Mo); Fig 5 EUendale,
i j'rv; nil rnmrnnns Sen 1895 (ANS): Fig. 6, Madalin, Dutchess Co.,
NY McVS 267r(^)%THLe ioBJ^ (NYS); Fig. 8 Dearn 35293
(XK'^^'vVa or Ehlers 1313 (A^S): Fig. 10, Miami, Dade Co., Fla., Meredith,
Ma?i7 1917 (ANS ; Fig. 11, Pennell 4186 (UP); Fig 12 Stravsba^h309
(WVa • Fig 13, Greenfield, HiUsboro Co., N. H., BaichelderAng.U, 1911
(ANS);' Fig." 14, BiZYmore h^h. 622c (UP); Fig. 15, House 19001 (NYS); Fig.

^\he'iS^^^ were all taken with the aid.of a Bausch and Lomb

48 mm microtessar lens. The magnification is approximately 12x.

Explanation op Text Figures

The lines showing the glacial moraines are taken from Antevs (2), and the

Dosition of the Fall Line is essentially that shown by Fenneman (J6). A few

Records on the maps are indicated by circles rather than solid dots; these are

doubtful either as to exact locality or as to exact identity of the plant m

question.

Literature Cited

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1936] McVaugh, — North American Species of Lobelia

361

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Pursh, F. Flora Americae septentrionalis. 2 vols. London, 1»14.

Rafinesque, C. S. Annals of Nature. 1: Le^^gjon, Ky., 1820. Re-
print by T. J. Fitzpatrick, Iowa City, la., Jan. 15. 190».

1 i

u

362

Rhodora

[October

58.

69.

60.
61.
62.
63.

64.
65.

66.
67.
68.

69.

70.
71.

72.
73.
74.
75.
76.

77.
78.

Atlantic Journal and Friend of Knowledge. 1: no. 4.

Philadelphia, winter of 1832.

New Flora and Botany of North America.

Part 2: Neo-

phyton. Philadelphia, 1836.
Ray, J. Historia plantanim. 1:731-747. London, 1686.

. 1. c, 2: 1325. London, 1688.

RiviNus, A. Q. Introductio generaHs in rem herbarium. Lipsiae, 1690.
Rock, J. F. A monographic Study of the Hawaiian Species of the Tribe

Lobelioideae, Family Campanulaceae. Mem. Bernice P. Bishop Mus.

7: no. 2. Honolulu, 1919.
RoEMER, J. J., and Schultes, J. A. Systema vegetabiUum. 5: 39.

Stuttgardtiae, 1819.
RuDBECK, O. Act. Ups. 1720. p. 97. t. 2. (according to Linnaeus, Spec.

PL 2: 929. 1753). This is apparently the following: Acta literaria

{et scientiarum) Sveciae of the Kongliga Vetenskaps-Societeten,

Upsala.
Rupp, H. B. Flora Jenensis. Francofurti & Lipsiae, 1718.
Rydberg, p. a. Catalogue of the Flora of Montana. New York, 1900.
Small, James. The Origin and Development of the Compositae. London,

1919. Reprint from New Phytologist vols. 16-18, 1917-1919.
Small, John K. Studies in the Botany of the Southeastern United States.

BuU. Torr. Bot. Club 24: no. 7. 338. 1897.

. Flora of the Southeastern United States. New York, 1903.

. Manual of the Southeastern Flora. 1293. New York,

1933.
TouRNEFORT, J. P. DE. Institutioncs rei herbariae. 3 vols. Paris, 1700.

. 1. c, 1: 163. Paris, 1719.

Walter, T. Flora Caroliniana. London, 1788.
Willdenow, K. L. Hortus BeroHnensis. 1: Berlin, 1806.
. 1. c, 8 : plates 85, 86. Berlin, 1809. Plate 85 is to accompany

the original description of L. fidgens; plate 86 is to accompany that of

L. splendens.
WiMMER, F. E. Studien zu einer Monographic der Lobelioideen. Fedd.

Rep. Spec. Nov. 26: 3-4. 1929.
Wood, A. Class Book of Botany. New York, 1861.

f

I i

V i >

V- >*'.▼

^ w*

u

THE INFLUENCE OF SEVERAL BENZENE DERIVATIVES

ON THE ROOTS OF LUPINUS ALBUS

4

* ♦

I* ♦

4

' !
jj

A DISSERTATION

IX BOTANY

PRKSEXTKI) TO THK FACULTY OF THE GUADIATK SCHOOL ((F THE

UMVEUSITY OF PENNSYLVANIA IN PARTIAL FULFILL.MLNT

OF THE KEQUIREMENTS FOR THE I)E(;REE

OF DOCTOR OF PHILOSOPHY

•^ *

V.

* '■A

MOTHER MARY CHRYSOSTOM

« V

>»!

Reprint from
The American Journal of Botany, Vol. 23, No. 7

PHILADELPHIA
1936

t

*f

Reininted from tho Amkkican Journal of Botany, Vol. 23, No. 7. 461-471, July, 1936

Printed in U. S. A.

THE INFLUENCE OF SEVERAL BENZENE DERIVATIVES ON THE

ROOTS OF LUPINUS ALBUS '
Mother Mary Chrysostom

Bkxzene substitution products have long afforded
a very attractive field of research to the chemist and
bacteriologist and, more recently, to the plant path-
ologist. To the chemist, the ease of preparation of
these substances and the evident correlation between
their chemical properties and their structure present
almost unlimited scope for investigation. The bac-
teriologist early discovered their antiseptic value and
is now following the chemist in testing the newer
derivatives for possible use as antiseptics and germi-
cides. Literature in this field is voluminous and con-
stantly being augmented. Owing to the fact that
various plant products are benzene derivatives often-
times formed in the course of destructive metabolism,
the plant pathologist is beginning to consider their
presence in the tis.^ue as the active principle of im-

1 Received for publication April 3, 1936.

The writer wi.shes to express her deep gratitude to
Protf'ssor Rodney H. True for oUHgt sting the prouicm
and for his unfailing interest during the course of the
work, and to Doctor John R. Skeen for his kind assist-
ance in reading the manuscript.

munity to certain diseases. Such investigations have
stimulated research on the use of many of these
substances as fungicides and germicides.

There is a decided paucity of literature, however,
on the benzene conii)oun(ls in relation to the higher
plants. True and llunkel (1898) investigated the
influence of several phenols, aromatic acids, and their
salts on Spirogyra and the roots of Lupinus albus.
They concluded that the toxic effect of the benzene
nucleus was modified to a greater or less degree by
the introduction of various radicals into it. Brown
and Tinker (lOKia, lOltJb), in their studies on the
I)ermeability of the seed coats of Hordeum vulgare,
found that various ])henols were rai)idly absorbed.
Brenchley (lOlS), in a study of the effects of phenols
on barley and i)eas with the possible use of these
substances for soil sterilization, found that high con-
centrations were very toxic to the roots, while weak
concentrations did not seem to retard growth. Re-
cently, Eisenmenger (1931) studied the effect of silica
gel in modifying the toxicity of phenol to soy bean

462

AMERICAN JOURXAL OF BOTAXV

seedlings. It would seem then that a quantitative
study of their influence might be of interest.

In their studies on the penetration of various
morganic substances into plant tissue, True and
Bartlett (1915), Osterhout (1922), and Stiles and
J0rgensen (1914) have measured the electrical con-
ductance of the tissue itself or of the solutions bath-
ing the tissues as an indication of changes in permea-
bility of the membranes. Tiie method gives a fair
quantitative measure of increase of electrolytes in the
tissue or in the liquid surrounding it. Thus, to con-
sider only the solutions bathing the roots, if' conduc-
tance increases there has been exosmosis of electrolytes
from the tissue; if it decreases there has been ab-
sorj)tion from the solution wilh or without exosmosis
of electrolytes. As a result of his study on exosmosis
of electrolytes from potato tissue, Stiles (1927) con-
cluded that increase in electrolytes accompanied by
evident turbidity might be considered the result of
mjury to the membrane and therefore a rough meas-
ure of toxicity. With this in mind electrical conduc-
tance of the solutions at constant time intervals will
be compared as a measure of exosmosis from the
roots; that is, of the toxic effect of the solution.

The solutions studied are of three classes: (a) the
acidic— the phenols; (b) the more strongly acidic—
the acids; (c) the weak bases— the amines. Conse-
quently a comparative study of changes in hydro-
gen-ion concentration of the solutions has been made.
To correlate the effects noted, the physiological con-
<lition of the roots was observed.

Experimental MATEHiAL.-^'eec/s.— Seeds of Lupi-
71US albus were selected and grown by the usual
method in chopi)ed sphagnum until the primary roots
were between 00 and 70 nun. in length. This stage
of growth was reached in about three davs. Seedlings
were washed free of sphagnum in distilled water ami
drieil with absorbent jiaper. The seed coats were
then carefully removed, and the seedlings were i)laced
m culture jars. These consisted of oOO-cc. beakers
covered by the lid of a Petri dish which was pierced
m four i)laces to allow the roots to go through. Each
seedling was fixed in place with absorbent cotton.

Solutions.— AW the solutions used in this study were
made volumetrically from stock solutions of C.P.
chemicals. The cresol i)urchased was a U.S.P. mixture
of ortho-, para-, and meta-cresol. The water used
was redistilled from Pyrex glass and its conductance
varied between 1 and 2 X 10-6 mhos. As it was not
desired to kill the roots immediately, a series of i)re-
liminary experiments was performed using at least
ten concentrations of the solutions. As a result, the
following concentrations were chosen for phenol,
cresol, resorcinol, pyrogallol, aniline, methvl aniline-'

20 X 10-4 :^r^ 8 X 10-4 yi^ .^^ 2 x 10-4 :m. p^r

benzoic, salicylic, and gallic acids, 16 X 10 -s yi
8 X 10-5 ^r, and 2 X lO^s M were used. ' '

Seedlings were also grown in a standard nutrient
solution and in distilled uitpr fnr ni,rt„. ., r^( p„^
parison.

[Vol.23,

Methods and procedure. -Resistances were read
on a Leeds and Xorthrup conductivity apparatus
and recorded as mhos.

Ilydrogen-ion readings were made by the quin-

h^-drone titration method using Mcllvaine's standard

buffers-citric acid and disodium hydrogen phosphate

All cultures were kept in the dark in a constant

emperature room. During one series of experiments

he temi)erature of the solutions remained at 23-

1 C, and durmg the series ])erformed at what is

designated ' high " temperature at 28 ± PC

The conductance and hjdrogen-ion concentrations
of the solutions were taken, and the roots were fixed
m i,os,ti«n immediately. At the end of one hour
con, lu-tance was taken by removing the roots to an
empty beaker and inserting a dip electrode in the
solution. Ihe roots were replaced immediately. In
all solutions except that of the acids there was a slicrht
iiK'rease in conductance, },ut this was not significant
enough to be incorporated in the curves. The acids
revealed a slight decrease. Then conductances were
taken at regular time intervals of 24 hours for ei-ht
|lays. At the same jjeriods hydrogen-ion concentni-
ions were read on a 10 cc. sam])le taken from the
< ilture jars. This sample was rei)laced bv ten cc
ol the original solution jjIus a quantitv of water
necessary to bring the level back to 500 cc

A second similar series was studied, but the samples
aken Irom the culture jars for hydrogen-ion readings
u ere boiled or one minute, cooled to original tem-
perature and then titrated. It was hoped to drive
on cu^ b}' this procedure.

results

For this i,resentation it is desirable to treat the
ellect of the solutions in three sjjecific groups: (1) the
phenols; (2) the aromatic acids; (.3) the amine^
I nder each sei)arate heading will |,e considered con-
ductance hydrogen-ion concentration, and jihysio-
logical effects. As rate is more interesting in 'this
study than gross change, grajihs are i)lotted as semi-
og— I.e., log conductance against constant time in-
tervals of twenty-four hours. All value< have been
corrected to 20 C.

Phenols. — Co/?r///r^///re. — Figures 1 '^ 3 and 4
show the effects of the four phenols used' on the con-
<luc'tance of the solutions. It will be noted that the
general shape of the curves is the same: A very great
nicrease in conductance during the first twenty-four
hours is followed by a gradual increase during the rest
ot the exj)eriment. Pyrogallol diflers from tho other
solutions m that its initial increase is extended over
a period of two days and that the increase for the
remainder of the experiment is much greater per day
than that of the other solutions. Except in the ca^e
o cresol and pyrogallol, concentration makes verv
little difterence. In the two solutions mentioned the
strongest concentration produced the greatest chancre
ui.u nils was decidedly more marked than in the
other two.

7

Fi,. l-6.-Chu,mos in conductance of soh.tions.~Fig. 1- P^ond (23°C.).- Fi.^ 2. R^^^^^^^^ (23°C.),

Cresol (23°C.).-Fig. 4. Pyrogallol (23^C.).-Fij?. 5. Phenol (28°C.).-Pig. 6. Crc^ol (28 C).

Fig. 3.

IRREGULAR PAGINATION

464

2xlO-4M

1»H pH
U. B.

Phenol

8xlO-4M

pH |)H
V. JJ.

AMERICAN JOURNAL OF BOTANY

Table 1. Hydrogen-ion concentration.'^

Cresol
20xlO-4M 2X10-4M 8xlO-4M 20xlO-4M 2xlO-4M

[Vol.23,

Resorcinol
8xlO-4M

1)H pH
U. h.

20xlO-4M

pH ],H
U. B.

Original
1st Diiv
2nd Day
31(1 Dav
8th Dav

5.6
5.4
5.4
5.2
5.5

5.6
4.9
6.1
5.9
6.1

5.4
5.4
5.2
5.5
5.4

5.4

4.8
6.1
5.8
6.4

5.5
5.2
5.2
5.5
5.2

5.5
4.8
6.3
6.0
6.1

5.5
5.4
5.6
5.1
5.1

5.5
5.1
5.9
6.1
6.0

pH
U.

5.4
5.5
5.4
5.2
5.4

1)H
B.

5.4
5.2
6.1
5.9
6.6

pH
U.

5.5
5.7
5.7
5.4
6.4

pH
B.

5.2
5.4
6.2
5.9
6.6

J>H
U.

5.3
5.8
5.5
5.7
5.5

pH
B.

5.5
5.7
5.8
6.5
6.4

pH
U.

5.4
6.3
5.6
5.6
5.6

pH
B.

5.4
5.6
6.0
6.5
6.3

pH
U.

Pj'roKallol

2xlO-4M 8xlO-4M 20xlO-4M 2x10 5M

j)H pH
U. B.

1)H pH
U. B.

I)H pH
U. B.

pH j)H
U. Ji.

Benzoic

8xl0-SM

1)H i)H
U. B.

16xlO-5M

1)H pH
U. B.

2xlO-5M

Original
l.st Dav
2nd Day
3rd Dav
81 h Dav

4.8
5.2
5.6
5.6
6.3

4.8
4.5
5.0
5.4
6.3

4.6
5.1
5.6
5.2
5.5

4.6
4.8
5.5
5.6
5.7

4.4
5.2
4.7
4.6
5.7

4.4
5.4
4.3
4.6

4.8

5.0

5.8
5.7

5.8
5.8

5.0
5.7
6.2
6.0
6.4

4.3
5.6
5.6
5.6

5.8

4.3
4.8
6.0
6.0
6.5

3.8
5.8
5.7
5.7
5.8

3.8
4.6
5.7
6.3
6.4

pH
U.

5.7

5.8
5.6
6.0
6.2

1>H
B.

5.7
5.9
6.2
6.2
6.6

Salicylic

8xlO-5M

pH i)H
U. B.

pH
B.

5.5
5.6
5.6
6.6
6.4

16xlO-5M

1)H pH
U. B.

4.9
4.9
5.5
6.4
6.1

4.9
4.9
6.3
6.1
6.4

4.2
4.2
5.7
6.4
6.1

4.2
4.2
6.2
6.1
6.6

Gallic

Aniline

2xlO-5M 8xlO-5M 16xlO-5M 2xlO-4M 8xlO-4M

Methyl aniline

pH pH
U. B.

Original
l.st Day
2nd Day
3rd Day
8th Day

4.8
5.2
5.5
5.3
5.7

4.8
5.6
5.5
6.5
6.2

pll
U.

4.3
4.5
5.4
5.2

5.8

1>H
B.

4.3
4.7
5.4
6.2
6.2

pH i)H
U. B.

I.H pH
U. B.

I.H j.H
U. B.

20\10-4M

pH pH~
U. n.

2xlO-4M 8xlO-4M 20xlO-4M

4.0
4.3
5.6
5.4
5.7

4.0
4.7
5.2
6.1
6.1

5.1
5.5
5.5
5.3
5.7

5.1
5.9
6.1
6.3
7.1

5.4
5.6
5.5
5.3
6.2

«U niean.s unb()il(>d; Ji luean.s boiled.

5.4
6.0
6.3
6.3
7.3

5.8
5.8
5.8
5.5
5.1

5.8
6.2
6.2
6.4
7.2

i>ri

U.

5.6
5.6
5.6
5.5
6.0

pH
B.

5.6
6.3
6.3
6.3
7.2

pH
U.

5.7
5.3
5.3
5.4
6.3

pH
B.

5.7
6.2
6.2
6.3
7.5

pH pH
U. B.

6.0
5.6
5.6

5.8
6.8

6.0
6.3
6.3
6.5
7.3

A closer study reveals the further fact that jdienol
and resorcinol are quite similar in initial effect, but
considerably different in end results; cresol causes a
greater increa.se durinji; the first twenty-four hours
than j)henol or resorcinol.

Temperature effects. — Figures 5 and 6 show the
results of the series of e.\j)ennients conducted with
I)henol and cresol at a tenii)erature of 2S\ In all
cases except the most concentrated cresol solution,
there is a greater e.xosmosis from the roots than at
the lower temi)erature. This may account for the
increase noted in the grai)hs.

In general, then, a study of the changes in conduc-
tance of the.se solutions reveals that: (1) There was
an increa.se in conductance in all concentrations;
(2) this increa.se in conductance was greatest for all
solutions (luring the first twenty-four hours; (li) there
was no marked difference as a result of concentration
except in the stronge.st cresol .-md pyrogallol .solutions;
(4) e.xosmosis from the roots increa.sed in the follow-
mg order: i)henol, resorcinol, cresol, i)yrogalIol; (5)
exosmosis was greater from the roots at 28^ into
phenol and cresol solutions than at the lower tem-
perature.

Hydrogen-ion concentrntiom.—Two sets of reading.s

• - "* ^-^^ '' --/iiiiiMi.. me jii.>i, tie-ignated as

unboiled, is the reading as it was taken on the solu-

tions immediately after they were removed from the
cu.ture jars. The second, designated as boiled, is the
reading taken on a sample which had been boiled for
one m:nute. In i)re.senting the first set of readiness

'VrvT"^'"'"^'^ ^^'""^ ^^'^ '^^'^-^ ''''^ constantly throwing
off C( X, into the .solutions and that the quantity has
been .shown to be greatest during the first twenty-four
o"r? '"V^ gradually fall to a minimum (Aslander,
J.w.i). iiuis the amount jiresent nmst tend to equi-
lil)num They are expected therefore to be more
acid. It was hoped that Imiling would drive off the
LU formed. It would seem, however, that durincr
l)oiling other changes may have taken place at least
in some of the solutions -e.g., tho.se which became
dis inctly more acid during the first twenty-four hours
and remained acid even after boiling. Beadin-s are
presented for the first three days and for the end' of
the ex])eriment.

Phk.xol.— ('on.<idpring the unboiled solutions first
It will ],e noticed that there is a tendency to greater
acidity, a fact which would be expected.' But there
IS no neutralizing action in the.se solutions and -.11
end more acid or as acid as the original. This may
be due to increa.sed respiration with con.sequent evo-
lution of more carbon dioxide than normal

in the boiled solutions there is a definite drop to
much greater acidity at the end of twenty-four hciurs

)

July. 19361

CHRYSOSTOM — BENZENE DERIVATIVES AND ROf

465

The contrast here with the unboiled solutions is so neutralizing action ' roots in ciilture solutions to

'reat that there must have been more changes in the Nil, production. ,sequently there is a sharp in-

^^kiticln^;^ than driving off CO... Perhaps the roots crea.se to 6.1 or 6.:i id all solutions end considerably

have excreted some other volatile compound which more basic than th( began,

r^iiains 1 n(listur]>ecl in the unboiled, but is driven off CnEsoL.-There >■ onsiderable variation in the un-

in boiling Priani.schnikow (1928) attributed the boiled solutions. \^ ,h some fluctuations, the solu-

T.^BLK 2. Physioloffical effects of the phnutls
Roots

Primary root

Mature

Phknol

Tip

re^jion

2xlO-4M

Flaccid
5th day

Thick and
knotted

8xlO-4M

Flaccid
4th day

Thick and
knotted

20xlO-4M

Flaccid
4th day

Thick and
knotted

Cresol

2xlO-4M

Flaccid
3rd day

Normal

8xlO-4M

Flaccid
attenu-
ated
2nd day

Normal

20xlO-4M

Flaccid
attenu-
at(Ml
1st day

Flaccid
4th day

Resorcinol

2xlO-4M

Flaccid
3rd day

Normal

8xlO-4M

Flaccid
2nd day

Normal

20x1 0-4 M

Flaccid
2nd day

Normal

Pyrog.allol

2xlO-4M

Flaccid
1st day

Flaccid

8xlO-4M

Flaccid
1st day

Flaccid

20xlO-4M

Flaccid
1st day

Flaccid

Benzoic Acir

2xlO-5M

Flaccid
3rd day

Normal

8xlO-5M

Flaccid
2nd day

Normal

16xlO-5M

Flaccid
1st day

Normal

S.ALIC^XIC Ac

[D

2xlO-5M

Flaccid
3rd day

Flaccid

8x10-5 M

Flaccid
2nd day

Flaccid

16xlO-M

Flaccid
l.st day

Flaccid

Laterals

Time of

appearance

1st day

1st day
1st day

3rd day
3rd day

3rd day

Character
Scattered

Scattered
bunched at

top
Bunched at

top

Scattered

Bunched at
top

Bunched
few along
roots

Length
3-4 cm.

3-4 cm.
2 cm.

3-4 cm.
2 cm.

1 cm.

1st day

Normal

4-5 cm.

Ist day

Normal

4-5 cm.

l.st day

Bunched at
top

4-5 cm.

1st day

Few

1-2 cm.

1st day

Few at top

1 cm.

3rd day

Bunched at
top

No
{irowih

3rd day

Scattered

3-4 cm

3rd day

Scattered

3-4 (in

3rd day

Aggrega'ed
at top

3-4 cm

1st day

Scattered

3 cm.

l.st day

Agprenated

at top

3 cm.

3rd day

VV iioried

4 cut.

Cotyledon?

Epicotyl

Normal
Normal

Normal

Growth
Growth

Growth

Slight
separation

Slight
separation

No growth
No growth

Slight No growth

separation

Normal Slight

growth

Normal Growth

Normal

Growth

Normal

Growth

Slight No growih

.se])aration

No No growth

.separation

Normal

Growth

Normal

Growth

Normal

Very lilt le

growth

Slight

Slight

separation

growth

Slight

Slight

separation

growth

Nu

No

separation

growth

466

AMERICAN JOURNAL OF BOTANY

Tahlk 2. Con finned.

Root;

Gallic Acid
2xlO-'^M

8xlO-5M

16xlO-5M

Aniline
2xlO-4M

8xlO-*M
20x1 0-4 M

Mkthyl
Aniline

2xlO--*M

8xlO--*M

20xlO-4M

Tip

F'laccid
4th day

P^laecid
2nd day

Fhipcid
2ik1 day

Flaccid
5th day

Flaccid
4th day

Flaccid
3id day

Priinaiy loot

Mature
i't'y;ian

[Vol. 23,

Cotyledons Epicotyl

July, 1936]

CHRYSOSTOM— BENZENE DERIVATIVES AND R0(

Laterals

Normal

Normal

Normal

Time of
appearance

Ist day
l.st day
1st day

Normal

Flaccid
Va lens I h

Flaccid whole
length

2nd day

2nd day
2nd day

Flaccid
2nd day

Flaccid
2nd day

Flaccid
1st day

Flaccid whole
lenfith

Flaccid whole
lenv;th

Flaccid whole
length

2nd day
2nd day
2nd day

tions reach their most acid condition on the third
day. The two lower concentration?; end more arid
than the original or just the same, while the highest
concentration l)ecomes as basic as the boiled solutions.
In the l)oiled solutions except for the first tlrop in
acidity there is an almost unbroken jirogression to a
more basic condition. Thus phenol and cresol may
be looked upon as having practically the same effect
in stimulating changes of hydrogen-ion concentration.
Kksohcinol. — Results in the strongest unboiled
solution are not available, owing to unavoidable mis-
haps. In the two weakest concentrations there is a
progression with nuich fluctuation to more basic
solutions. After boiling there is an almost unbroken
sequence to pH of 6.3 and 6.4.

PYROfiALLOL. — Original solutions were much more
acid than any of the phenols. There is a more gradual
change to a more basic solution in all cases. The
strongest boiled solution ends decidedly acid. Since
pyrogallol changes chemically in air and on boiling,
other factors are affecting hydrogen-ion readings, and'
neutralization is probably the resultant of many
factors.

Phif.siologicnl effects. — Tho jihysiological effects of

the phenols are summarized in table 2. It will be

noted that the strongest cresol solution and all nvro-
fill.' « -- -■•

^ II

gcuiui .>(HiHiuij.- .snow marked toxic etfects. Jn these
solutions the root tips became flaccid almost imme-

Charact(>r

Normal
along whole
length

Normal

whole

length

Normal
half of
region

Scattered
bunched at
top

Runched at
top

Very small
bunches

Length
3-^ cm.

3-4 cm,

3-4 cm.

Normal

Normal

Normal

Growth

Growth

Growth

1-2 cm.

1 cm.
1 cm.

Slight

Slight

separation

growth

Slight

No

separation

growth

Slight

No

sejjaration

growth

No sej)-
arution
No sep-
aration

No sep-
aration

(liately and this condition extended to the mature
region ot the root. Laterals were tardy in appearing
HI the cresol solutions and in the most concentrated
pyrogallol solution. Subsecjuent development con-
sistetl in an aggregation of very stunted laterals at
the top ot the roots. Seedlings in these .solutions
showed very little toj) development.

In the phenol an<i resorcinol solutions, development
was more or less normal.

Turbidity ai)peared at about the same time in all
solutions.

Aromatic Acw^.—Comlnctnnce.—CuvyeH for these
results are presente.l in figures 7, 8, and 9. The strik-
nig contrast between the.se grai)hs and those of the
phenols IS very eA'ident. Considering the three acids
together there is, in general, a decrease in conduc-
tance followed by an increase and a final levelling
except in the highest concentration of salicylic acid "'
Bvnzmc add. — In the lowest concentration, the
Hiitial decrease in conductance extends over a period
of twenty-four hours, in the next concentration over
a i)erio(l of forty-eight hours, and in the hicrhest
concentration, ninety-six hours. After this peHod
conductance increases until the sixth dav when all
curves show a constant level until the end 'of the ex-
periment. It will }»e noted fh^f fhere i- very little
increase in the strongest solution, more in the inter-
mediate and most in the weakest. Except in the

3

latter solution, leach does not compensate for initial

absorption. ,

Salicylic acid.— Decrease in conductance in salicyhc
acid solutions is similar to that in benzoic acid in
point of time, but much greater in amount. In the
weakest concentration there is a decrease for twenty-
four hours, in the intermediate for forty-eight hours,
and in the strongest for one hundred twenty hours.
The two lower concentrations then increase in con-
ductance until the sixth and seventh day, when they
remain at the same level. The highest concentration
gradually rises in conductance until the last day,
when it shows a very sharp increase. Only the weak-
est solution reaches a conductance greater than the

original value.

Gallic acid. — This acid produces much less effect
than the other two. There is no decrease in conduc-
tance in the weakest concentration, a slight decrease
for forty-eight hours in the intermediate, and for
seventy-two hours in the highest. The lowest con-
centration then gives the most increase, while the
two higher increase slightly for two and three days,
respectively, and then remain at comparatively the

same level .

' In all of the acid solutions only the weakest con-
centrations reach a final conductance which is greater
than that of the original solutions. It would seem,
then, that whatever ions were removed from the
solution by the plants were not compensated for
completely by the end of the experiment. In order
of increasing effect these acids would be arranged as
follows: gallic, benzoic, salicylic.

In general, graphs for the acids show: (1) A de-
crease in conductance at first for varying periods,
.seemingly dependent on concentration; (2) a very
decided concentration effect; (3) a final increase in
conductance above its initial point.

Hydrogen-ion concentration.- Changes in hydro-
gen-ion concentration for the three acids may be
presented in a concisely comparative study. In all
solutions there is a distinct neutralization reaction
on the part of the roots under the influence of which
the solutions end much more basic than they were

originally.

Unboiled solutio?}s. —li will be o])served from table
1 that the greatest change in the hydrogen-ion con-
centration of all the acids takes place during the first
forty-eight hours. From then to the end of the ex-
periment there is a progressive tendency to a more
neutral end point with only slight variations. This
point is the same for all concentrations of each acid—
e.g,, 5.8 for benzoic, 6.1 for salicylic, and 5.7 for
gallic. The initial change in acidity can be corre-
lated with decrease in conductance during the first

two days.

Boiled solutions.— Y.xcepi for the fact that all read-
ings are more basic than those of the unboiled, the
results seem to be the same as in the latter solutions.
I he striKiMg cii.iJi^t i»i .HHiiit i4>iiiii^ li.v^^ •'. ....-..,.,
four hours in the unboiled solutions seems to require
at least forty-eight hours in the case of benzoic and

sahcylic acids and
gallic. End i)oint.-
the same — e.g., ().4

467

:^nty-two hours in the case of
the solutions of each acid are

6.5 for benzoic, 6.4 or 6,6 for
for gallic.

-If the effects listed in table 2

e noted that salicylic acid is
benzoic or gallic. There is a
root in all the concentrations

igation. In the most concen-
have developed, but are

salicylic, and 6.2 or
Physiological efjv
are studied, it wil:
more toxic than eii
softening of the eni
with no apparent ( .^c
trated solution l.iti ;ls

whorled in arrangcinriit since there was no accom-
panying elongation oi the root. In the same .solution
cotyledons did not m |>arate and there was no growth
of the epicotyl.

Benzoic acid is new greatest in toxic effect on the
roots. Tips are flacnd the first day and laterals are
late in appearing and then scattered or aggregated
at the top of the re'.iion of growth. Tops show de-
velopment in all the solutions.

Gallic acid seems to have less effect than the other
two. Only the root tips .seem affected and the region
of side root growth is shorter in the strongest con-
centration.

All solutions were turbid from about the third day.
In order of increasing toxicity the solutions are:
gallic acid, benzoic acid, salicylic acid.

The amines, — Conductance. — Figures 10 and 11
show changes in conductance of the solutions of the
amines. In general, there is a very great increase
in conductance; much greater than in the phenols.
The rate of change in both solutions is greatest dur-
ing the first twenty-four hours. Concentration of
these .solutions .«eems to have a much greater effect
on the roots than that of any other substance studied.
Using the amount of exosmosis of electrolytes as a
criterion, methyl aniline is much more toxic than
aniline.

Temperature ,'?.9\— Just as with phenol and cresol,
higher temperature causes a greater increase in con-
ductance in the solutions. With both .substances the
curves are very mut h steeper than those at the
lower temperature. If will be noted that the curve
for the strongest methyl aniline .solution leveled at
the end of the fourth day. This was probably due
to the fact that roots in solutions of that strength
very quickly reached a state of disintegration. Per-
haps the ec|uilibrium cstabli.shed at that point is an
indication of death. No other solution reaches equi-
librium so early in the experiment.

Hydrogen-ion concentration. — With the general-
ization that the boiled .solutions are very much more
basic than the unboilcl, the two sets of solutions may
be considered togetlx r. The two unboiled .solutions
of methyl aniline became slightly more acid the first
day. This was prob.d^ly due to the carbon dioxide
factor. The other solutions changed steadily in the
direction of neutrality. End points for all these
«njiitinn< ire more hisic than anv vet considered.
The readings for the first three days and for the
last day are include* I in table 1.

tnr

Ui

0

z

<

»-
o

3
P
Z
o
o

(9
O

50

50

20

: t I ' 1
i I I J '

III

"+tT

|11t

ill

! i ! !

fn-t-f

-H++

J
f4-H-+

: ! I t I

w

III

: f i

+4-

i i i ! !

1 ijf

: 1 I I 1

Tjrn

i+-

4

j I

-ft

tsk

vr

m

^4

i t ! :

m

TlMe IN DAY5

ti-Mfflt

iffil

I r i I

T»r-i£ IN t>AY5

TIME

ir4

15 Af 5

TIME IM 'DA^'S

3

July, 19361

CHRYSOSTOM — BENZENE DERIVATIVES AND Hoi

468

Physiological efjects.— The results listed in table 2
point to tlie fact that these solutions are nuich more
toxic than anv of the others studied with the possible
exception of * pyrogallol. Root tips became flaccid
tJiifihtlv later than in the aromatic acids, but this
softening was progressive, except in the weakest ani-
line solution. Finally the whole root was affected.
Laterals grew to some extent in the two weaker ani-
line solutions but not at all in the strongest except
in small aggregations at the top of the root. In
methyl aniline solutions, laterals were inhibited imme-
diately on formation. Tops showed a slight growth
in aniline, but none in methyl aniline. The loss of
turgidity in the latter solutions extended throughout

the seedling's.

^UTIUE^T— Conductance. — The nutrient solution

showed a decrease in conductance for the first five
days, then an increase. This is in accordance with
what might be expected, since the plants are con-
stantly taking ions from the solutions. The increase
at the end of the experiment was probably caused by
unequal exchange of ions.

HlliJrogcn-ion concentration. — Thc^^e solutions be-
came progressively more acid, even those which were
boiled to remove the carbon dioxide. Evidently the
cause of acidity in the unboiled as well as the boiled
solutions was not due entirely to carbon dioxide.

Fhysioloqical effects.— In the lupines grown in the
nutrient solution there was a plentiful growth of
lateral roots and a ('ecided elongation of the primary
root. Ilypocotyls elongated, the cotyledons opened,
nnd there was conside.-able growth of the epicotyl.
l^xcept for the yellow color of the leaves, the ]ilants
could be considered in good condition. The root>^
were turgid and the solutions clear at the end of the

experiment.

Distilled water.— ro^Jz/rfa/^rr— The conductance

of distilled water in which plants have grown in-
creases in an almost constant ratio from day to day.
This e!Tect is in accordance with the work of various
investigators who attribute the rise to electrolytes
dilTusing from the roots.

Hydro'ien-ion concentration.— Thv^e readings show
very little difference from those already discussed.
There is a neutralizing tendency much more marked
in the l)o'le I than in the unboiled solutions. Few
fluctuation^ a»-e noted in the readings from day to day.

Plnisiolofiical efjects. — Plants grown in distilled
water, although not normal in develoi)ment, were in
very good condition in comiiarison with those grown
in toxic solutions. There was a decided elongation
of the i^riniary root with a scatte-erl growth of lat-
erals. These, however, attained considerable length.
At the close of the ex])eriment the t'ps hid become
poft. ?ome of the hvijo-otvls showed clearing and
had collapsed at the end of the time, the primary
bud onened and the stem elon'^nted sliditly. Solu-
tions did not become turbid until the fifth day.

Fie 7 8 10 M 12 1.3.— ChiUi-o^ in conductanrr of
acid (23^C.') -Fi^. 10. Aniline (23"C.).-Fig. 11. Mc hyl
aniline (28°C.).

From the result-
exosmosis from the
result of cell in,iur\
"I he materials difhi
lytes, since they <•
tance of the solutin
all solutions showe,!
substances may li;i\'
of the cells, makiuu
other substances to
];ermeable. This v\\'\
Stiles and .T0rgens('n I
of exosmosis from rot
the other hand, the i
absorbed by the roni-
may have caused the
substances. This ex
Brooks (1916) to hi-

409

SCUSSION

sented, it is very evident that
ts of the seedlings studied is a
leh ultimately results in death.

from the roots were electro-
•d an increase in the conduc-
and colloidal substances, since
persistent turbidity. The toxic
irst attacked the surface layer

permeable to electrolytes and
iiich it would normally be im-
mation has been advanced by
1917) as a result of their study
)ts into organic solutions. On
oxic substance may have been

and attacking the cytoplasm,

formation of easily diffusible
1 Sanation has been given by
: results from studies on exos-

Fip. 9. Chanjrcs in tonductance of gallic acid solntion.s
(23°(\).

mosis from roots, 'flu- present study seems to present
evidence for both explanations. In the concentration
of the ])henols studic^l. the effects on the roots seem
to be i)urely local, pet niit ting good top develoj^ment,
but killing the root iijis almost immediately. Phenols
are known to i)recipiiate and coagulate proteins as
well as other labile (clloidal systems. If the external
surface of the cytopl i-in is colloidal, as many inves-
tigators believe, the tiist effect of the solutions would
be external, but thcv would then be a gradual attack
on the cytoplasm wb h would finally kill. Pyrogallol
killed much more i i idly than the other solutions.
It is more soluble ind more sen^^itive to oxidation
than the other i)li*'; 'N u^ed. Its penetration was

"ct more evident.

- as to what factor caused

u^ed in this study are only

! that there are in the solution

ireat number of undissociated

mo'e rapid and its c
"idle question ari
toxicity. The acid-
slightly dissociated. -
hvdrogen-ions and •

f;^ q cj-.ii

aniline (23°C.).-Fig. 12 Aniline (28°C.).— Fig. 13. Methyl

470

AMERICAX JOURNAL OF BOTANY

[Vol.23,

inolecules. The hydrogen-ions cannot be the only
toxic agent. If they were, benzoic and gallic acids
would bo more toxic than salicylic, but this contra-
dicts the results. It seems that the evidence here is
in iavor ol' undissociated molecules as the i)rincipal
loxic I'.icior. 'J his opinion has been advanced by True
(llOOj lor Lupinus albiis, by Clark (1S99) for fila-
mentous fungi, by Collett (1919) for Paramoecium
t\nd Euplotes, and has been held by many other in-
vestigators.

Ihe amines were most devastating in their action.
The toxic effect was general. They are rapidlv ab-
sorbed and rapidly transj)or<ed to all parts of the
seedlings. They seem to ])roduce a general disinte-
gration of the i)rotoi)lasm with very great leach
greater than in any of the solutions in .si)ite of the
much smaller absorbing surface exposed. They are
both water-soluble and fat-soluble, a fact which' may
account for their ra])id jienetration. Rapidity of
I-enetration of weak bases and alijihatic amines^into
cells has been reported by Harvey (1911) for animals
and ])lants and by Hrof)ks (1923) for plants. This
investigation points to a similar characteristic of the
aromatic amines studied.

Hydrogen-ion concentration-? recorded for this study
cannot be considered significant excejjt in the aciil
solutions. In these the evident agreement between
fall in pll and decrease in conductance i)oints to the
abs()ri)tion of hydrogen-ions as the mechanism of
neutralizaticm. In the other solutions, with the ex-
coi.tion (.f pyrogallol, the .lecrease in pll cannot be
attributed to the absorption of hydrogen-ions as there
IS no evidence to support such a hyjiothesis. Rather,
the fact of exosmosis of colloidal substances from the
roots points to a coml)ination of the.se substances
w:th the hydrogen-ions of the .solution to form more
shghflv dissociated compounds. Ilydrogen-ion.s in
the solution would therefore become fewer. This ex-
planation has been advanced by Mevius (1927) in
h's stiKhe-- of root growth in various solutions."

This investigation shows a correlation l)etween the
rhysiological action and the structure of the molecule
in both the phenol series nnd the amine .«eries
Cresol, which is formed by the substitution of a
meth\l group for one of the hydrogen atoms of the
nuc'eiH, IS more toxic than i)henol. True and Ilunkel
nS9S) noted this fact as a result of their studies on
Lypinuft albus, and Woodward, Kingerv, and Williams

(1934) as a result of their studies on fungi. With
the i)henols having increasing number of hydroxyl
groups— namely, i)henol, resorcinol, and i)yrogallol—
there is a correlation similar to that found by Walker
and Link (1935) in their studies with onion bulb
l)arasites. When the hydroxyl groups are in the ortho
l)osition, there is an increase in toxicity — namely,
l)yrogallol is more toxic than phenol; when the
hydroxyl groups are in the meta i)osition, there is
little or no increase in toxicity— namely, resorcinol is
less to.xic than phenol or as toxic. It is interesting
to add that salicylic acid, the most toxic of the acids
studietl, has the hydroxyl group in the ortho position.
Among the amines a methyl group in the aniline
series increased toxicity. This has been observed to
be true for bacteria also by Bachman (1928).

SUMMARY

A study has been made of the eflfects of several
benzene derivatives on the roots of Lupinm albus—
namely, four i)henols (jihenol, cresol, resorcinol, and
pyrogallolj ; three acids (benzoic, salicylic, and ga'lhc) ;
two amines (aniline and methyl aniline).

The effects have been expressed in changes of con-
ductance and hydrogen-ion concentrations of the
solutions as well as in the physiological influence on
the i)lants.

In general, all solutions increased in conductance
excei)t the acids, which decreased for varying periods
of time and then increased. Except in the lowest
concentrations of the acids, the ions lost to the .<5olu-
tions were not comi)ensated for by the end of the
ex])eriment.

Ilydrogen-ion concentration became considerably
less in all i=!olutions excei)t the most concentrated of
the inrogallol solutions.

There seems to l)e an agreement l^etween toxicitv
and comi)lexity of structure in all of these compounds
Among the jihenols studied, the order of increasing
toxicity IS phenol, resorcinol, cresol, pvrogallol. Amon^
the acids studied, the order is gallic, benzoic, and
salicylic, and among the amines, aniline and methvl
aniline.

There is .some indication that rise in temperature
increases the toxic effect in the case of phenol, cresol
anihne. methyl aniline.

University of Penn.sylvania,
Philadelphia, Pennsylvania

LITERATURE CITED

AsLANDKR. A. 1933. XontralizinK action of plants on
;'(••(! THitnont solutions. Sven.sk. Bot. Tidskr 27-
257-272.

Bachman. O. W. 1928. Chemical constitution and
ponnicidal action. Proc. Soc. Exp. Biol, and Mod
25: 249-250.

Bre.vchley. W. E. 1918. Organic plant i)oison.s. II
I hrnol. Annals Hot. 32: 259-278.

Brooks. M. M. 192.'? .^h„i;, . ^„ thr. ^^^.^p^K.-i:*.. <■
livms and dead colls. III. Tho penetration of certain
alkahos and ammonium s'dts into living and dead
cells. Pub. Health Ro],. 866, p. 2074-2086.

Brooks, S. C. 1916. Studies on exo.smo.sjs. Amer Jour
Bot. 3: 483-492.

Brown. A. J., and F. Tinker. 1916a. Rate of ab.sorption
ot various phenolic solutions bv the .seeds of
llonkum vulunrc and tho factors go\orning tho rate
of diffusion of aqueous .solutions across semi-porme-
a bio membranes. Proc. Roy. Soc. London B, 89:
119-135.

--. AND F. Tinker. 1916b. Selective pormeabilitv.
Iho absorption of i)honoI and other solutions by the

TtVL^^'o^^''''''' ^'"^^'"■'^- ^'°'^- ^°>'- Soc. London
■15, Hy : o7«j — o79.

July. 19361

MARTIN — TOXICITY OF SELENIUM TO PLANTS AND vIALS

471

Cl.\rk, J. F. 1899. On the toxic effect of deleterious
agents on tho germination and dovelopment of
certain filamentous fungi. Bot. Gaz. 28: 289-327,
378-404.

Collett M. E. 1919. Toxicitv of acids to ciliatc in-
fusoVia. I, II. Jour. Exp. Zool. 29: 443-472.

Eisenmenoer, W. S. 1931. Factors modifying the
toxicity of phenol. Plant Physiol. 6: 325-332.

Harvey. E. N. 1911. Studies in tho permeability of
cell's. Jour. Exp. Zool. 10: 507-556.

Mevius, W. 1927. Kalziuni Ion und Wurzehvachstum.
Jahrb. Wiss. Bot. 66: 183-253.

OsTERHOUT, W. J. V. 1922. Injury, recovery and death
in relation to conductivity and p(>rineability. Mono-
grai)hs on Experimental Biology 8: 1-259. Phila-
delphia and London.

Prianischnikow, D. 1928. I'ber die Ausscheidung von
Ammonia durch PHanzenwurzeIn der Saurevergif-
tung. Biochom. Zeitschr. 193: 211-215.

Stiles, W. 1927. Exosmosis of dis.-^olved substances
from storage tissue into water. Piotoplasma 2: 577-

601.

-, AND I. J0R(iENSEN. 1914. Tho measun-iucnt of

electrical condiu v as a method of investigation
in plant physiol, " Now Phytol. 13: 226-242.
-, AND I. Joi; v.s. 1917. Studies in perine-

ability. IV. Tl: ction of various organic sub-
stances on the 1 ability of the plant cell and its
])oaring on ( "zm| . t heory of the plasma membrane.
Annals Bot. 31: i 6.

True, R. H. 1900. toxic action of a series of acids

and of their sod salts on Lupinus albu.s. Aiucr.
Jour. Sci. 9: is;;

. AND H. H. 1 . iLETT. 1915. Exchange of ions

l)etwe(>u tli(^ itHM Lupinus albus and culture solu-
tions containing iv.o nutrient salts. Amer. Jour.
Bot. 2: 311-323.

AND C. (;. lirNKEL. 1898. The poisonous

effects exerted on living plants by phenols. Bot,
Centralbl. 75. 76: 2S<)-295. 321-327, 391-398.
W.ALKER, J. C.. AM) K. LiNK. 1935. Toxicity of phe-
nolic compoiuids to certain onion bulb parasites.
Bot. Gaz. 96: lOS 1S4.

WOODW.AHD, G. I.. L. b. KiNGERY, AND R. J. WiLLIAMS.

19.34. The fungiii(l;il i^ower of phenol derivatives.
I Effect of introducing alkvl groups and halogens.
Jour. Lab. and Clin. Med. 19: 1216-1223.

~ -^

Reprinted from Plant Physiology, 11: 195-200, 1936.

- A

- *

I* »

L- V

-" *■

r^ 4

' r
I" ♦

1 ^

" >

■ •• >'

BREAKDOWN OF FRUIT AND VEGETABLE TISSUE DUE TO AN

ELECTRIC CURRENT

William Seifriz
(with two figures)

Experiments in electrodialysis involving the use of natural membranes
revealed the interesting fact that the tissues of fruits and vegetables break
down very rapidly when they are the cathode of an electric circuit, but do
so only to a slight degree, if at all, when they are the anode.

An apple resting in a bowl containing a weak salt solution, with one elec-
trode of an electric circuit penetrating it and the other electrode projecting
into the surrounding solution, will ''rot" after eighteen to twenty-four
hours if the apple is the cathode or negative pole (fig. 1). If the apple is

Fig. 1. Effect of the negative electrode of a 110-volt direct current on an apple
standing in 0.1 N NaCl for twenty hours. The cross-lined portion is well "rotted'* (soft
and dark brown in color) ; the heavily stippled region is considerably discolored (to
green), and the lightly stippled portion is slightly discolored.

the anode or positive pole, the tissue is little damaged. The foregoing was
the original experiment ; it was repeated many times and then duplicated
with other fruits and vegetables, always with the same result, except that
the degree and kind of degeneration of tissue varied with the material.
Particularly striking were the results obtained with the persimmon and

potato.

■ The experimental findings are listed below. The electrodes were of
platinum and the circuit a 110-volt direct current, with an intervening 0.5
ampere electric-light bulb. The surrounding solution was 0.1 N NaCl, and
the length of the run, eighteen to twenty-four hours. The fruit was im-
mersed in the solution one-half to two-thirds of its height. The degree of
acidity of the normal fruit and of the tissues surrounding the respective

195

<f

IRREGULAR PAGINATION

^ ^

Reprinted from Plant Physiology, 11: 195-200, 1936.

- ^

- 4

I* »

-" *•

U- 4

I-* ♦

V- A

1 ^

*«i

^ >

.' ^

>— /

T

BREAKDOWN OF FRUIT AND VEGETABLE TISSUE DUE TO AN

ELECTRIC CURRENT

William Seifriz
(with two figures)

Experiments in electrodialysis involving the use of natural membranes
revealed the interesting fact that the tissues of fruits and vegetables break
down very rapidly when they are the cathode of an electric circuit, but do
so only to a slight degree, if at all, when they are the anode.

An apple resting in a bowl containing a weak salt solution, with one elec-
trode of an electric circuit penetrating it and the other electrode projecting
into the surrounding solution, will ''rot" after eighteen to twenty-four
hours if the apple is the cathode or negative pole (fig. 1). If the apple is

Fio. 1. Effect of the negative electrode of a 110-volt direct current on an apple
standing in 0.1 N NaCl for twenty hours. The cross-lined portion is well "rotted" (soft
and dark brown in color) ; the heavily stippled region is considerably discolored (to
green), and the lightly stippled portion is slightly discolored.

the anode or positive pole, the tissue is little damaged. The foregoing was
the original experiment ; it was repeated many times and then duplicated
with other fruits and vegetables, always with the same result, except that
the degree and kind of degeneration of tissue varied with the material.
Particularly striking were the results obtained with the persimmon and

potato.

The experimental findings are listed below. The electrodes were of
platinum and the circuit a 110-volt direct current, with an intervening 0.5
ampere electric-light bulb. The surrounding solution was 0.1 N NaCl, and
the length of the run, eighteen to twenty-four hours. The fruit was im-
mersed in the solution one-half to two-thirds of its height. The degree of
acidity of the normal fruit and of the tissues surrounding the respective

195

196

PLANT PHYSIOLOGY

SEIFRIZ

BREAKDOWN OF TISSUE DUE TO ELECTRIC CURRENT 197

poles was obtained by macerating the tissue in a minimum amount of water.
The values, in terms of pH, are given, that of the normal fruit first the
treated last. The word ''rotting" will be used as a convenient term to
express a breakdown or change in the tissue without in any way suggesting
the true nature of the change.

Apple (pH S.6)~Cathode: pronounced rotting ; tissue soft and discolored
(dark brown) ; skin turns green; tissue basic (pH 12.2).

Anode: slight rotting; tissue soft but not much discolored; skin turns
pink; tissue acidic (pH 3.2).
The experiment was varied by placing the electrodes in separate apples
resting in the same solution, and by placing both electrodes in opposite sides
of the same apple ; the results were identical in all three cases.

Pear (pH 5.2)— Cathode: severe rotting; tissue basic (pH 12.2).

Anode: slight softening; tissue acidic (pH 3.6).
Persimmon (pH 5.6)— Cathode: rotting complete; tissue becomes black
exceedingly soft, gelatinous in part, collapsed and much shrunken ; basic
(pH 12.4).

Anode: slight softening and discoloration; acidic (pH 3.0) ; otherwise
normal.

Orange (pH 3.8)— Cathode: some discoloration and softening; tissue
somewhat gelatinous; basic (pH 12.2).

Anode: no pronounced change; tissue slightly more acidic (pH 3.0).
Banana (pH 5.2)— Cathode: rotting throughout the greater part of the
fruit; color nearly black; basic (pH 12.2).

Anode: slight softening and discoloration; tissue acidic.
Osage orange (pH S.2)—Cathode: marked gelatinization ; basic (pH

Anode: tissue not noticeably altered ; acidic.
Onion (pH 5.4)— Cathode: gelatinization of tissue; basic (pH 12.2).
Anode: slight softening; acidic (pH 3.0).

Carrot (pH 4.8)— Cathode: marked rotting, discoloration and shrink-
age; basic (pH 12.0).

Anode: tissue normal, no noticeable change ; acidic (pH 5.3) .
Potato (pH 6.2)— Cathode: complete gelatinization of half of the potato
(fig. 2); basic (pH12.0).

Anode: slight softening; otherwise unaffected; acidic (pH 5.4)—
(Sweet potato reacts in a manner similar to white potato.)
These results indicate that high alkalinity is constantly associated with
breakdown in fruit and vegetable tissue due to the passage of an electric
current.

The negative pole or cathode of an electric circuit is alkaline where elec-
trolytes are involved. This is true because the positive ions of alkali metals

■y

i

+

Fig 2. Effect on potatoes of the negative (left) and the positive (right) electrodes
of a no-volt circuit. The potato (left) in which the cathode was imbedded has become
a soft and glutinous jelly throughout more than half its volume (there is no discoloration),
while the potato (right) containing the anode was but very slightly affected.

wander to the cathode and there unite with the dissociated hydroxyl ions of
the water to form a base. Alkalinity at the cathode may be proved experi-
mentally by adding phenolphthalein to an electrolytic solution into which
two electrodes project ; the color changes to pink at the negative pole.

As the negative pole is alkaline and as decomposition of the tissues took
place at this pole, it is possible that alkalinity per se or some reaction which
is accelerated by alkalinity, is responsible for the decomposition. It is par-
ticularly significant that in all fruits and vegetables with but three excep-
tions, the pH of the tissue at the cathode was 12.2; in two cases it was 12.0
and in one case 12.4, while the original pH values of the tissues differed

considerably. , , , ^ • „*

In addition to alkalinity the causative agent may be the electric current
• as such, heat, bacteria, oxidation, or autolysis. The electric current prob-
ably functions only through some condition which it establishes, such as
reduction. Heat was given special consideration because the fruit and
vegetables often rose to quite high temperatures owing to high resistance ot
the tissues, but as the tissue at both positive and negative poles became hot
and as heat applied separately caused no breakdown in tissue, heat cannot
be a primary cause. As for bacteria, the reaction is too rapid for them

alone to be responsible. .

It appears that the reactions which most probably cause a breakdown in
tissue due to a flow of current are oxidation and autolysis (proteolytic self -
digestion) The alkaline condition developing at the cathode might be
expected to hasten non-enzymic oxidations, as there are many instances in
which alkalinity is known to promote oxidation ; for example a metal
bucket rusts more rapidly when lime is left within it; pyrogallol oxidizes
more readily in the presence of alkali. However, oxidation (involving oxy-

198

PLANT PHYSIOLOGY

gen) as the primary cause of initial degeneration seems unlikely because
no oxygen is formed at the cathode during electrolysis. There is also no

theTthode *''* °"'''"°= ""^"'^ "■■" ''''''''"' ^P^^'''"^ ««*-« ^t

A more probable explanation of the effects observed is suggested bv the

rtrto b^ ' 'T",r'°°- "'^ *'^^"^ ''"^'^ *° •'-"- sorwhlhCa
result to be expected from autolytic changes, especially proteolysis, involv-

ng breakdown of tissue structure. Intracellular proteolydc ;nzymes
(papain, cathepsin) are activated by various reducing substances and
autolysis sets in rapidly when the oxygen supply to tissues is cut off Thus
reducing conditions generally seem to favor autolysis or proteolytic break
down, and the cathode, where electrons are liberated, is the electrode where
reducing conditions would be expected to prevail. Proteolysis, therefore
is possibly the initial reaction. mereiore,

Following this self-digestive breakdown of the tissues, various autoxi-

dizable substances would be liberated, and atmospheric ox;gen would pene-

rate more easily. The coloration observed might then be due to a se ondary

dation by H,0, (formed as a by-product) catalyzed by peroxidase The
substances oxidized by H.O, and peroxidase are all phe'Lo' e S " acMs
tyrosine catechol, etc.) and the oxidation products are almos always o^

catalyzed largely by peroxidase, a preliminary step being the formation of
hydrogen peroxide by a respiratory enzyme.) « formation ot

If the strongly alkaline condition near the electrode develops imme

ttir T""^' " " """"^'^ ''''' "«*^'^- '« -P--ble for the changes
in the tissue because enzymic reactions could not take place. The chanS

wS bHrue t tr''' '" f '"" '"'"'''''' "' ''' -" -terials s:d
would be true m the ease of proteins and amino acids. Proteins on hy-
drolysis yield ammo acids of which tyrosine is one of the most ^bundan^
An oxidation product of tyrosine is quinone which is of brown cot Pec
tins may also be involved; they form pectic acid and methyl alcohol 'in tht
presence of dilute alkalies. Starches, however are ve^v . fwl !v

presence of dilute alkali. Alkaline hydro^ of p tLs ^^^^^^^^^^
be responsible for the breakdown of fruit and certain vegetable ii sues but
this cannot be the cause of the change in the potato in wUch sta ch 1 ' „"
manly involved and where no discoloration takes place. We return ther "
ore to the possibility that the high alkalinity observed deTelopsouly after"
the tissue IS already partly liquefied and the current flowing stronWvTn
which ease autolysis (enzymic proteolysis) could take pla^r ^'''

SEIFRIZ: BREAKDOWN OF TISSUE DUE TO ELECTRIC CURRENT

199

Certain features of the experiments here described appear to be new,
certain others have an historical background. In 1791 Galvani^ made the
observation that experimental animals which have been previously subjected
to electrical treatment, spoil much faster than those not so treated. Later,
Humboldt^ reported some observations of Benjamin Franklin, who, he
says, was the first to remark that putrification takes place very quickly in
animals or parts of animals in which the irritability has been destroyed by
strong electrical shock or wasted by repeated contractions. Humboldt
adds that he had often made the same observation in regard to animal tissues
used in galvanic experiments. The legs of frogs to which a current had
been applied for a long time, putrefied several days sooner than those which

had not been so treated.

The observations of Galvani, Franklin, and Humboldt are other in-
stances of the acceleration of the rotting of tissues as a result of the passage
of a current. The current as such is not the primary cause of rotting in
these historical experiments any more so than in the experiments reported
here. That the current itself is not responsible is shown by the observation
of Franklin that muscle which has been wasted by repeated contractions
also putrefies very quickly just as does muscle which has been subjected to
an electric shock.

Summary

1. The tissues of fruits (apple, persimmon, etc.) and vegetables (potato,
etc.) are broken down to a soft and usually dark (brown to black) colored
mass when they are the cathode of a 110-volt direct electric current. The
anode produces little or no effect.

2. All tissues assume a remarkably uniform pH value (of 12.2) at the

cathode.

3. The primary cause of the degeneration is thought to be autolysis (pro-
teolysis) catalyzed by intracellular proteolytic enzymes, which are rendered
active by the reducing conditions existing at the cathode. Proteolysis is
(in most instances) followed by oxidation (and discoloration) which is
accelerated by the highly alkaline condition existing at the cathode. These
oxidations are catalyzed by oxidases and peroxidases and probably involve

phenolic compounds.

4. If the highly alkaline state existing at the cathode is immediately
produced, then proteolysis, which would be inhibited by so high a pH as

2 Galvani, a. Abhandlung uber die Kriifte der Elektrizitat bei der Muskelbewegung,
1791 (W. Ostwald's Klassiker der exakten Wissenschaften, no. 52, Leipzig, 1804.)

3 Humboldt, F. A. Experiences sur le Galvanisme, et en general sur 1 'irritation des
fibres musculaires et nerveuses. Paris. 1799.

200

PLANT PHYSIOLOGY

12.2, probably does not occur; in this case direct alkaline hydrolysis of pro-
teins may be responsible for the breakdown in tissue. It is believed, however
that alkalinity follows proteolytic degeneration.

5 Potato and onion tissue gelatinize at the cathode into a typical jelly
and do not become discolored (oxidized) when traversed by an electric'
current.

^ I am indebted to Dr. K. A. C. Elliott for suggestions bearing on the
interpretation of the foregoing experimental results.
University of Pennsylvania
Philadelphu, Pennsylvania

)

IRREGULAR PAGINATION

s

[Reprinted from The Geographical Review, Vol. XXVI, No. i, January, 1936.]

VEGETATION ZONES IN THE CAUCASUS

William Seifriz

University of Pennsylvania

VEGETATION zones as determined by altitude are well marked
in certain regions, especially on tropical mountains, but
elsewhere they so merge into one another or are so influenced
by exposure, precipitation, and glaciation that they cannot be sharply
delimited. The latter condition is more typical of the Caucasus.
However, altitude is always a factor to be reckoned with, and, in

8

4\ V

." ^ .. t ~

FiG i-Panorama of Tskhra-Tskharo. Minor Caucasus, showing altitudinal plant zones: i. lower
fields with wild apple and pear; 2. deciduous forest; 3. tree line of birch; 4. rhododendron thickets; 5.
high-grass subalpine fields; 6, alpine pastures.

spite of exceptions and confusions, the zone is a convenient geographic
unit by which to characterize plant distribution on mountain slopes.
The Caucasus (Fig. 2) offers every conceivable problem in plant
geography. 1 South of the main range lie the semideserts of Tiflis,
the subtropical shores of the Black Sea, the luxuriant forests and
fields of Svanetia, and, in the Minor Caucasus (or Transcaucasus),
the garden spots of Borzhom and Bakuriani and the parched hills
along the Armenian border. The north slopes of the main range, which
fall away to the flat expanses of the Russian steppes, are more uniform
among themselves but differ wholly from anything to be found on
the south side. From Mt. Kazbek on the east to Mt, Elbrus on the
west, these slopes are rocky and precipitous, with grass fields insuf-
ficient for grazing. The southern Caucasus supplies the northern
with hay; but although the south surpasses the north in wealth of
forests and pastures, the north has some of the finest natural flower

gardens in the world.

In the Caucasus there is an extraordinary variety of plants Many
of these are endemic, as is evident from the frequent occurrence of the

. For a more detailed study see the writers "Sketches of the Vegetation of ^o-^ Southern Provin-
ces of Soviet Russia.- II. 1 II. VI. and VII. Jo,m,. o/£ro/ogy. Vol. 19. 1931. pp. 372-382; Vol. 20. 1932.

PP 53-68; Vol. 23, 1935. PP- 140-160.

59

6o

THE GEOGRAPHICAL REVIEW

Specific term caiicasica in botanical nomenclature. Plants, like
people, seem to have stopped here in their migratory journeys (nine-
teen different languages are spoken). The distinctive plant life is
due primarily to the high barrier formed by the mountains between
regions of wholly different climate.

I* ■ I. ■' I )

80 MILES

80 KILOMETERS

•AO

Erzerum

6E06W REVPEW, JAW. I9M

Fig. 2— Location map of the Caucasus. Scale approximately i : 8.000.000

Vegetation Zones of the Main Range

The Georgian Military \\ ay (Fig. 2) from Ordzhonikidze (Vladi-
kavkaz) to Tiflis passes through the renowned Daryal Gorge. A
journey along this route provides an opportunity for studying the
typical vegetation zones of the main range. North of Ordzhonikidze
spread the great steppes, which, where not in cultivation, support
that most characteristic of all steppe plants, artemisia. The best-
known of the Russian artemisias is A. absinthium {A. tridentata is the
sagebrush of southwestern North America).

Where the steppes rise to meet the slopes of the lower foothills
and forestation has left open areas, a number of small plants find a
home and convert the steppe-forest transitional subzone into a
flonstically attractive region. Here, at 2400 feet, the dioecious shrub
Ilippophae rhamnoides grows in extraordinary abundance, and her-
baceous plants with showy flowers make their first appearance, though
they are more abundant in the slightly higher and cooler ravines of
the foothills.

The lowland forest, a herbaceous woodland in the main, is entered
at an altitude of 2500 feet, near the former village of Balta. The black
poplar {Populus ni^ra), monarch among the trees, though not abun-
dant, has a wide distribution in the Caucasus. Its close relative P
pyramtdalis is more numerous and gives to the landscape something

vegetation zones in the CAUCASUS

61

of the appearance of southern France. Abundant also are the willow
iSalix alba), common along the banks, the locust {Robinia pseudo-
acacia), and the wild apple {Pyrus Malus). Most prolific of all is
the hawthorn (Crataegus melanocarpa) , which grows into sturdy trees
five meters in height and is very characteristic of the foothills.

piG. 3-Tree line of birch on the moraines of the Bash-Kara (.iac.ci ... ihe main Caucasus range in
the valley of the Adyl-su.

Away from the road and well within the denser lowland forest there
occurs an association of deciduous trees consisting chiefly of the linden
{Tilia caucasica), beech {Fagus orientalis), elm (Ulmus miens), and

oak {Quercus iberica).

On emerging from the lowland forest one enters a deep valley
surrounded by massive hills (ca. 3000 feet) devoid of all forms of
arboreal vegetation. These bald mountains, some of them perfect
cones, make a striking panorama. Whether they were once forested
and later denuded (as botanists maintain) or have never been covered
with trees (as the natives believe), it is impossible to say. They
support chiefly the grasses, poa, festuca, and brome and, along the
stream banks, a wealth of richly colored flowers, including Althaea
ficifolia, A. rosea, Inula helenium. Asparagus vertictUatus, and Physalts
alkekengi. A lower herbaceous zone of this kind is found nowhere

else in the Caucasus.

Eighteen miles from Ordzhonikidze, near the village of Lars, the
upper, primarily evergreen, forest begins at an altitude of about 4000
feet. The dominant plant is a form of the European pme Pinus
svlvestris) and has been named P. caucasica and P, hamata. ^^ PJ^'
carious positions at higher altitudes, or often because of another kind
of soil, grows a scrubby form, the mountain pine, Ptniis montana
(P. svlvestris var. alpimi). The species, taxonomically doubtful, is

I

l^'

62

THE GEOGRAPHICAL REVIEW

I r, .< t.*'

V?iW ^a**"

characterized by its soil
preference, since the mon-
tana form grows on granite.
Three species of juniper
and a yew (Taxus baccata)
complete the list of conifers
along the Georgian Mili-
tary Way.

The deep Daryal Gorge
is a truly awe-inspiring
canyon. Vegetation be-
comes sparse: mountain
pines perch upon ledges
here and there, and an oc-
casional alpine flower cling-
ing to a crevice is a fore-
runner of the flora of still
higher altitudes. A trun-
cated cone of rock, crowned
by the ruins of a castle
credited to the legendary
Queen Tamara, marks the
upper, or south, end of the
gorge and the limit of the
evergreen forests. Several
species of deciduous trees,
particularly the birch {Bet-

rlo ine on Xir f T*""' '" *''' ^""'^'^^"^ '""^ pine ascends to
Strcted to ntT T I ''•'°'"''^' ^^''"'^ '''^ •^''''^h is sharply
Ind a ero, n 7 '"'''• ^" «^<^'^^'""-' "^^Ple (Acer TrauH^Ueri)

and a group of mountain poplars (Poptdus tremula), protected bv a

ormer monastery, are indications of the presence in hi zone of trees
tha n, other parts of the Caucasus are abundant and highly typical
In valleys leading off from the Gcwgian Military VvJf,; Tui

and thesmaller villageofGviletyherbacLuspt t flL i h^ulu^i'^^^^^^^
•n suba,p,„e pastures at altitudes of about 6000 feet. Fw gains
equal the profusion of color of the Caucasian subalpine vegmtion

hth :::s:zrar'th'"^^" ^-^t '': «'^-- ^^^^^^^s^

eentifns l\: r ''™""''' '^•«- ^'■'""'»'» caucasicum), poppies

The high alpine fields of the Caucasus at about 8000 feet take two
forms, pasture and tundra rhigh moor). Belonging typSly to ti:

BaTkarir^'"' ^"'""' "" «""th-facing slope at Tegenekli.

\

VEGETATION ZONES IN THE CAUCASUS

63

tundra but constituting a
community of their own
are large, scrubby patches
of rhododendrons, azaleas,
and daphnes. The alpine
pastures are much like
those of the Alps, with
similar genera dominating.
In addition to the grasses
(e.g. Poa alpina), the ger-
anium {G. pasterns) is par-
ticularly abundant in cer-
tain localities; typical also
are Alchemilla sericea, My-
osotis alpestris, and Saxi-
fraga sibirica. In moist
depressions luxuriant
growths of Crocus flavus
occur up to 10,000 feet, and
the autumn crocus {Colchi-
cum autumnale) may also
be found.

Minor (Southern)
Caucasus

The Minor Caucasus lie F'^- S— Birch forest on north-facing slope, valley of the

^ . 11 ^1 Uzengi, Balkaria.

m Georgia and along the

border of Armenia. Less rugged than the main range, they offer some
unusually delightful regions for rest and phytogeographical study. The
forests of Borzhom are renowned. At Bakuriani an exceedingly pleas-
ing valley presents many problems of ecological interest, especially that
of the distribution of the pine. The pine is sharply restricted to south-
facing slopes both here and in Balkaria (northern Caucasus). On
the knoll Mucheri, near Bakuriani, an east-west line may be drawn
down the middle of the east slope so that to the south of this line
there is an almost pure stand of pine while to the north there is but
a single pine tree.

The pine cannot stand competition but tolerates great dryness;
consequently where precipitation is low or the drying effect of the sun
pronounced it finds little competition and does well. These factors
put it on the southern slopes in the Caucasus. In Armenia, on the
other hand, it is found on northern slopes. North-facing slopes of the
Caucasus have enough moisture for all forms of vegetation ; the pine,
therefore, is driven out by competition. The south-facing slopes have

64

THE GEOGRAPHICAL REVIEW

'retoSlltlotsT ''T ''"r-^^ ^- '''- ^" Armenia
therefore h!r '^ '"'' '^"°"«'' ^"^ P'"^ but for little else-

resir.i£tr~^ - -- ^^ aao^Tiirters?;^^^^^^^^^^^

one PassesVoL itTp -stut th^ °' Tskhra-Tskharo (Fi,. ,)

through deciduous fore's "be" h li Z ^^^^^ .^'^^">' "^
(pine and spruce on south f^^.-n i ? °" ""■"'h-facing slopes

of maple and ,he^"nto tt > ^^^^ '"'"''" "PPer deciduous forest
line at 70^0 f4t S^ h J '^'^'^'?/"d --owan woods that form the tree

di 7000 teet. bubalpine fields fo bw with gentians r^^r. 1
pnmulas, geraniums, aconites, and poppies AsT. "^""P^""'^^'
proached, large thickets of rhododendron I'nH •'""""" " ^P"

more precipitous slopes just be^t he ridgr'pig^^^^^^ Tl '''

SLrwinXTi'irf xir -^^^^^£^^^^:
the delightful c::cr;:.2i:rst^:^^^^^^^^^

compact rosette of leaves, and taproot so '^ rcS.triip'i^:

Balkaria

a-piS tltLf Balkan: TFir.^ Th"' ^""^ ''°'''''-' ^^ ^"^
and the province has the W o nclud nrMrETbr^'^lH ^u'^'
summit in Europe (18 470 feetl T>rT, \ "^' ^''^ ^'^^^^^

many. The sharp restrctL of t^e '" "'""' ^'^^"''"tion are

as well defined her'e as in ^ M n'o "cau^sus" %u'''"l ''°''' '^
forested northern slones ^r^ n^, ^ u f ^^ '"'^''^ luxuriantly

birch at higher althude ." d th I is t rt o'f Th^V""" ^^'°^'^'' ^"^
So pronounced is this restriction th.tTi? ^^"'^^'"' '" «'^"^^'»'-

almost pure stands of pS o^the So, ,h ^p'™".""""'"'" "^^^ ^ave
north (Fig. 5). ^ ^^^ '°""' (f'-K- 4) and of birch on the

thei:gt;tst?trtrr,h:s.u""^^""^ '" "^'^-■■^- "-^

the alpine and subalpine oLs are h^ el'"" '" T"^ ^'^^^^^ ^'''^- ^^ •
forced down to and below'L Lf qI "'' ■' " "' '"" ''"' "
are absent the tree line ris.Z :T.,,2::Ts:^Z.'''''' ''''''''

tionirruS:,;rvrr:: ^tTb" ''t '-^^'-^ -^'■

as Krugo^or, where^neTf\he glaciers If^Elbr?" ''; ^'^T ''"°^"

VEGETATION ZONES IN THE CAUCASUS

65

Fig. 6 — Valley of the Ingur, Svanetia. Slopes covered with fir and spruce.

SVANETIA

The valley of Svanetia (Fig. 2) combines the tranquil beauty of
a New England meadow with the awe-inspiring grandeur of snow-
covered peaks. The province is part of the "Repubhc" of Georgia
and lies immediately south of the main Caucasus ridge. On entering
it from the north by the Donguz-Orun Pass, one leaves the over-
whelming rocky pinnacles and ice sheets of the northern Caucasus
and descends rapidly into a pastoral country (Fig. 6).

The forests here differ in several respects from those of the northern
Caucasus. The fir {Abies Nordmanniana) , lacking in the northern
Caucasus and scarce in the Minor Caucasus, grows to great size. With

Fig. 7 — A pure stand of Populus tremiila at Tavrari, Svanetia.

66

THE GEOCiRAPHICAL REVIEW

it occurs the spruce {Picea orientalis), which is also absent in the north-
ern Caucasus but common in the Minor Caucasus. Deciduous trees
are many; they include all the genera so far mentioned and numerous
others, notably oaks, of which there are three species. The dominant
deciduous tree of the mountain slopes is the trembling aspen (Populus
tremtila), which reaches great size and where occurring in pure stands
is a prominent feature of the landscape (Fig. 7)- The alpine zone
takes the form of pasture or tundra. In the higher valleys of the
main range north of Mestiya one finds an association of plants typical
of Arctic moors. A small community, covering not more than several
square yards, contained the following typically Arctic forms: Linnaea
borealis, Pyrola seainda, Empetrum nigrum, and the widespread moor
heath Vaccinium vitis idaea. Altitude gives these plants here in
latitude 42° the climate that they have at sea level in latitude 69°.

The Shores of the Black Sea

The coastal fringe of the Black Sea is an entirely different vegeta-
tional region. This is the Riviera of Russia. Grapes are grown ex-
tensively; ripe figs are common in the markets; the ailanthus and the
leguminous tree albizzia are reminders of the Near East. Junipers
are represented by five species, Jiiniperus excelsa being the most
abundant. Oaks {Querctis pubescens), maples (Acer campestre), the
hornbeam (Carpinus orientnlis), the ash {Fraxinus excelsior), and the
arbutus tree (Arbutus andrachne) are common where conditions are
favorable.

M-

J

THOMAS JEFFERSON»S GARDEN BOOK

RODNEY H. TRUE

(.Read April 23, 193^

Abstract

Thomas Jefferson kept a -ne^orandun, book ^^^'f^C^l':^'';:^^^:
after his return from William and Mao' College ° '^H, when he 5 y ^^^^^.^_^

This book contains jottings of interest b«>"'<^ fj '^f^^"'*"" ?„"„hSi he was interested.

they bear to matters with which Jf"-";^ ;";:": ^es are accompanied by observa-
Notes on the ornamentals and on garden ^^setaD'es , "^^^^^^^ p„ts of the

tions on the weather by informat,on on Pjantmrod^^ ^_^ ^^P^^

world, and by notes bearmg on "^"' °' "J, , V j , f upland rice to amelio-

i:-=irtr=r^l5^^^^ (3) .h: possib. ntnity o.

in relation to landscape problems, methods of work, quantiu
other practical affairs.

TT>^ rccorSv, when some years ago this manuscnpt

1 ne DooK / J through the year, while during

and are unevenly scaiiercu hh^^b j . pu;io_

ana are / Jefferson was living in Fhila-

long periods ot time, as wucn j . i i ^r „^tp«; The

nil- A \r. Pnrk there is an entire lack of notes, i ne
2 reco"d W.^:;;;et:im fro. ,766, when as a young ma
TeTaL back from William and Ma^y 1^2^^;^^^^!
Shadwell, to 1 824, two years before his death 5« X ^^^

KepnnUi iron Proceedings A,nerican PMosopHical Society.
^ Vol Ixxvt., No. 6, 1936

Printed in U. S. \.

940

RODNEY H. TRUE

The notes throughout are in Jefferson's own handwriting,
and offer an interesting series of samples showing the changes
with the years.

The subjects seeming to him worth noting are usually
commonplace, and the entries probably grew in some measure
out of the note-taking habit of a man industrious with his pen
throughout his active life. At times, however, observations
are included that hint at other and more significant things.
His first annotation is one of the less usual observations bear-
ing on the esthetic interest of the subject. On March 30,
1766, he writes: "Purple hyacinth begins to bloom." During
the following spring months, narcissus, puckoon, the purple
flag, violets, and honeysuckles, are noted, as well as "a bluish
colored, funnel-formed flower in low grounds." He notes the
times of coming into bloom, and the end of the flowering
periods. These wild plants here share interest with garden
varieties.

Notes made in 1767 are more numerous and diversified.
A large number of sorts of old-fashioned flowers are noted,
usually with the time of blooming, but the vegetable garden
claims more lines than the ornamentals. In February, he Is
sowing two kinds of garden peas, the progress of which is
followed until the earlier lot "comes to the table" on April
24th. Apparently, while dealing with the pea seeds to be
planted, a curious and steadily present inclination to statis-
tical precision first appears. A line records that " 500 of these
peas weighed 3 ozs., 18 pwt., about 2500 fill a pint." This
type of accurate dealing crops out often and records, appar-
ently for later reference, facts that may be useful in future
planning. One can here learn how long it took Jefferson's
seeds of celery, asparagus, peas, Spanish onions, and lettuce,
to come up and to become useful.

The flowering times of Sweet William, lunaria, snapdragon,
lychnis, larkspur and poppies, are recorded, facts of interest
to one designing a flower garden. He looks further ahead and
plants trees and shrubs, lilacs, roses, laurel, Spanish broom,
plums and gooseberries, almonds and altheas.

^»»

THOMAS JEFFERSON'S GARDEN BOOK 941

Strawberries planted in 1766 now bear for the first time,
and again, Jefferson becomes statistical. "The plants bear
2 o. strawberries each, 100 fill half a pint." This bit of infor-
mation shows how this fruit has been increased in size through
the skill of gardeners since his time.

The book now begins to be a repository for occasional
records of importance in the economy of a plantation; e.g.,
"8 or ID bundles of fodder (corn RHT) are as much as a horse
will generally eat through the night; 9 bundles X 130 days
= 1 170 for the winter."

In 1768, his records are brief and confined to the spring
months. Again peas are counted, Charlton Hotspur—" 500
of these peas weighed 3 ozs., 7 pwt., 2000 filled a pint ac-
curately."

With 1769 his notes begin to concern Monticello entirely,
his new home growing on the height above Charlottesville.
With March 14th, planting begins, trees and shrubs receiving
exclusive attention; pears, one row of them grafted, cherries.
New York apples, peach stocks, nectarines, quinces, pome-
granates, figs, walnuts and apricots, being enumerated.

Notes on the quantity of lime to lay 2000 bricks indicate
the beginning of construction, either on the house or walls
about the place. He notes what Nicholas Meriweather says
about the area needed to produce watermelons enough for a
family " not very large." He quotes Miller's Gardener's Dic-
tionary on the yielding capacity of 50 hills of cucumbers,
" I hill yields 40 cucumbers."

In 1771, notes on weather conditions begin to enter, and
record items that seem to sound like the weather reports of

today.

I have quoted these items to show the character of much
of the content of this Garden Book, and rather commonplace
the material seems to be. However, certain entries are con-
nected with matters not mentioned, and hint at subjects of

wider interest.

In 1771, Jefferson makes a note of shrubs not exceeding
10 feet in height, in great part native plants. Another list

IRREGULAR PAGINATION

942

RODNEY H. TRUE

of trees is recorded, in part native, including dogwood, red
birch, catalpa, magnolia, mulberry and locust among the num-
ber. Other lists of climbing shrubs, evergreens and hardy
perennials follow. This seems to be a mustering of materials
to be considered in his plans for planting his grounds.

Next year, he married Martha Skelton and brought her
home to the partly finished house and unfinished surroundings.

In that year, 1772, there was much activity on the little
mountain. Earth was being moved, walls were being laid,
and the new road up the mountain was being built. Here
again, the statistical method applied to the task suggests the
modern efficiency expert. "Julius Shard fills the two-wheeled
barrow in 3 minutes and carries it 30 yards in i§ minutes
more. Now this is four loads of the common barrow with
one wheel. So that suppose the 4 loads put in, in the same
time, viz., 3 minutes, 4 trips will take 4 X i^ minutes — 6,
which added to 3 filling is — 9 to fill and carry the same earth
which was filled and carried in the two-wheeled barrow in
4§ minutes. From a trial I made with the same two-wheeled
barrow, I found that a man would dig and carry to the dis-
tance of 50 yards, 5 cubical yards of earth in a day of 12 hours'
length. Ford's Phill did it, not overlooked, and having to
mount his loaded barrow up a bank 2 feet high and tolerably
steep." This sample may illustrate the type of observation
and annotation frequently occurring before 1809. After that
date, the material is generally compactly tabulated, and ex-
tended notes are unusual.

In 1773, he continues to plant fruit trees, while grape vines
and a few notes on wine appear.

An interesting light is thrown on the domestic arrange-
ments on a large plantation by a list of "articles for contracts
with overseers."

"He (the overseer RHT) shall let his employer have his share of
grain if he chooses it at a fixed rate.

"He shall not have his share until enough is taken out to sow,
and then only of what is sold or eaten by measure.

"Allow }/2 a share for every horse and same for a plough-boy.

THOMAS JEFFERSON'S GARDEN BOOK 943

"To have at the rate of a share for every 8 hands, but never to
have more than 2 shares if there be ever so many hands.

"Provision 400 lbs. pork if single, 500 lbs. if married."

"To be turned off at any time of the year if employer disapproves
of his conduct, on paying a proportion of what shall be made,
according to the time he has staid.

"To pay for carrying his share of the crop to market.

"To pay for carnage of all refused tobo (tobacco RHT).

"To pay his own levies.

"To pay his share of liquor and hiring at harvest and never

bleed a negro."

In 1774, the book begins to show traces of an activity
that grew with Jefferson and became a matter of public inter-
est. His Garden Book shows a long series of entries in Italian,
apparently growing out of a shipment of garden seeds from
Mazzei, who had gone to Europe on a governmental errand.
This list was a long one, and introduced a variety of onions,
endive, parsley, spinach, broccoli, radishes, bush fruits, mel-
ons, cantaloupes, squashes and grapes. The grapes were
planted by "some Tuscan vignerons who came over with

Mr. Mazzei."

Jefferson's interest in grape growing and in wine making
as an American industry is here indicated. That interest
grew as the devastating effects of drinks having a high alco-
holic content were realized. He looked on the domestic
making of pure wine for family consumption or the use of
cheap imported French wines as temperance measures. His
agreement with Timothy Pickering on these matters throws
an interesting light on Jefferson's concern for genuine public
welfare and his idea of the type of remedy to be applied.

After recording in full how the Tuscan vignerons planted
the grapes, he records unusual weather conditions, effects of
the destructive frosts of May being given in some detail. ^

In 1775, again he records destructive frosts in the spring,
but little else is recorded.

No records appear in 1776, for reasons that are easily

understood.

I will pass over the remaining years, in which the usual

V

944

RODNEY H. TRUE

types of items are found. To the plant student, much of
interest is to be found in Jefferson's notes. The dates of the
appearance of a great variety of imported plants and trees
are historically interesting, the result of Jefferson's desire to
introduce the best products of the Old World into the young
country. He was equally concerned to see what might be
done with the native plants.

His efforts to establish the growing of upland rice in Amer-
ica grew out of his realization of the loss of health and of life
in the malarial rice fields of Carolina. He had ship captains
sailing to other lands enlisted as observers and agents. Seeds
of upland rice were brought from the Pacific islands and from
Asia, and he himself visited the Italian regions growing dry
rice, and started an active local cultivation from a little of the
seed that he brought home in his pocket. Upland rice did
not displace that of the wet lands until long after Jefferson's
day, but small areas of these early sorts are still grown in the
Georgia mountains.

The Garden Book records: "May 7, 18 10, sowed upland
rice at the mouth of the meadow branch."

In 1794, Jefferson was planting sugar maples at Monticello
in the hope of using them as a source of sweetening. He
frequently urged the possible value of this tree for this pur-
pose. In the same year he was planting pecans, that later he
seems to have distributed to Washington, Judge Duval, and
others. He was again trying to develop the use of a native
product.

In 1794, he was quoting temperatures observed on Doctor
Walker's thermometer, he himself lacking one. By 1803, he
seems to have supplied the missing instrument, since he is
quoting temperatures freely.

I know of no earlier planting of a nursery of forest trees in
this country than that of Jefferson on April 12, 1804, when he
planted hemlock and white pine "near the aspen thicket."
Grapes, pecans, and fruit trees continue to furnish main items
at this period, with minor plantings of a great number of
garden plants from foreign lands. Monticello had become an

J

I*-

THOMAS JEFFERSON'S GARDEN BOOK 945

actively operated experiment station, where plants from all
corners of the earth were planted and tested.

References in the records of the Lewis & Clark Expedition
to seeds brought back from the west gave rise to a search into
the subsequent history of those seeds. The results of that
search were reported some years ago to this Society. The
Garden Book shows that a few of these seeds were taken to
Monticello, where Jefferson put them into his experimental
garden. " Pani corn, " noted in 1 8 1 1 , was planted with native
Cherokee corn, with Quarantine corn from his friend Thouin
at the head of the Jardin des Plantes in Paris. "Columbian
salsify," one of the Lewis & Clark plants of the far west, having
a fleshy root, was grown and tested on the table at Monticello.
The verdict was adverse.

In 181 2, more of the Lewis & Clark material appeared at
Monticello. Red gooseberry, Lewis' sweet-scented currant,
Lewis' snowberry bush, and Lewis' yellow currant indicate the
propagation of a group of ornamental shrubs, several of which
in time took an established place in ornamental planting.

He sums up his long experience with plants at Monticello
in 1823, as his life neared its close, in a condensed table of
what should be done month by month. He calls it "Com-
pend of a Calendar."

As time passes, and the nation goes through its strenuous
earlier years, the Garden Book of Jefferson continues to tell
a quiet story of grapes and sea kale for the garden, of carp and
chub for the fishpond, of plums and peaches, peas, radishes
and squashes, and one can understand his longing to leave the
man-made tumult of Philadelphia and Washington. The
worn Garden Book seems to give a glimpse into the back-
ground of the life of one of the most significant and storm-
tossed characters in our history, and in this close contact with
plants and with the earth, we seem to find one of the real
sources of his strength.

IRREGULAR PAGINATION

Reprinted from the Brooklyn Botanic Garden Memoirs, Volume IV,

published May 7, 1936.
Printed in U. S. A.

TWENTY-FIVE YEARS OF PLANT PHYSIOLOGY, 1910-1935

1

rv

i

Rodney H. True

Professor of Botany and Director of the Botanic Garden,
University of Pennsylvania

Two persons have an element of clanger attached to their utterances — ^the
historian and the prophet. The historian may meet objection from those who
weigh occurrences of the past in a different balance from that used by himself.
The prophet is in an even more dangerous position, because the hold of the
future on the past is none too stable a one, and the future sometimes breaks
sharply with the past. However, while this applies to the world of politics,
as we have seen, it would seem less likely to happen in the world of science.
Still, it is possible even here, because one hardly knows what the development
of the sister sciences may bring by way of new viewpoints and new methods.

In the hope of getting a starting point, I would like to review in an outline
way the situation with Plant Physiology twenty-five years ago.

Twenty-five years ago, plant physiology was one of the younger children
of the botanical family, and it existed rather independently of the older mem-
bers— taxonomy and anatomy. These separate branches made relatively little
use of each other, but during the past twenty-five years, while individual
botanists have specialized and drifted apart until almost out of touch with
each other at times, the subject of Botany as a whole has been slowly develop-
ing a unity through the recognition of mutually helpful relations.

By the methods of physiology a complete and coherent system of plant
relationships has been built up by Metz and his fellow-workers at Konigsberg.
Taxonomy and physiology have thus come closer together, and since the re-
sults of taxonomy based on physiology agree closely with the scheme of rela-
tionships worked out on structural relations, these latter are seen also to be
involved in the synthesis.

Formerly, the plant pathologist studied his disease-producing organisms
quite apart from the reaction that the parasitized host might develop, and it
was not until the chief aspects of the physiology of the parasite had become
fairly well known that the host reaction — the physiology of the thing para-
sitized— has come to attract attention. The relation now between physiology
and pathology is very close. The host reaction is being studied and pre-
disposing causes are being sought in the environment through the physiologi-
st

82

BROOKLYN BOTANIC GARDEN MEMOIRS

TRUE TWENTY-FIVE YEARS OF PLANT PHYSIOLOGY

83

cal conduct of the host under given circumstances. More and more, a physi-
ological basis is recognized as necessary to a sound pathology.

Of course, I hardly need mention the fact that agriculture and all practical
aspects of plant cultivation have long been known to reflect the great facts of
physiology, and it was not by accident that physiology made its first great
gams m mstitutions investigating the problems of agriculture.

When the Brooklyn Botanic Garden was established, plant physiology had
already come out of the rather larval condition which characterized its earlier
development in America. During the preceding twenty-five years— 1885 to
1910— plant physiology was already receiving recognition in a number of the
more promment American universities. However, it diflFered from other
phases of botany in America in that it had a rather more direct relation with
Europe, particularly Germany, than other phases of the science had A few
Americans, studying with Sachs and PfefTer, had brought to this country
much of the German methods and some of the German viewpoints and
American physiology twenty-five years ago was continuing to build directly
on the foundations laid in Europe.

Let me list some of the men and the problems in which conspicuous prog-
ress was being made in a period centering around 1910.

PHOTOSYNTHESIS

Perhaps one of the most important things was the work of Willstatter
and his associates on chlorophyll. Chlorophyll had already been investigated
from the standpoint of its general methods of functioning, but the chemical
make-up of chlorophyll itself still remained uncertain. Willstatter, associated
with Stoll and other of his contemporaries, put the chemical understanding of
chlorophyll on a sound basis at this period. The knowledge of the action of
chlorophyll was advanced by Lubimenko and Stahl, who studied it mainly
from the ecological standpoint. Along with the clearing up of chlorophyll
chemistry, conspicuous progress had already been made in the investigation of
the physiological processes dependent on chlorophyll. Bayer had advanced
his formaldehyde-condensation theory of the origin of glucose in 1864
Brown and Morris in 1893 had claimed priority for cane sugar as the first
carbohydrate; Dixon and Mason in 1916 were proposing an alternative.
Some doubt still remained as to what the first carbohydrate found in plants
might be. *

Twenty-five years ago the presence of formaldehyde in the green parts of
plants was receiving close attention from Schryver in America and from
Curtius, Franzen and Grafe in Germany.

x

' '

I •

•> I « *

Other plant pigments were also receiving more thorough attention than
before. Escher, working on carotin and lycopin, Hanson studying phyco-
erythrin, Molisch investigating purple bacteria, illustrated this greater interest
in pigments.

CHEMOSYNTHESIS

Along with the work on chlorophyll and its significance for green plants,
we may recall that chemosynthesis, developed in various of the minor or-
ganisms, was also receiving attention. The work of Molisch on sulphur and
iron bacteria, the work of Lieske on iron bacteria, and Nikelewski's work on
hydrogen oxidation, would illustrate this phase of plant physiology.

SOIL PHYSIOLOGY

Twenty-five years ago, the soil was being investigated from a new view-
point. This time the work was done chiefly in America, at the Bureau of
Soils in Washington, under the leadership of Milton Whitney. The theory
of toxic substances in the soil as a cause of a lack of fertility was being ad-
vanced through the studies of Schreiner, Skinner and Shorey from the
chemical standpoint, and by Livington from the physiological angle. To this
body of work belongs also Cameron's study of the soil solution. This prob-
lem still remains in the region of debate, and is likely to receive further study.

MINERAL NUTRITION

The work on mineral nutrition, started in the laboratory of Sachs when he
was a young man, advanced but little after his time. Of course, Knop's and
Pfeflfer's solutions and other salt balances had been used and the theory de-
veloped that only a small and definite number of inorganic constituents were
required for the successful production of green plants. The function of the
diflFerent elements was surmised in a general way. Potassium and nitrogen
were associated with the increase in volume ; iron was known to be necessary
for chlorophyll formation, and the presence of magnesium in the chlorophyll
molecule had been demonstrated by Willstatter.

The great influence of calcium on cell permeability and on the absorption
of other ions of the nutrient medium was being investigated by the use of
conductivity methods, first in America by True and Briggs and Bartlett and
Osterhout, and in England by Stiles and Jergensen.

The main facts of toxicity and of antagonism were being studied by the
same investigators following the pioneer work of Kahlenberg and True,
through whose study the significance of the ion was shown to be general in
the physiology of plants and of animals.

84

BROOKLYN BOTANIC GARDEN MEMOIRS

In another part of the field of plant physiology, the exact methods of the
physicist were yielding important results in the studies of osmosis by Morse
and his associates at Johns Hopkins, and by Berkeley and Hartley in Eng-
land, and by Renner in Germany. The study of the lowering of the freezmg
point of plant saps by Dixon and Atkins was supplemented by the very in-
tensive work carried on in America by Harris and his associates.

The very important work of Ursprung on suction tension was probably
begun at this period, but did not come to full consideration until several years
after the date of the present cross section.

WATER PHYSIOLOGY

The water question was receiving much attention. Chemically, a defi-
nitely important advance was made by the work of Babcock on the water of
metabolism and its relation to the processes of nutrition. The water require-
ments of plants were being studied eiTectively and extensively by Livingston
and by Briggs and Shantz. Out of these studies came light on wilting and
other consequences of water shortage. The question of the course of water
up the stem and out through the leaves was being approached from several
angles. Renner published on the physics of transpiration in deser plan s,
while Lloyd was investigating the relation of stomatal movements to the prob-
lem Sir Francis Darwin was contributing to the study of methods. The
veteran Schwendender made what was perhaps his last contribution to plant
physiology in a paper on transpiration. The work of H. H. Dixon seems to
liave done much to co-ordinate the various facts known regarding the ascent
of sap and the process of transpiration. It seems likely, however, that this
is not a final statement. Hasselbring, a little later, was investigating the
relation of salt absorption to transpiration.

PROTEINS

The deeper things of plant nutrition were being attacked from the chemi-
cal vantage ground furnished by the fundamental work of Emil Fischer on
protein chemistry that had just preceded our period. The work by Fischer
was most effectively followed up by his student Abderhalden in his studies on
the animal side of the problem. The study of vegetable proteins was being
advanced notably in America by Thomas Osborne. ,„,.,, ,

Asparagin physiology was being studied in Russia by Butkewitsch and
bv Prianischnikow and his co-workers. Walters, Krasnoselsky Maximow,
and others, followed the earlier work of Treub, whose chief contribution was
an attempt to relate HCN to protein formation.

TRUE TWENTY-FIVE YEARS OF PLANT PHYSIOLOGY

LIPOIDS

85

>> ♦

^ •

■(■

Perhaps here should be mentioned the subject of lipoid physiology, devel-
oped in this period chiefly by Ernst Overton, Bang, Palladin, Rosenbloom
and others.

ADVANCES IN IRRITABILITY

During the past twenty-five years the relations of plant physiology to
chemistry have multiplied and expanded until now a textbook on this subject
would show little space devoted to other aspects of the subject, including
irritability. This fact is perhaps due to the tremendous progress that chem-
istry has made during this period in such ways as have furnished methods
available in the exact investigation of physiological problems. There has been
no corresponding advance in methods of studying irritability. However, one
conspicuous advance should be noted in this field. In the absence of anything
more definite the transmission of stimuli has been assigned to the mysterious
conducting powers of living protoplasm, it being perhaps assumed that some-
thing like a rudimentary nerve action might be here involved. However, the
earlier work of Julius von Sachs contained the germ of later developments.
In his assumption of organ-forming substances, he laid the basis for the hor-
mone theory. This has been developed by Went and other investigators,
chiefly Dutch, into a very interesting and fairly satisfactory explanation of
stimulus conduction. Whether in other works of the early masters fertile
germs still persist which may develop in this field, can hardly be foretold.

Turning toward the outlook for future development in plant physiology,
the prospect for another era of progress seems bright. This opinion is based
on the great advances being made by physics in the study of aspects of energy
revealed in the recent work of physicists, chemists and astronomers. What
the quantum theory may mean to plant physiology, as physiologists get into
working relations with it, remains to be seen. Perhaps as work in energetics
goes further, the old suggestion of Arthus may be found to have something
of a basis. He contended that enzymes owe their marvelous ability, not to
their chemical nature, but to the way in which the material is energized.

One of the greatest gains of plant physiology is the growing recognition
that almost imponderable quantities of certain substances are found to have a
very great influence on plant life. It seems likely that the ideal culture solu-
tion will eventually contain not merely the seven ions conventionally sup-
posed to be necessary, but also a host of others in minute traces that in their
own peculiar and fitting way influence the process of life. This great effec-
tiveness of imponderables calls loudly for methods of determining the pres-
ence of substances in minute traces. Perhaps the magneto-optical analytical

S6

BROOKLYN BOTANIC GARDEN MEMOIRS

methods of Alhson may be so improved as to yield results free from an
ob,ect,o„able subject.vity, and when minima can be recorded by some non-
psychological process, .t is quite possible that a door may open to an entirely
new part of the realm of plant nutrition. At present, problems of perm abt

^ ^^:^::l:r'"" -''-' -'-''-'' "^^*°^^ ^- •■- ■•^-'«-'-

chemistrv"il"'"' "'"',"' T '"'""'"^ '"'" " "'"^ ''^' ^ ^^' =»= Physics and

in th e th n e^? ' "' " " "'""' ""="" *''=" ^"^ •"='i°^ deviopments

stered t'o hv Z "T""'"' ^T ^'''^ ^'"'' '" **= P^""^'^"' '<^i«"c« min-

istered to by them, and one of the chief of these will be the field of plant

physiology. It seems increasingly certain that the physiologist of the future

.Try and CVl .T"' "1'/"'"^'^ '" '""'^"''^ "^ ^°"^ P"^^^- -^ cS!
.stry and he w.ll be st.ll more hkely to advance the science if he has command
of the abstract methods of mathematics tommana

reJe^.eZTT°! ^'^T""" '° ''" ^°'' ^^'^^ ^"""'''^^ "^ -" *e sciences and
represents the high goal toward which all sciences strive.

W

i

I

t#,ft

I

1

1

s

I

DISTRIBUTION OF THE
NORTH AMERICAN PITCHERPLANTS

BY EDGAR T. WHERRY

Associatt Proftssor of Botany, Univirsity of Pttmsylvania

The pitcherplant family [Sanacniaaac) consists of three genera, Sarracmia, Chrysamphora,
and miamphora. Nine species of Sanacma have been recognized thus far, one of them rang-
ing from the Gulf Oast far north into Canada, the others restriaed to the southeastern
United States. The genus Chrysamphora, also known as Darlinffonia, is monotyp.c its smgle
species occurring in northern California and southwestern Oregoa One spec.es of Hcham-
La was described in 1840 and for ninety years remained the only known represenm.ve
of the genus, but three more have now been found. These are limited to northern South
Ameria, however, and wiU not be discussed in this article.

KEY TO THE NORTH AMERICAN PITCHERPLANTS

Pitcher hood bearing a fish.ail-shaped appendage; scape bracted; petals bronzy yellow style
ricciici iiuwM 6 Chrysamphora caltfomtca

Pi::Jerhood not' appendaged^scape^ naked; style nonnally expanded into an -bre.h-shaped

Structure

Pitchers erect or essentially so.

Pitcher orifice incompletely covered by hood.

Margins of the large hood more or less reflexed.

Hood yellow-green, veined or suffused with red.
Petal-color of a yellow type.

Reficxing of hood margins slight; petal texture firm.

Flat leaves abundant; petals greenish yellow . . S. oreophtla

Flat leaves sparse; petals creamy yellow ^ ^^'^i"

Refiexing conspicuous; petals delicate, yellow S. flava

r i . . S. ionesii

Petal-color of a red type j-

Hood white, green- and red-veined; petals red T^ZL

Margins of the small hood not reflexed; petals red s Z*»^

Pitcher orifice well covered by hood; petals yellow ^^^""^

Pitchers decumbent; petals red or exceptionally yellow. ^ .^^

Orificelateral, small; hood closed

Orifice terminal, large, hood open. ^ ^^^^^^

Pkcheroutlme short and broad s purpurea g^osa

Pitcher outline long and narrow ^ '^

Data as to the relationships and distribution of the species have been obtained mairdy
from studies in the herbaria of the Academy of Natural Sciences o^P^^^P^-f-^^
National Museum, CorneU University, Gray Herbarium. New York Botanical Garden,

1

1

•l

DISTRIBUTION OF THE
NORTH AMERICAN PITCHERPLANTS

BY EDGAR T. WHERRY

Assoclau Proj,s,cr oj Botany, Unmruty of Pcmsylvama
The pitchcrplant famUy iSarrace«.a«ae) consists of thr^ genera ^-^ 'J^^^^^I

S:: SJ^. ?"sr X.^.-. a. -own as O..^ -°--;--^

rheTnt^^nr r Ic nL ^: .una. These a. ....cea . no«he. Soueh
America, however, and wiU not be discussed in this article.

KEY TO THE NORTH AMERICAN PITCHERPLANTS

PUch. hood bearing a fish.aU-shapcd appendage; scape bracced. ^^-^^^J^-^f::^!:^

radiate ! , • ^.^oiw ^vnanded into an umbrella-shaped

Pitcher hood not appendaged; scape naked; style normally expanded mto ^^^^^^^^

Structure

Pitchers erect or essentially so.

Pitcher orifice incompletely covered by hood.

Margins of the large hood more or less reflexed.

Hood yellow-green, veined or suffused with red.
Petal-color of a yellow type.

Reflexing of hood margins slight; petal texture firm.
Flat leaves abundant; petals greenish yellow
Flat leaves sparse; petals creamy yellow . .
Reflexing conspicuous ; petals delicate, yellow . .

Petal-color of a red type

Hood white, green- and red-veined; petals red . • • •

Margins of the small hood not refiexed; petals red ....

Pitcher orifice well covered by hood; petals yellow

Pitchers decumbent; petals red or exceptionally yellow.

Orifice lateral, small; hood closed

Orifice terminal, large, hood open.

Pitcher outline short and broad

Pitcher outline long and narrow

. S. oreophila
. . S. sledgei
. S. flava
. S. jonesii
S. drummondii
. S. rubra
. S. minor

. 5". psittacina

S. purpurea venosa
S. purpurea gibbosa

INTENTIONAL SECOND EXPOSURE

IPPPriTTT AP PAnTMATTONT

United States National Museum, and University of Pennsylvania, and from the following
publications:

Sarraccniaccae, by John M. Macfarlane. Engler's Pflanzcnreich, vol. iv. no, 1908.

The American pitcher-plants, by Roland M. Harper. Journal of the Elisha Mitchell Scientific

Society, vol. 34, p. 110, 1918.
The biochemistry of the American pitcher-plants, by Joseph S. Hepburn, Frank M. Jones, and

Elizabeth Q. St. John. Transactions of the Wagner Free Institute of Science, vol. 11, 192.7.

Four articles relating to pitcherplants have previously been published by the writer:

Acidity relations of the Sarracenias. Journal of the Washington Academy of Sciences, vol. 19,
p. 379, 192.9. (The species ranges given in that paper are somewhat modified in the present
one in accordance with data subsequently obtained.)

The geographic relations of Sarracenia purpurea. The Appalachian relative of Sarracenia flava.
Bartonia, vol. 15, p. i, 1933.

Exploring for plants in the Southeastern States. Scientific Monthly, vol. 38, p. 80, 1934.

As pointed out in the paper by the writer on Sarracenia purpurea^ some recently pub-
lished maps of the distribution of the members of the Sarraceniaceae (Die Pflanzenarealc,
vol. 3, pt. I, 193 1 ) are not satisfactory, having been constructed from incomplete data for
most of the species. A new set of maps is accordingly presented herewith, based on a thor-
ough review of the literature, a cataloging of numerous herbarium records, and extensive
field observations.

The base map shows, in addition to State boundaries, two geologic lines of plant-geo-
graphic significance. The more northern of these represents the limit reached by the last, or
Wisconsin, ice sheet, as mapped by Antevs (Bulletin of the Geological Society of America,
vol. 40, p. 63 1, 192.9). The southern line is the fall line, taken from the map of Physiographic
Divisions of the United States, published by Fenneman (Annals of the Association of American
Geographers, vol. 6, p. 19, 19 17). Most of the territory enclosed between these lines has been
continuously available for occupancy by plants since Cretaceous time, when the development
of our modem flora began.

On the individual maps, areas where the species is frequent in favorable habitats are
dotted. When the boundaries have been fairly definitely determined, they are marked by
solid lines, and when only approximately known, by dash lines. Presumable migration routes
are indicated by arrows.

Before the individual species are discussed, the geologic relations of the group as a whole
require brief consideration. Physiographic studies indicate that prior to and during the Creta-
ceous much of eastern North America was reduced to a peneplain. The sea then extended
up to what is now the fall line, and for some distance northward and westward from this
boundary the conditions for plant growth were similar to those of our present-day Coastal
Plain. There were vast level areas covered by alluvial sands and clays, traversed by sluggish,

meandering streams, and dotted with bogs and swamp. This was the ancestral home of

''i;^:t^:l^ ehe land too. place, and et^ion develop the ptese.

topTr phy Some of 7he plants adapted themselves to the cxx,ler climate of -°- f ^'^
X nd grew there throughout the course of the development of our Blue lUdge and
ZaLchian Mounta.ns. oThers proved unable to adjust their temperature reqmremen.
fnd'ti out from their ancestral regions. MeanwhUe. however, the sea was gradually
creating leaving land open to occupation by plants, and seeds or other d^mrnules of
r st^rfound their^way down various river vaUeys and developed colomes on the
Twy formed Coastal Plain. Most of the Sarracenias evidently migrated m th. manner.

CALIFORNIA PITCHERPLANT

Chtysamfhora califomica (Torrcy) Greene

Though currently believed to be restricted to a few mountain bogs in Olifornia and

Though ^""^" y ^' J^^^ ^^ „f herbarium records has revealed that ahforma

southern Oregon, a study of the 1. e^tu ^^^^ .^ ^^^^^ ^^

nirrherDlant erowsm numerous localities tromriaceri^uiiiy, .■:,.,-

pitcnerpiant grow r«„nr„ Oregon where the wnter has coUected it at

Kid .«! .h= «a «mp.™" F*"* •"« '"^ '5' f'!""'";: 1^

T-. f" -"t -^t "^ :^.tJ:ZJ^!l'Z^^:^^

T«m, .iml « b, *= «l™" of "» " '"^ """^ «'""' "^

GREEN PITCHERPLANT

Sanacmia oreophila (Kearney) Wherry
r r^ nitcherolant the most primitive of the eastern sj^ies, was apparently first coUeaed
J^^c^tX^f the cLtal p.^ .n Taylor County G-gi.^-^!- P^
tLr chiefly in the Appalach^n Mou„«ins ^^^^^^^^^^^ ^^^ 2^
found thus far in a number of stream vaU^s m ^''^-''^e^^t s;iX«'ws in allu-
Counties. and may be considerably more widespread* Unhke ^'J^^^
vial sands and gravels on stream banks, rather than m bogs or swamp. The soU on.

however, is rather intensely acid. „™,(,h above sea level for

province only after it had become too conservative to spread ther^

in Elmore County, in central Alabama.

FIGURE I

Distribution of green pitcherplant, Sarracenia oreophila

5

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II

H

PALE PITCHERPLANT

Sarracenia sledgei Macfarlane

The ancestral home of pale pitcherplant was presumably in what is now the Cumb.^
land Plateau of Tennessee, where one or two colomes are report«l »;™ ^'^^^
down the Tenner and the Mississippi River systems, after they developed. Becoming
SbnizS ol the Coastal Plain, .t then spread eastward into Alabart^ there approaching
but apparently not mterminghng with its eastern relative Sanacemaflava.

It 1 sprLd westward across Louisiana and is the only spec^nown to have r^chd^
Texa. The westernmost station from which a specimen has been seen >s n^ A he^
Henderson County, nearly at longitude 96' W. There are credible records of this pitcher
olTnt fZ alC distanL farther west, but, growing as it does m acid swamps and springy
Lelt'ci; unable to enter the more arid ^rtions of Texas, and reports of its occurrence

it prlbly grows farther north in the Mississippi VaUey and has merely faded to be
collected by the few botanists who have explored that regioa

FIGURE 1

Distribution of pale pitcherplant, Sarracenia sledgei

YELLOW PITCHERPLANT

Sarracenia flava Linnaeus

The yeUow pitcherpknt developed farther east than its telative.the pale pitcherplant and
in the course of the Tertiary uplift and e.«ion which resulted in the development of the
southern highlands it managed to survive in a number of places in North arohna. As the sea
gradually withdrew from the old shore line, leaving new land open for occupation by plant^
fhe seeds of this species traveled down various river systems in North and South Carolma and
Georgia, and starS^lonies in the Coastal Plain. Uteral spreading from these colonies abo
occuL, and the species reached on the one hand, nearly to the Alabama River, and on the

Other to the James River in Virginia. , „ ,

Like most plants with such a geologic history, this pitcherplant gradually lost its aggres-
siveness, and by middle Tertiary time its colonies apparently ceased to expand further.
Accolgly. alt'hough it grows in abundant in moist meadows and dep^io^ m Poland,
it has never been able to enter lower p«iinsular Florida, which emerged from the sea only
toward the close of the Tertiary.

FIGURE 3

Distribution of yellow pitcherplant, Sarracenia fl.

ava

8

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FIGURE 4

Distribution of red pitchcrplant, Sarracenia jonesii

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RED PITCHERPLANT

Sarracenia jonesii Wherry

occurred, seeds reached the neaawatc _ , , , • , ^ Abhama Piedmont province and,

f.«he. down .^^. le ^ J^J^^l^^t I ^\t^^^^^^^ W uL sp.ead.g
when the Coastal Plain emerged from 'he^'^^" ^ ^^, J jj^^n^ i„,o Florida and

Sit™. r-"^»^»- - --* - "" *' " -"" " ""^

further.

II

4

♦? 1 >

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WHITETOP PITCHERPLANT

Sarracenia drummondii Croom

The evolutionary changes which resulted in the development of the showy red-flowered
Sarracenia jonesii from a rather inconspicuous green-flowered ancestor did not come to an
end with that species, but continued along several lines. The tendency toward increased size
and coloration reached a culmination in whitetop pitcherplant, the showiest of our species.
The common name selected for it refers to the predominance of white in the hood.

Development of this pitcherplant evidently took place somewhere in the headwaters of
the Alabama River system. The Tertiary mountain-making exterminated it from its ancestral
home, but seeds traveled downstream and soon colonized the Coastal Plain. Spreading laterally
from this river valley, it migrated a short distance westward into Mississippi and somewhat
farther toward the east, into the western extension of Florida. It also formed two isolated
colonies, one in Madison County, Florida, the other in Sumter County, Georgia. Reports of
its occurrence farther northeast seem to be based on misidentification of other species.

FIGURE 5

Distribution of whitetop pitcherplant, Sarracenia drummondii

12.

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SWEET PITCHERPLANT

Sarracenia rubra Walter

As shown by the accompanying map, sweet pitcherplant occurs mostly near the fall line
in Georgia and South Carolina, although in North Carolina it extends to the coast. Its north-
ernmost known station is at Southern Pines, in Moore County. There is also an apparently

isolated area in western Florida.

This distribution indicates that the species originated, as a descendant of Sarracenia jonesii,
somewhere on the headwaters of the Santee River system. Being very sensitive to cold, it
was exterminated in its ancestral home by the climatic changes accompanying the Tertiary
uplift, but seeds meanwhile drifted downstream and colonized the Coastal Plain. Its devel-
opment chiefly near the fall line indicates its early arrival after the retreat of the sea, but it
soon lost its aggressiveness and only locaUy reached the outer part of the Coastal Plain.

The favorite habitat of the sweet pitcherplant is a moist, grassy thicket near the margin
of a swamp, although it can grow also in dense shade. The soU is usuaUy peaty and in-
tensely acid.

FIGURE 6

Distribution of sweet pitcherplant, Sarracenia rubra

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14

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FIOURB 7

Distribution of hooded pitchcrplant, Sarracenia minor

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I

HOODED PITCHERPLANT

Sarracenia minor Walter

Peninsular Florida emerged from the sea only toward the close of the Tertiary age, and
as by that time most of the Sarracenias had apparently lost their ability to colonize new
territory, they are not to be looked for in that region. The one exception to this rule is the
hooded pitcherplant, which extends far southward over the State, being reported even in
Palm Beach County, at latitude 2.6'' N. In other directions its range is more restricted, how-
ever; it is the only species which has, so far as known, failed to reach the State of Alabama.
It is frequent in southern Georgia, but gradually diminishes in abundance northeastward
and barely enters North Carolina.

Such a distribution indicates that the species originated on that part of the Cretaceous
peneplain which has since become the Georgia Piedmont. Although unable to survive the
geologic changes there, its seeds found their way down the Altamaha River system, and
colonized the Coastal Plain. It thrives best in moist meadows or open pinelands, underlain
by loamy but intensely acid soil, and as such habitats are common, it has attained a wide range.

1

PARROT PITCHERPLANT

Sarracenia psittacina Michaux

The distribution indicated on the accompanying map shows that the parrot pitcherplant
originated at some point in the ancient peneplain near what is now the northwestern corner
of Georgia From this region it disappeared as a result of the geologic and climatic changes,
but its seeds found their way down both the Alabama and Chattahoochee Valleys. The
colonies thereby formed on the Coastal Plain expanded laterally, reaching the vicinity of
New Orleans on the west and the coast of Georgia on the east. Search for it near Augusta,
Georgia, where it was stated by Michaux to occur, has proved unsuccessful, the northeastern-
most colony thus far found lying lo miles southwest of Mdlen, in Jenkins County.

Like most of the others, this species apparently lost the abUity to expand its range farther
before the end of the Tertiary. It thrives best in low meadows subject to frequent munda-
tion by acid waters from nearby swamps, but in spite of the abundance of such habiuts
which developed in peninsular Florida after it emerged from the sea, the plant has never
succeeded in entering that region.

FIGURE 8

Distribution of parrot pitchcrplant, Sarracmia psittacina

i8

I

19

SOUTHERN PITCHERPLANT

Sarracenia purpurea venosa (Rafincsque) Wherry

Southern pitcherplant occurs in swamj^. bogs, and wet meadows in nearly every ^rt of
the State of North arolina. It probably originated in the region now constitutmg Henderson
and adjacent counties, but when this territory was uplifted, the immediate ancestors of the
plant were exterminated. There can be little doubt, however, that it is a remote descendant
from Sarracenia jo,usii,y.^Kh has managed to survive in the same general region, for the
flowers of the two are almost identical, in spite of the dissimilar leaf shape.

In the course of time its seeds were transported down several of the eastward-flowmg
rivers forming colonies on the Piedmont and ultimately on the newly emergmg Coastal Plam.
Some' seeds also found their way down the Chattahoochee River system and an extensive
series of colonies developed near the coast on either side of this valley. In addition, unhke other
species, it migrated from northeastern North Carolina to southern New Jersey, over a smp of
land which apparently connected these States during Tertiary and early glacul time^

FIGURE 9

Distribution of southern pitcherplant, Sarracenia purpurea venosa

lO

XI

NORTHERN PITCHERPLANT

Sarracenia purpurea gibbosa (Rafinesque) Wherry

Northern pitcherplant ranges from Maryland, Delaware, and New Jersey northward over
a vast territory in this country and Onada. The geographic relations, together with the
presence of intermediates between the northern and southern subspecies in southern New
Jersey indicate that the northern one originated in that region. Evidently the southern an-
cestor' after arriving in New Jersey, became variable, and gave rise to descendants differing
more or less in morphological and physiological characters. Most of these, lacking ability to
extend their ranges, have remained where they originated. One, however, became highly
aggressive, and as it also differs in pitcher shape, it is classed as a distinct subspecies.

During the ice advances of the glacial epoch this pitcherplant was of course unable to
migrate northward, and merely spread a short distance into Maryland and southern Pennsyl-
vania, but after the retreat of the last ice sheet it soon occupied the newly developed bogs,
and even managed to reach sub-Arctic Canada.

FIGURE lO

Distribution of northern pitcherplant, Sarracenia purpurea gtbbosa

IX

Reprinted by Permission from

Illustrations of

NORTH AMERICAN PITCHERPLANTS

By Mary Vaux Walcott; with Descriptions and Notes on
Distribution, by Edgar T. Wherry, and Notes on Insect
Associates, by Frank Morton Jones.

Published by the

SMITHSONIAN INSTITUTION

Washington, D. C. 1935.

Il

Reprinted from American Fern Journal, Vol. 25, No. 4,
Gctober-Deceniber, 1935.

Fern Field Notes, 1935

Edgar T. Wherry^

A Kentucky station for Adiantum Capillus-
VENERis.— Learning that the Southern Maidenhair had
been reported from the vicinity of Burnside, Pulaski
County, Kentucky, a visit was made to that town in July,
in company with Professors S. C. Palmer of Swarthmore
College and Thomas N. McCoy of Catlettsburg, who is
writing up the ferns of the state. On inquiring as to the
possibFe presence of any cataracts in that vicinity, we
were directed to one about one and one-half miles to the
west, along State Highway No. 90, on the south bank of
Cumberland River below Bronston P. O. On going there
we found a large spring emerging from a limestone cav-
ern, but thorough search of the cliffs and talus failed to
yield this or any other noteworthy fern. Just as we were
leavin*^, I noticed that although a current of very cold
air accompanied the outflowing water, a counter-current
of decidedlv warm air could be felt. Realizing that the
place to look for a subtropical plant in this northern lati-
tude would be in the warmest situation, the stream was
followed to the point where it descended over a series of
limestone ledges into the river and no influence of the cold
cavern air remained. Here the fern sought proved to be
crrowin- in the greatest luxuriance. On previous occa-
sions I have found rare plants by flashlight, and once
detected an as yet undescribed Bryopterh in Alabama by
seeincr its reflection in a mirror while shaving beside a
purling brook, but this was the first time I had ever
located anything by the aid of hot air! ^ ^ ., _
AsPLENiUM Bradleyi IN New Jersey.— On April 7,
1935 a -roup of six members of the American Fern
Society visited the cliffs of a spur of Kittatinny Moun-
~ 1 Contribution from the Botanical Laboratory and Morris
Arboretum of the University of Pennsylvania.

|1

II

American Fern Journal

Volume 25, Plate 12

Fig. 1. — Pellaea densa at its southeastern limit, Black
Lake, Quebec.

m

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■ai.i«ii

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WLx

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if'

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f

Fig. 2. — Asplenium resiliens at its northeastern limit,
Franklin Co., Pennsylvania.

Fern Field Notes

125

tain, about 5 miles northwest of Blairstown, Warren
County, New Jersey, in the hope of rediscovering Asplen-
ium TrudelU, collected there by Carhart many years
ago. Asplenium montanum was found in abundance,
several miles east of its previously known colonies near
Delaware Water Gap, but A. TrudelU eluded us. Then
just as we were about to leave, a single plant of A. Brad-
leyi, not previously reported from the state, was noticed
in a vertical joint-crevice in the quartzite cliff. Five
months later the place was revisited, in company with IMr.
Harry W. Trudell, and the cliff was more fully explored.
Not only were six additional plants of this species found.
in other joint-crevices, but also a colony of the hybrid
A. Bradleyi x montanum. The station is 40 miles, as the
spore blows, east of the nearest known Pennsylvania
occurrence of A. Bradleyi at Glen Onoko, Carbon County,
and about 75 miles southwest of the long-lost colony in
the Shawangunk Mountains in Ulster County, New York.
Asplenium resiliens in Pennsylvania.— Ever since
the finding of the black-stem spleenwort in Maryland last
year I have cherished the hope of extending its range still
farther north, and on September 12 made a systematic
search for it. In the main limestone valley around Green-
castle there seemed to be no cliffs of sufficient size to fur-
nish a favorable habitat for it, but in a narrower strip of
limestone east of Mercersburg several promising cliffs
were located and searched, one after another. Although
the only abundant fern of this group was A. trichomanes,
a small colony of the species sought was finally found on
one sheltered cliff. (See Plate 12, fig. 2.) To avoid pos-
sible destruction by over-enthusiastic collectors its exact
location will not be divulged, but it lies in Franklin
County, well north of the Mason and Dixon line.

Pellaea densa and some other ferns in southern
Quebec— On August 25 a visit was paid to Black Lake,
Mec-antic County, Quebec, to search for the southeastern-

IRREGULAR PAGINATION

A.MKkirAx Fkhx JorHXAr-

Volume 2;', Plate 12

Fi:kx Fikli) Xotf.s

1

'J.')

Fl(!. 1. — PkLLAEA HEXSA at its SOUTHEAKTEKX limit, liLACK

Lake, Qiehec.

I

m
1

tain, about 5 milos iiortliwost of lilairstowii, Warroii
(\)iiiity, Xow Jorsey, in the lioix' of r('(lisfovorin«,r AspJeu-
ium TruileUi, collected there by (^arliart many years
ago. Aspleniiim montanum was found in abundance,
several miles east of its previously known colonies near
Delaware Water Gap, but A. TrudeUi eluded us. Then
just as we were about to leave, a single plant of .1. lirud-
Icfji, not previously reported from the state, was noticed
in a vertical joint-crevice in the (juartzite clitt'. Five
months later the place was revisited, in comi)any with Mr.
Harry W. Trudell, and the clifl^" was more fully exi)lored.
Xot only were six additional plants of this s|)ecies found,
in other joint-crevices, but also a colony of the liybrid
A. Bradleyixmontmivm. The station is 40 miles, as the
spore blows, east of the nearest known Pennsylvania
occurrence of A. Bradlejfi at Glen Onoko, Carbon County,
and about 75 miles southwest of the long-lost colony in
the Shawangunk Mountains in Ulster (V)unty, Xew York.
AsPLKXiUM Ri:siLii:xs IX Pi:xxsYLVAN"iA.— Ever since
the finding of the black-stem spleenwort in Maryland last
year I have cherished the hope of extending its range still
farther north, and on September 12 made a systematic
search for it. In the main limestone valley around (Ireen-
castle there seemed to be no cliffs of sufficient size to fur-
nish a favorable habitat for it, but in a narrower strip of
limestone east of :Mercersburg several promising cliifs
were located and searched, one after another. Although
the only abundant fern of this group was A. fyirhommirs.
a small' colonv of the species sought was finally found on
one sheltered cliff. (See Plate Pi, fig. 2.) To avoid pos-
sible destruction by over-enthusiastic collectors its exact
location will not be divulged, but it lies in Franklin
Countv, well north of the Mason and Dixon line.

Pellaea dexsa and some other ferxs ix sotttiterx
QiTEBEC— On August 25 a visit was paid to Black T^ake,
^re<rantie County, Quebec, to search for the southeastern-

Fig. 2, — Asplenium resiliex.s at its xorttiearterx limit,
Fraxklix Co., Pfxxsylvaxia.

INTENTIONAL SECOND EXPOSURE

126

American Fern Journal

Reprinted from American Fern Journal, Vol 26 No. 4,
October-December, 1936, pages 127-131, 141-142

I

most recorded station for the fern best known as Pellaea
densa (Brack.) Hook., also bearing the name Cheilanthes
siliquosa Maxon, although it perhaps does not belong to
either of these genera. A large colony of it (see Plate
12, fig. 1) was finally located after a long and strenuous
trip through the woods and swamps, and in order to save
others unnecessary effort the following directions may be
given. From Black Lake village drive south along high-
way No. 1 for two miles, when an electric power line will
be seen crossing the lake and extending westward into the
hills. Follow this for a mile, and a prominent cliff of
serpentine will come into view to the north. Cross the
intervening swamp and climb to the top of the talus slope.

The following also seem worth recording: Crypto-
gramma Stelleri and Woodsia glabella, three miles north
of Leeds Station, Beauce County, on a phyllitic mica
schist which looks non-calcareous, but contains enough
lime to neutralize the crevice-soil; Dryopteris fragrans
and Woodsia glabella on a serpentine cliff four miles
southwest of the same Station; and Dryopteris Boottii
in a dense alder thicket at the south end of Black Lake.

Lycopodium porophilum. — The Cliff Staghornmoss
lias been variously interpreted as a distinct species, a
variety of L. lucidulum, and a variety of L. Selago. In
the hope that field study might aid in deciding which
view to accept, visits were made to several stations for it
in Kentucky, Ohio, Pennsylvania, Virginia, and Wiscon-
sin. Although it is sometimes alleged to grow on cal-
careous substrata, in every colony seen the rock was
siliceous and the soil reaction mediacid. Everything
about its habitat and aspect indicated it to represent an
ecological derivative of the dwarf er and more appressed-
leaved L. Selago (typicMm), resulting from the falling
of spores of this occupant of bleak situations into more
sheltered crevices at lower altitudes or latitudes.

Philadelphia, Pa.

Fern Field Notes, 1936

Edgar T. Wherry^

Two CALCAREOUS-SOIL FERNS IN LOWER SoUTH CARO-
LINA.—In listing the ferns of the coast region of South
Carolina, Miss Bragg^ included Dryopteris patens as col-
lected by Ravenel at Eutaw Springs and by herself as an
escape at Otranto. The plant concerned is what was sub-
sequently segregated by Christensen as D. normalis. On
July 6, 1936, Mr. J. E. Benedict, Jr., and I explored the
bluff on the west side of Biggin Swamp, through which
the old Santee Canal runs, about 2 miles northeast of
Moncks Corner. Here an outcrop of Cooper Marl yields
an abundance of calcareous soil, and this fern occurs in
great luxuriance and profusion, forming an especially
beautiful display near the Cooper River. On a hard
marl ledge there were also found several plants of
Asplenium resiliens, which was not listed by Miss Bragg
(though possibly represented by the Bachman report of
A. trichomanes) .

Woodsia ilvensis rediscovered in North Carolina.—
Among the notable plants observed in the vicinity of Jef-
ferson, Ashe County, North Carolina, in 1841, Asa Gray»

1 Contribution from the Botanical Laboratory and Morris Arbo-
retum of the University of Pennsylvania.

2Thi8 Journal 4: 92. 1914. o .q iRflo

8 Am. J. Sci. 24: 27. 1842 ; Sci. Papers Asa Gray 2: 49. 1889.

I' -^

128

American Fern Journal

Fern Field Notes

129

listed Woodsia ilvensis, without, however, giving any
definite locality for it. Being unable to find any speci-
mens preserved, I expressed doubt as to the occurrence of
the species in the State in my Fern Field Notes for 1933.'*
Subsequently, however, Mr. Weatherby advised me that
one of the fronds on a mixed sheet in the Gray Herbarium
is ascribed to North Carolina. On July 14 Mr. Benedijt
and I followed the more or less distinct trail southward
from Jefferson up Nigger Mountain. Our first find con-
sisted of Lycopodium selago (which Gray had found on
Phoenix Mountain) in a vertical crevice close to the trail.
For a long time our search of the rock ledges was other-
wise fruitless, but by a fortunate chance we worked west-
ward from the trail between the two levels where exten-
sive rocky flats are developed; and here on a gentle
northwest slope, with scattered shrubs forming some
shelter, we came on a colony comprising about 10 clumps
of the Woodsia sought. Whether this represents the
occurrence found by Gray 95 years previously is ques-
tionable, but at any rate the species can now be correctly
included in the list of North Carolina ferns.

Other southern records of Woodsia ilvensis deserve
brief discussion. Because of its inclusion by Williamson^
many compilers of data ascribe it to Kentucky, overlook-
ing that author's admission that **I have not been able
definitely to give this beautiful little fern * a local habita-
tion and a place' in Kentucky. ' ' In reporting it in West
Virginia® I made an error in the county; the locality, 2
miles southwest of Landes, is not in Pendleton as pre-
viously stated, but about i mile within Grant County.
Unfortunately this place has been visited in recent years
by overzealous collectors, who have removed the fern

* This Journal 23 : 112. 1933.
8 Ferns of Ky.: 115. 1878.
» This Journal 13: 108. 1923.

r -1

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— *,

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completely. At least 7 stations in Virginia are now
known, in the following counties: Augusta, 2; Giles, 1;
Page, 3 ; and Shenandoah, 1.

An intermediate beech-fern in Macon County,
North Carolina. — Our two eastern beech- ferns are often
termed respectively the ''long" and the "broad" beech-
ferns, but as they do not differ in outline as consistently
as these adjectives imply, I prefer to term Dryoptcris
Phegopteris {Phegopteris polypodioides) the northern,
and D. hexagonoptera the southern beech-fern. At the
time Dr. Blomquist^ was compiling his Ferns of North
Carolina, he showed specimens from the Dry Falls of the
Cullasaja, 3 miles west of Highlands, Macon Co., N. C,
to Dr. Maxon and myself, and on our recommendation in-
cluded it as a scaly form of the southern beech-fern. On
August 30th of the present year I visited the place, and
found all the plants there (Plate 15, fig. 1) to show one
of the diagnostic features of the northern species, namely
absence of wing on the rachis between the lowest and the
second pair of pinnae. Although not so hairy as usual
for the northern plant, the laminas do bear some broad
brown scales on the under side, so I now think that the
occurrence should be regarded as a southern range-exten-
sion of the northern beech-fern. Evidently it is enabled
to grow this far south by the continuous current of cool
air produced by the waterfall behind which it grows.

A remarkable Fern-Association on Avery Island,
Louisiana.— On September 9, 1936, a visit was made to
Avery Island, in Iberia Parish, on the coast of Louisiana,
under the guidance of Mrs. Dan Debaillon. In the steep-
walled ravines grow several species of ferns, including
Adiantum pedatum, not previously reported this far

TFernsof N. C: 72. 1934.

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130

American Fern Journal

south. At one point were seen several plants of Pteris
cretica, presumably an escape from a greenhouse of some
nearby estate. Alongside of one clump of this sub-
tropical species there was found most unexpectedly a
small plant of the decidedly northern ** narrow-leaf
spleenwort ' ' or glade-fern, Athyrium pycnocarpon. The
southernmost station of this fern previously known is In
West Feliciana Parish, 75 miles away; its ability to sur-
vive at this still more southern point is evidently due to
the ravine being very narrow, completely shaded from
the sun, and well cooled by air currents. A photograph
showing the two ferns of such divergent geographic rela-
tionship growing side-by-side is reproduced as Plate 15,
figure 2.

Trichomanes petersii at Saratoga, Mississippi. — On
September 5, 1936, I drove to Saratoga station, Simpson
County, Mississippi, to try to rediscover the locality from
which this fern was distributed to herbaria some years
ago. It was found by first walking a few hundred feet
north along the track to a point opposite the big spring
(after which the place was presumably named), then
turning northeast and following the route of a long aban-
doned railroad siding about a thousand feet. Ledges of
ferruginous sandstone outcrop here, and toward their
northeast end a small spring oozes out. The fern occurs
on shaded vertical faces of the rock above this springy
place, in moderate abundance. Mitchella repens was
noted as a close associate, corresponding to the soil being
rather dry and strongly acid in reaction. Various indif-
ferent or acid-soil ferns occurred in the vicinity, includ-
ing a colony of a score of plants of Botrychium dissect um
var. tenuifolium (Underw.) Farw., remarkable in fruit-
ing while of very small size, 7.5 to 15 cm. high over all. — ■
Edgar T. Wherry, Philadelphia, Pa.

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American Fern Journal

Volume 26, Plate 15

Fig. 1 (UPPER).— Northern Beech-fern at its southernmost
known station, Dry Falls, Ma( on Co., North Carolina.

Fig. 2 (lower).— The sub tropical Pteris cretica growing be-
side THE northern Gladekern, Athyrium pycnocarpon, on
Avery Island, Louisiana.

J

It

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American Fkrx Journal

VoLUMK 20, Tlatf. IT)

i:]()

Amehk'An Fehn Journal

soutli. At one point were seen several plants of Pterin
cretka, presnniably an escape from a greenhonse of some
nearby estate. Alongside of one clump of this snb-
tr-opical species there was found most unexpectedly a
small plant of the decidedly northern *' narrow-leaf
spleenwort ' ' or glade-fern, Athyrium pycnocarpon. The
southernmost station of this fern previously known is In
West Feliciana Parish, 75 miles away; its ability to sur-
vive at this still more southern point is evidentl.y due to
the j-avine being very narrow, completely shaded from
the sun, and well cooled by air currents. A photograph
showing the two ferns of such divergent geographic rela-
tionship growing side-by-side is reproduced as Plate 15,
figure 2.

Trkmiomaxks pktersii at Saratoga, Mississhmm. — On
Septcmlx'r 5, l!);j(), 1 drove to Saratoga stati<m, Simps(m
County, Mississippi, to try to rediscover the locality from
which this fern was distributed to herbaria some vears
Hgo. it was found by first walking a few hundred feet
north along the track to a point opposite the big si)ring
(after which the place was presumably named), then
turning northeast and following the route of a long aban-
doiK'd railroad siding about a thousand feet. Ledges of
ferruginous sand.stone (mtcrop here, and toward their
northeast (Mid a small spring oozes out. The fern occurs
on shaded vertical faces of the rock above this springy
place, in moderate abundance. Mitchella repens was
noted as a close associate, corresponding to the soil being
rather dry and strongly acid in reaction. Various indif-
ferent or acid-soil ferns occurred in the vicinity, includ-
ing a colony of a score of plants of lintryvhium dissectum
var. ffnuifoliuni (rnderw.) Farw., remarkable in fruit-
ing while of very .small size, 7.5 to 15 cm. high over all. — ■
Ed(}Ar T. Wiikrry, rhihidelphui. Pa.

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Fio. 1 (UPPKR).— North KRN liKK( ii kirn at ns soitiif-rnmost

KNOWN station. DRY FALLS, >rA( ON CO., NoRTII CAROLINA.

Fig. 2 (lowkr).— Tiik srn troi'K al Ptkris < ri-.tk a cRowiNi; hk

.SIDK TIIF, NORTIIF.RN (JlADE FF.RN, AtIIVRUM I'YCNOCARI'ON, ON

AvKRV Island. Loi isiana.

INTENTIONAL SECOND EXPOSURE

Reprinted from Bartonia No. 18, 1936.

Miscellaneous Eastern Polemoniaceae^

Edgar T. Wherry

In bringing to a close my series of studies on the members of the Polemonium
Family occurring east of the Mississippi River, a key to the genera concerned,
simplified to cover only eastern representatives, is first given:

POLEMONIACEAE: KEY TO NATIVE AND ADVENTIVE EASTERN GENERA

Sepals all alike, entire.
Calyx-tube remaining unbroken to maturity of capsule.
Leaves pinnately compound; calyx-tube herbaceous, markedly accrescent.

Duration perennial; flowers borne in cymose clusters; bracts small Polemonium.

Duration annual; flowers borne singly in the axils of large bracts Polemoniella.

Leaves entire to pinnatifid; calyx-tube more or less herbaceous, not markedly accrescent

Calyx-tube becoming ruptured by the maturing capsule.
Leaves entire or essentially so ; corolla salverform Phlox.

Leaves lobed or dissected.
Plants coarse biennials with numerous leaves and thyrsoid inflorescence ; corolla red

open salverform ; seeds obliquely polyhedral, buff, translucent Ipomopsis.

Plants delicate annuals with few leaves and corymbose to capitate ^"A^^^f ^/^f ^ ^°;;°"*
violet or lavender, open salverform to funnelform; seeds ellipsoidal, light to^^J^

brown, opaque * * . '

Sepals unequal, spinulose-lobed ; calyx-tube unbroken to maturity of capsule Navarretm.

Identity of Polemonium ciliatum Willd. ex Roem. & Schult.

In the course of collecting material for the present article, a question arose as
to the identity of " Polemonium ciliatum.'' This had been sent to Europe by
Muhlenberg from Pennsylvania, presumably from Lancaster County, where he
lived, and was described ' as having acute leaf-segments and calyx-lobes. In
compiling the Kew Index, Jackson « recognized that it represented a Phaceha but
referred it to P. fimbriata Michx., overlooking the facts that this has obtuse leaf-
segments and calyx-lobes, and grows only farther southwest. Brand * at first
accepted this interpretation, but later « allocated this supposed Polemonium to
Phacelia purshn Buckley. That this is the most satisfactory solution of the
problem is affirmed here, in order that future workers may avoid overlooking
Brand's final conclusion.

1 Contribution from the Botanical Laboratory and Morris Arboretum of the University
of Pennsylvania.

2Systema veg. 4: 792, 1819.

8 Index Kewensis 3: 584, 1894.

* In Engler'a Pflanzenreich IV. 250 : 47, 1907.

» In Engler'a Pflanzenreich IV. 251 : 62, 1913.

S^J

BABTONIA

COLLOMIA NUTTALL
In 1811 Thomas Nuttall collected in the northwestern prairie region, chiefly
along the lower Cheyenne valley in what is now South Dakota, a small annual
which obviously belonged to the Polemoniaceae, but was peculiar in that its seeds
became mucilaginous when moistened. He decided that this represented a pre-
viously unrecognized genus, including also one or two South American plants
which had been assigned to Phlox, and proposed^ for this the name Collomia,
from the Greek word for glutinous material. Gray * at first accepted the genus
as valid, transferring into it several plants previously classed as Gilias. Subse-
quently, however, he recognized * that what he had regarded as a differentiating
character, unequal insertion of stamens, was shown by certain unmistakable
Gilias ; despairing of finding any other distinctive feature, he proceeded to reduce
Collomia to the status of a Section under Gilia. A valid and fundamental basis
for generic distinction was found soon afterward, however, by Greene,* — in typical
Gilias the calyx-tube is ruptured by the maturing capsule, while in the CoUomias
the calyx is sufficiently accrescent to remain intact to maturity. This was ignored
by Gray and by the editors of the 6th and 7th editions of his Manual of Botany,
where the sectional status of Collomia is maintained; but most recent workers
have admitted the generic independence of Collomia, which is emphatically
favored here. The genus comprises approximately 5 perennial and 5 annual
species of western North America, with 2 or 3 annuals in southern South America.
Only Nuttall's original species ranges sufficiently far to the east to be covered in
the present article.

Collomia linearis Nuttall.

History. — ^The original species on which the genus Collomia was based was
named by Nuttall * C. linearis in reference to its dominantly linear leaves. It was
mistakenly transferred to the genus Hoitzia ( Jussieu 1789) by Sprengel ' in 1825,
while many years later, under the circumstances above discussed, Gray ^ made it
Gilia linearis. It was renamed Collomia parviflora by Hooker® and several
variants have received special names: a broad-leaved phase C. lanceolata
Greene; ' a stunted one C. linearis var. humilis Brand,® one with the bracts some-
what mottled C. linearis var. picta Lunell,® and still another with profuse branch-
ing C. linearis var. congesta Lunell.^^

1 Genera N. Am. Plants 1 : 126, 1818.

2Proc. Amer. Acad. Arts Sci. 8: 258, 1870.

«Proc. Amer. Acad. Arts Sci. 17: 223, 1882.

*Pittonial: 122, 126, 1887.

B Systema veg. 1 : 626, 1825.

•Curtis's Botanical Magazine 56: pi. 2893, 1829.

7 Plantae Bakerianae ; ex Brand in Engler's Pflanzenreich IV. 250 : 49, 1907.

8 In Engler's Pflanzenreich IV. 250: 49, 1907.

9 Bull. Leeds Herb. 2: 7, 1908.

10 Am. Midi. Nat. 4: 512, 1916.

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BARTONIA

COLLOMIA NUTTALL
In 1811 Thomas Nuttall collected in the northwestern prairie region, chiefly
along the lower Cheyenne valley in what is now South Dakota, a small annual
which obviously belonged to the Polemoniaceae, but was peculiar in that its seeds
became mucilaginous when moistened. He decided that this represented a pre-
viously unrecognized genus, including also one or two South American plants
which had been assigned to Phlox, and proposed ^ for this the name Collomia,
from the Greek word for glutinous material. Gray ^ at first accepted the genus
as valid, transferring into it several plants previously classed as Gilias. Subse-
quently, however, he recognized ^ that what he had regarded as a differentiating
character, unequal insertion of stamens, was shown by certain unmistakable
Gilias ; despairing of finding any other distinctive feature, he proceeded to reduce
Collomia to the status of a Section under Gilia. A valid and fundamental basis
for generic distinction was found soon afterward, however, by Greene,* — in typical
Gilias the calyx-tube is ruptured by the maturing capsule, while in the Collomias
the calyx is sufficiently accrescent to remain intact to maturity. This was ignored
by Gray and by the editors of the 6th and 7th editions of his Manual of Botany,
where the sectional status of Collomia is maintained; but most recent workers
have admitted the generic independence of Collomia, which is emphatically
favored here. The genus comprises approximately 5 perennial and 5 annual
species of western North America, with 2 or 3 annuals in southern South America.
Only Nuttall's original species ranges sufficiently far to the east to be covered in
the present article.

Collomia linearis Nuttall.

History. — The original species on which the genus Collomia was based was
named by Nuttall ^ C. linearis in reference to its dominantly linear leaves. It was
mistakenly transferred to the genus Hoitzia (Jussieu 1789) by Sprengel '^ in 1825,
while many years later, under the circumstances above discussed, Gray ^ made it
Gilia linearis. It was renamed Collomia parviflora by Hooker® and several
variants have received special names: a broad-leaved phase C. lanceolata
Greene ; ' a stunted one C. linearis var. humilis Brand,^ one with the bracts some-
what mottled C. linearis var. picta Lunell,® and still another with profuse branch-
ing C linearis var. congesta Lunell.^^

1 Genera N. Am. Plants 1 : 126, 1818.

2Proc. Amer. Acad. Arts Sci. 8: 258, 1870.

aProc. Amer. Acad. Arts Sci. 17: 223, 1882.

* Pittonia 1 : 122, 126, 1887.

» Systema veg. 1 : 626, 1825.

«Curti8's Botanical Magazine 56: pi. 2893, 1829.

^ Plantae Bakerianae ; ex Brand in Engler's Pflanzenreich IV. 250 : 49, 1907.

8 In Engler's Pflanzenreich IV. 250: 49, 1907.

oBiill. Leeds Herb. 2: 7, 1908.

10 Am. Midi. Nat. 4: 512, 1916.

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INTENTIONAL SECOND EXPOSURE

il

MISCELLANEOUS EASTERN POLEMONIACEAB

m

Geography. — ^Botanical exploration of western North America subsequent to
Nuttall's time showed Collomia linearis to be widespread there, and by 1870
Gray ^ was able to state the range as " Both sides of the Rocky Mountains, north
to Mackenzie River; also on the shore of New Brunswick, Fowler, perhaps a
waif." With respect to the final item, it should be noted that subsequent collect-
ing has shown that the species is present at a number of stations in both New
Brunswick and Quebec, in a region where many northwestern plants occur as
relics of a former cross-continental distribution, disrupted by the last ice sheet.
These eastern colonies are accordingly here regarded as native, although the plant
is unquestionably of recent introduction at several intervening points. On the
accompanying map only records east of longitude 100° are shown.

Fia. 1. Distribution of CoUomia linearia.

[Illinois: Becoming increasingly abundant as a weed, especially along rail-
road Imes, in Cook-', Du Page-', Jo Daviess-', and Kane-' counties.]

Iowa: Recorded from Chickasaw*, Decatur, Dickinson, Emmet* and Lyon

counties.

[Maine: Adventive in York* County.]

Minnesota: Widespread northward, specimens having been seen from 10
counties: Aitkin, Becker, Chippewa, Goodhue, Jackson, Mille Lacs, Otter Tail,
Pipestone, St. Louis and Steams.

[Missouri: Entering along a railroad in Marion* County.]

Nebraska: Common westward, and ranging east of 100° in Antelope and

Cedar counties.

[New Jersey: Collected once as a weed in Camden* County.]
[New York.: Adventive along a railroad in Cattaraugus* County.]

I 1

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BAETONIA

North Dakota: The county list reaches a maximum of 15 in this state,
although only part of them are toward the eastern end: Barnes, Benson, Cass,
Grand Forks, La Moure, Pembina, Richland, and Stutsman.

[Pennsylvania: Found as a weed at opposite comers, in Crawford^ and
Philadelphia^ counties.]

South Dakota: Widespread in the western part, and collected in one eastern
county, Brookings.

[Vermont: Adventive in waste places in Bennington^ and Chittenden*
counties.]

Wisconsin: Recorded from Douglas, Jefferson^, Pierce, and Price counties,
and apparently native in most of them.

Canada: Extending from the northwest across Manitoba to westernmost
Ontario, in Thunder Bay County. In the St. Lawrence valley recorded from 3 or
4 stations each in Restigouche-Madawaska County, New Brunswick, and Bona-
venture County, Quebec.

Ecology,— Collomia linearis is a pioneer plant, appearing on gravel banks,
sand-hills, and burned-over land at an early stage in their revegetation. It per-
sists into intermediate successional stages, but tends to die out as climax con-
ditions develop. Some of man's activities favor its spread, and it appears to be
expanding its range over the northeastern states along fence-rows, railroad
embankments, and waste-places generally, although there is no indication of its
becoming a serious weed. Its tiny pink ^ flowers are produced for 2 or 3 months
in summer, but seem too inconspicuous to attract insects, and no data are avail-
able as to its pollination.

Vanation.— In response to differences in availability of moisture and of
nutrients, this species varies from a slender, narrow-leaved simple-stemmed plant
a few centimeters tall to'a robust lanceolate-leaved plant with a much-branched
inflorescence. The following forms may be distinguished:

FORMS OF COLLOMIA LINEARIS.
Stunted. C. I. var. humilis Brand.
Luxuriant. C. I. var. congesta Lunell.
Broad-leaved. C. lanceolata Greene.
Mottled-braeted. C. I. var. picta Lunell.

Cultivation.-ln the text accompanying the figure of this plant in the
Botanical Magazine, Hooker ^ noted that it had been " introduced by Mr Douglas
to the garden of the Horticultural Society," and that it had proved to be a " hardy
annual, flowering nearly the whole summer." Its flowers are too inconspicuous
however, for it to have found general favor, and its seeds are rarely offered for
sa e. Several other members of the genus, with more showy flowers, are often
cultivated.

1 In Maerz & Paul's Dictionary of Color 41-E-2 to 41-J-6 (or 49-E-6.)

2 Curtis's Botanical Magazine 56: pi. 2893, 1829.

MISCELLANEOUS EASTERN POLEMONIACEAE

IPOMOPSIS MICHAUX.
A showy red-flowered member of the Polemoniaceae native to the southeastern
United States, classed by Linne ^ as a member of his genus Polemonium and by
Lamarck ^ as belonging to Jussieu's Cantua, was made by Michaux ^ the type of
an independent genus, Ipomopsis. Two years later Persoon * referred this plant,
with a question mark, to the Ruiz and Pavon genus Gilia. In 1818 Nuttall,** " for
the sake of euphony," altered Michaux's name to Ipomeria, and still later
Rafinesque ® put forward a wholly new one, Batanthes. Bentham,^ Gray,^ and
most recent authors have concurred in Persoon's generic assignment, although
Greene ^ urged the acceptance of Rafinesque's genus name for some of the species.
On the basis of the characters given in the key, the plant is here regarded as
probably worthy of generic segregation, and a return to the earliest name for it,
that of Michaux, is favored. The genus comprises about 10 western species, and
a single eastern one.

Ipomopsis rubra (Linne) Wherry, comb. nov.

History. — ^An extraordinary number of name-combinations have been applied
to the species under discussion, the more noteworthy being: Polemonium rubrum
Linne,^ Cantua pinnatifida Lam.,^ C. coronopifolia Willd.,^® Ipomopsis elegans
Michx.," Cantua thyrsoidea Juss., ^^ Gilia coronopifolia Pers.,* Cantua elegans
Poir.,^* Ipomeria coronopifolia Nutt.,*^ Navarretia rubra Kuntze ^* and Gilia
rubra Heller.^*^ Since no one has heretofore associated the earliest species name
with that of the earliest genus, a new combination is here called for. A slender
variant which develops in Florida sand barrens has been named successively
Cantua floridana Nutt.,^® Gilia floridana Don," and G. rubra var. capillacea
Brand.^* Several color forms have also been assigned horticultural names, such
as " picta " for one with the corolla yellow within, " lutea " yellow throughout,
and " superba " purplish red.

1 Species Plantarum: 163, 1753.

2Tabl. Encycl., 111. gen. 1: 473, 1791.

8 Flora Boreali-Americana 1 : 141, 1803.

* S3aiopsis plantarum 1 : 187, 1805.

» Genera N. Am. Plants 1 : 124, 1818.

•Atlantic Joum.: 145, 1832.

T In De Candolle's Prodromus 9: 313, 1845.

8 Proc. Amer. Acad. Arts Sci. 8: 275, 1870.

» Leaflets Botan. Obs. & Grit. 1 : 224, 1906.

10 Species Plantarum 1 pt. 2: 879, 1797.

" Flora Boreali-Americana 1: 142, 1803.

12 Ann. Mus. Paris 3: 119, 1804.

" Encycl. meth., Bot., Suppl. 2: 80, 1811.

"Rev. Gen. Plant. (1): 433, 1891.

" Contr. Herb. Franklin & Marshall 1 : 81, 1895.

"J. Acad. Nat. Sci. Phila. 7: 110, 1834.

" Gen. Hist. Dichl. Plants 4: 245, 1838.

"In Engler's Pflanzenreich IV.2S0: 116, 1907.

Ki.

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BABTONIA

Geography.— As a native plant, Ipomopsis rubra ranges from central Texas to
Florida, and sporadically northward to southern Oklahoma and southeastern
North Carolina. It has also escaped from cultivation northward:

Alabama: Reported in but four counties, Autauga, Baldwin, Bibb, and Mobile.

Arkansas: Seen by Nuttall in the southeastern prairies.

[Delaware: Escaped in Sussex^ County.]

Florida: Widespread yet not frequent, specimens having been seen from but
8 counties: Brevard, Duval, Escambia, Lake, Levy, St. Johns, Sumter, and

Volusia.

Georgia: Known in Chatham, Clarke, Cobb, and Gwinnett counties.

[Illinois: Adventive in Winnebago* County.]

[Maryland: An escape in Wicomico* and perhaps other counties.]

[Massachusetts: Recorded from Franklin* County.]

[Michigan: Has succeeded in establishing itself in Allegan* and Oakland*.]

Fig. 2. Distribution of Ipomopsis rubra.

Mississippi: Known thus far only along the coast in Jackson County.
[Missouri: Apparently escaped in Clark*, Greene* and Holt* counties.]
[New Jersey: Established at several points in Atlantic* and Cumberland*.]
[New York: Escaped in Westchester* County.]

North Carolina: Specimens preserved from Craven County probably repre-
sent a native colony. The type locality was given as " Carolina."

[Ohio: Recorded as escaping in Erie*, Lake*, and Sandusky* counties.]
Oklahoma: Specimens seen from Choctaw, Comanche, Jackson and Murray.
South Carolina: Definitely reported only from Fairfield and Oconee counties.
Tennessee: Recorded in McMinn County.

s-^

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miscellaneous eastern polemonuceae

Texas: Abundant and widespread, there being at least 25 county records:
Anderson, Austin, Bexar, Brazos, Burnet, Cherokee, Collin, Comal, Dallas,
Fannin, Galveston, Gillespie, Harris, Kendall, Kerr, Kimble, McLennan, Mitchell,
Montgomery, Morris, Parker, Rusk, San Augustine, Tarrant and Travis.

[Virginia: Escaped in Chesterfield* County.]

Ecology. — Ipomopsis rubra is chiefly an occupant of sandy or gravelly land,
where mineral-rich subsoil or admixture of shells prevent the development of high
acidity. When it invades real sand-barrens it becomes rather delicate in aspect,
and as noted under History, has received varietal or even specific segregation.
Its biennial duration and the reduced transpiration surface of its pectinately
dissected foliage mark it as a moderate xerophyte, although it scarcely invades
the drier parts of Texas. Requiring abundant light, it is best developed in early
successional stages, and dies out from climax forests. The flowers begin to open
in late May toward the southern border of the range, in late June further north,
and continue to appear for two or even three months. Their open-salverform
corolla with red exterior ^ and more or less orange interior are adapted to attract
humming-birds, which visit the colonies in abundance. Large quantities of viable
seeds are produced, which germinate freely in the Spring, forming the first season
dense rosettes ; many of these are killed during the succeeding winter, but those
which survive begin in early Spring to send up the season's blooming stalks.

Variation. — This species varies considerably in size in relation to availability
of nutrients in the soil, but the differences are not here regarded as sufficiently
significant to justify nomenclatorial recognition.

Cultivation.— This showy plant early attracted the attention of Catesby, who
sent seeds to the Eltham garden, where it was studied and described by Dillenius *
in 1732. It soon found its way into many other gardens, and several of the
species names assigned to it were based on cultivated material. In the trade,
both in Europe and America, the seeds are usually offered under the name GUia
coronopifolia. It is of easy culture, though succeeding best in a rather dry, sandy
soil, and tending to succumb to fungus attack in moist locations.

GILIA RUIZ AND PAVON
The genus name Gilia was proposed by Ruiz and Pavon » in 1794, a single
South American species, G. laciniata, being described by them * five years later.
That representatives of this genus occur also in North America appears to have
been first recognized by Sprengel,** who in 1825 listed 5 species, four of which had
been originally described under other genera. From that time on the size of the

1 The color of the exterior is close to the pure scarlet, l-L-12, of Maerf & Paul's Dictionary
of Color.

2Hortus Elthamensis 2: 321, pi. 241, f. 312, 1732, as Quamoclit pennatum, etc.
8 Flora Peruv. & Chilen. Prodr. 1 : 25, pi. 4, 1794.
* Flora Peruv. & Chilen. 2: 17, pi. 123 f. b, 1799.
fi Systema veg. 1 : 625, 1825.

0

i >, BARTONIA

genus increased rapidly, especially through the merging ^f^)^ it of many other
lenera, until it now, as often interpreted, composes over 100 species. Though
freelv admitting that valid grounds for generic segregation are difficult to hnd, 1
feel thaUhe " lumping" process has gone rather too far, and prefer to maintain
Riirh ffrouDS as Ivomopsis and Navarretia as distinct. ,. ^ . i x

None 0 the typical Gilias are native to the eastern United States, but at leas
five have been Reported as escapes; these all have corollas of a more or less
funnelform shape and violet or lavender color, but differ m the respects listed.

GUia achUkifolia Benth. with deep violet broad-petalled flowers m few-

flowered heads, around New York City. „ , «

TcaSta Dougl. ex Hook., with lavender narrow-petalled flowers, m many-

flowered heads around New York and in Highland County, Ohio.

^ atlXa (Smith) Dougl. ex Hook, a small plant with most of is leaves

basal and pale lavender flowers in lax corymbs, in Middlesex County, Mass.

'"g. iita Benth., a much-branched plant with numerous purplish flowers
grouped in a large cyme, in Cumberland County, Mame and Middlesex, Mass

0 tricolr Benth , a spreading-branched plant with the corymbose-cymose
flowe;s yeUow in the center" dark purple-blotched in the throat, and pale lavender
!n the Umb, in Essex County, Mass. The introduced material was recogn . d
b; G eenman^ as representing a variety, which he named 27'"^ 1 ia
nativity was at the time unknown, but it was subsequently found m California.

NAVARRETIA RUIZ AND PAVON
Like the next-preceding genus, this one was proposed » in IJ^J and a species
native to Chile and Argentina, Navarretia involucratav,,^ J;;"^^^ ui 1799 , a
number of species were subsequently recognized m the western United States
Zm6 Endlicher ' reduced its status to that of a section under Gtha, and has
beef follSed by some authors, although others have maintained its genenc dis-
ttactness. KunL » pointed out that the name has page priority over GUra, and

renamed many species usually assigned to the latter.

Two species have been reported as adventive in the east:

Navarretia ir^tertexta (Benth.) Hook., with the calyx ^"'""^/^^^''^f j\£
the violet corollas nearly tubiform, in Lee and Page counties, Iowa, and Allen

""Xl^^r^etia leucocevhala Benth., in which the calyx is viHo- f Y j^^^^^^
costally and the corollas are whitish and funnelform, m Middlesex County,

Massachusetts.

iRhodoraS: 154,1904.

2 Flora Peruv. & Chilen. Prodr. 1 : 20, 1794.

8 Flora Peruv. & Chilen. 2: 8, 1799.

♦Genera plantanim: 657, 1836.

»Rev. Gen. Plant. (1): 432, 1891.

1 -

Reprinted without change of pagination from Jour. So. App. Bot. Club
^ Vol. I. 1936.

i

(

4

^ A

If

THE JOURNAL

of the

Southern Appalachian Botanical Club

Vol.1 October, 1936 No. 6

' ,*■

Observations on Selaginella tortipila'
Edgar T. Whkrry

In the course of a discussion of the Selaginellas of New Grenada
in 1865 A Braun^ called attention to the presence in North America
of representatives of SeluginelUi rupestris P tropica Spring, "quae
specierum dignitatem affectant." For one with contorted leaf-awns
he proposed the species name S. tortipila, indicating that he had on
hand specimens from two places: "In Carolina australi legit Dr. Cur-
tis in Carolina septentrionali Rugel." This plant was reduced to
varietal status, as 8. rupestris var. tortipiUi, by Underwood.

Many years later Underwood* described a Selaginella with the
same sort of awns under the name S. sJierwoodii, its type locality be-
ing near Highland.s, Macon County, North Carolina. It seemed o him
quite diifercnt from the one i«st discussed. "In place of the slender
lax sprawling habit of S. tortipila with enlarged though short stro-
biles, we have a very compact bushy or tree-like plant with stout stems
many.ranked leaves, and strobiles which are .scarcely noticeable as the
branches graduate imperceptibly into them without enlargement^
TWs was in turn classed as a variety, S. rupestris var. sKerwoodn, by

*^^"*These two Selaginellas were classed as distinct from one another
and from S rupestris by Small' in the Flora of the Southeastern

.Contrlbation from the BoUnlcal Laboratory and Morris Arboretum of
'•"* FA°r st'Nlt!?er V^Cnlque 3: 271. 1865 (not p. 2 a, often stated).

3Nart:ive Ferns, ed. 3: 140. 1888.

♦Torreya 2: 172. 1902.

5Fern Allies: 142. 1905 , , „ -qiq.

eFlora SE. U. S. :29. 1903; (ed. 2. 1913).

*i*

66

United States. The key characters separating them were given' as :
Stems erect or ascending. Cones 5 mm. long or less, as

stout as the diameter of the stem S. sherwocxin.

Stems spreading or creeping.

Stems rooting only near the base: leaves with tor-
tuous hairs .1 S- tortipula.

Stems rooting throughout, usually less than 1 dm.
. long: cones 10-15 mm. long: leaves ending in a
slender awn - ^' rupestris.

Their distinctness was also maintained by Van Eseltine,' but his
key was a more natural one :

Setae tortuous; plants prostrate or ascending; mega-
spores rugose-tuberculate.

Plants prostrate, spreading; leaves 8 ranked S. tortipda.

Plants ascending, densely cespitose; leaves 13-

ranked ^- sherwoodii.

Setae straight ; megaspores alveolate S. rupestns.

The differences in seta and spore characters obviously set off
S. rupestns wholly from the other two, and the contrasted habits of
Sf tortipila and S. sherwoodii suggest that they are distinct from one
another. When, however, an attempt is made to apply the characters
given to actual specimens, either in the field or the herbarium, diffi-
culties immediately arise. The material which has been collected
from most of the localities of the plants with tortuous setae consists of
a mixture of prostrate-spreading and ascending-cespitose plants, and
the leaves vary from 8 to 15-ranked on different fragments. No cor-
relation can be found between strobile outline, (emphasized by Under-
wood) and either habit or leaf-characters. Finally, the differences in
spore-characters given in the text of Van Eseltine's article do not
exceed the variability observable within material from the same
colony. Further study of these two supposed species is evidently

needed.

Herbarium material of the tortuous-awned Selaginellas being but

scanty, field study was called for, and visits have been made during

the past few field seasons to practically all their known localities.

Specimens obtained have been placed in the herbarium of the Acad-

emy of Natural Sciences of Philadelphia, which now includes the most

complete series of these plants known to exist. In all cases these

Selaginellas grow on bare slopes of granite or granite-gneiss rocks at

TContr. U. S. Nat. Herb. 20: 161. 1918.

67

elevations of 1500 to 4500 feet above sea-level. In general southwest-
northeast order its localities are :

GEORGIA. Rabun Co.-. The southwesternmost known occur-
rence of tortuous-awned Selaginellas is on the west bank of Chattooga
River, 1 mile south of Pine Mountain P. O.

SOUTH CAROLINA. Unfortunate^' none of the material col-
lected by M A. Curtis, referred to by Braun in his original descrip-
tion of SelaginelUi tortipila seems to be preserved in American her-
baria, and the locality from which it came is not certainly known.
There are however only two places in the state where the conditions
are favorable for its growth, and the second of these listed, being more
accessible, seems its most probable type locality: ,, „ ,

Pickens Co.: A specimen in the Gray Herbarium is labelled as
from Table R^ck, near the northern edge of this county, and not tar
from the following station.

Greenville Co. : Selaginellas with tortuous awns are abundan
on the famous Caesar's Head Mountain from the top of the cliff at
3200 feet (not 4500 as given on some labels) down to about 1750 feet,
four miles by road below the summit. A report by ea tie« from
Hogback Mt. was based on misidentification of my collection of S.
rupestris there.

NORTH CAROLINA. Macon Co. : Bare Ri-anite s'opos on whir-l.
Selaginellas with tortuous awns srow occur on Satula ami other motin-
tains near Highlands. In 1901 frag.nents of such a V^^J^^^
cred in that vicinity by W. h. Sherwo^.d. and shown to liof.sso.
Underwood, who considered it worthy of further study. Sherwood
thn wrote to Thon.as G. Harbison, who in April 1 )02 eo lee ted and
sent in good n.aterial. This Sherwood took to Underwood clamun.
That he wa-s the collector, and shortly thereafter the plant was de-
scribed as the new species S. sheru-oodn.

" „ 1934 Mr. Harbison kindly guided n,e to ^XT.XiT.
which he had collected those specinu-ns. on Sunset Rock, at 4000 fe t
rimude Although vandalisn. on the part of the pubhc was hardly to
have been expected in the ea.se of such an inconspicuous. non-Hower.n.
pint manv of the clumps proved to have been stnppe.l off dunn.
fnteie^ng years for use as fuel in a stone fireplace. A sheet of
TopoTyTe mat' rial was nevertheless collected, and a sun.lar one was ob-

— T^a Tryon Region, J. El.sha Mitchell Sci. Soc. 44: 120, 1928.

S

V7

IRREGULAR PAGINATION

68
tained from the summit of Satula Mt., altitude 4550 feet.

Jackson Co. ; Tortuous-awned Selaginellas have been collected
from at least 6 stations in this county, the most extensive being on
Whiteside Mt. at 4000 to 4500 feet, the others around Cashiers and
Glenville.

Transylvania Co.: On slopes near Lake Faii'field and 1^ miles
north of the villaj^c of Cedar Mt., elevation 2750 feet.

Henderson Co. : Collected from a now lost locality near Hender-
sonville by Huger in 1898 (N. Y. Bot. Gard.)

Rutherford Co. : The second collection noted by Braun in the
original description of S. tortipila was made by Ferdinand Rugel ; his
labels bear the data '' Lycopodium rupestre L. In locis rupestris humi-
dis montium ad Broad River, Carolina Sept. legit Rugel Jul. 1841."
Some of his sheets in American herbaria contain only 8. rupestris,
well known along the Broad River, but others bear a tortuous-awned
plant regarded by subsequent writers as typical S. tortipila. The lat-
ter occurs along that river only at Chimney Rock, at 1500 to 2000 feet
altitude, which is undoubtedly Rugel's locality.

Burke Co. : The earliest known collection of a tortuous-awned
Selaginella is one in the Academy of National Sciences of Philadelphia
herbarium labelled ''Table Rock, N. C. leg. Rinehart 1839." In 1841
it was again obtained there by Gray and Carey, and has been subse-
quently collected several times. Other stations in this county com-
prise Linville Falls, Wiseman's View, and Ilawksbill Mt., the last a
northern limit.

Ashe Co. : I have seen no material from this county to substan-
tiate reports, and could find only 8. rupestris on Phoenix Mt., one of
the few on which bare rock slopes are developed. Avery Co. has also
failed to yield it.

TENNESSEE. Polk Co.: 8. tortipila was reported from ''dry
rocks in Ocoee Valley" by Gattinger,^ but no herbarium specimens
appear to have been preserved, and the species may well have been
S. rupestris.

In every one of these localities of tortuous-awned Selaginellas
which was visited, the relations proved to be much the same. At the
top and in the center of each clump, the stems were found to be erect
or nearly so, the leaves crowded an ' many-ranked, and the strobili
not markedly set off from the stem below. At the bottom and around

eFlora Tenn.: 31, 1901.

69

the margins of the clumps, or throughout such clumps as chanced to
be shaded or crowded by rocks or by other plants, the stems are pros-
trate, the leaves well spaced and few-ranked, and the strobili rather
distinctly set off. In the drier and more exposed situations, the first
type of development is most conspicuous, in moister or more sheltered
places the second type. At Chimney Rock in Rutherford Co., N. C,
where the slope is unusual in facing north, and the plants are corres-
pondingly well shaded, the slender prostrate type is dominant, which
accounts for the fact that Rugel 's collections were wholly of this type ;
on the sunny ledges of the "Chimney", however, stoutish upright
stems occur as well. Around Highlands, on the other hand, the up--
right habit is most frequent, and is represented in many collections
made there, including the type of ''8. sherwoodn.''

Recently Professor R. M. Reeve has been makuig a study ot
Selaginella spores, and has been so good as to report to me, m advance
of his own publication, his finding that those of both supposed tor-
tuous-awned species are identical. Since both -S.tortrpda and ^S
.sherwoodii- occur, then, in the same colonies and even in the sam c
clumps, the second is to be cl^^sed as a mere ecologica form of the
first and withdrawn from the list of species of SelagmeUa.

IRREGULAR PAGINATION

Reprinted from

'THE AMERICAN MIDLAND NATURALIST"
VoL 16. No. 3, pp. 413-416, 1935

AN OZARK VARIETY OF PHLOX PILOSA-

EDGAR T. WHE.RRY

Some ten years ago the writer undertook a taxonomic study of the genus
Phlox, starting with the species native east of the Mississippi River. The
fifteen species distinguishable there have been discussed, a few at a time, in
a series of articles in Bartonia (1929-1935). Of these, seven also cross that
river and range into the Ozarkian Highlands of Missouri and Arkansas:
P bifida Beck, P. dtvaricata L., P. pilosa L., P. glaberrtma L., P. maculata L.,
P. ampltfolia Britton, and P. pamculata L. So far as can be ascertained from
material at hand, all but one of these are represented in the Ozark region by
varieties also known further east, although a more detailed study of this point
might well be made by someone who can carry on field comparison ot the

Fig. I. Phlox pilose ozarkana north of Shrcveport. Caddo pa r..h. Louisiana.
Photograph by Edgar T. Wherry. April 20. 1934.

the U

* Contribution from the Botanical Laboratory and Morn. ArWctum of
verity of Pennsylvania. A trip to the Ozark reg.on m Apr.l 1934 to obta.n fie d data
on the plant here described was made possible by a grant from the Board of Graduat.

Education and Research.

(413)

licpiintcd fiiiMi

THE AMERICAN MIDLAND NATURALIST"
Vol. 16, No. 3, pp. 413-416, 1935

AN OZARK VARIETY OF PHLOX PILOSA

EDGAR T. WHERRY'

Some ten years ago the writer undertook a taxonomic study of the genus
Phlox, starting with the species native east of the Mississippi River The
fifteen species distinguishable there have been discussed, a few at a time in
a series of articles in Bartonia (192^-1935). Of these, seven also cross thai
river and range into the Ozarkian Highlands of Missouri and Arkansas:
P. bifida Beck, P. diraricata L, P pdosa L„ P. gbberrtma L., P. maculata L.,
P amplifolta Britton, and P. panuidata L. So far as can be ascertained from
material at hand, all but one of these arc represented in the Ozark region by
varieties also known further east, although a more detailed study of this point
might well be made by someone who can carry on field comparison of the

Fin. 1. Phlox pilosa ozarkana north of Shreveport. ^a^do parish. Louisiana.
Photograph by Edyar T. Wherry. April 20. 1934.

"'^Contribution from the Botanical Laboratory and Morris Arboretum of '^e Uni-
versity of Pennsylvania. A trip to the Ozark region m April 1934 to obtain field data
on the plant here described was made possible by a grant from the Board of GraduaU
Education and Research.

INTENTIONAL SECOND EXPOSURE

414

THE AMERICAN MIDLAND NATURALIST

plants of the respective regions. The one exception is Phlox pilosa, which hIS
previously noted (1931, p. 46) here presents a well-markecl local variety.
This may be known as:

Phlox pilosa ozarkana var. nov.

Ozark Phlox

Plant a glandular-pubescent or rarely glabrate perennial with short thickish
rootstocks; stems single or often several from one crown, simple or somewhat
branched, thickish, averaging 40 cm. tall, with 6 to 10 nodes up to the inflo-
rescence; leaves opposite or the uppermost alternate, thinnish, acute to acumi-
nate, passing from linear at the base of the stem to oblong-lanceolate or
broadly ovate and cordate-clasping, on the average 60 mm. long and 15 mm.
wide, just below the inflorescence; flower* borne in small cymes grouped in a
rather lax corymb or panicle; bracts conspicuous, the lowermost similar to the
upper leaves; sepals narrow, 9.5 to 13.5 mm. long, united into a tube for
about 1/3 their length, and tipped with an awn 1 to 2 mm. long; corolla-tube
12 to 16 mm. long, usually pubescent, the lobes obovate, about 12 by 8 mm.,
terminally obtuse or somewhat apiculate; corolla-color phlox-purple through-
out some colonies, near-white throughout others, both extremes together with
intermediate hues appearing in still other colonics; eye often pale, and marked
with 2-3 conspicuous striae at the base of each lobe, ranging in hue from
tyrian rose through purple to violet; stamens rather short, the anther-tips
lying below the corolla-tube orifice at average depths of 2, 2.5, 4, 6, and 7 mm.
respectively; styles short and about half -united, the compound style averaging
1 mm. in length, and the stigmas slightly longer or rarely shorter.

Caulibus crassiusculis, simplicibus vel pauciramosis, glandulosis vel rarius
glabris; foliis superioribus oblongo-Ianceolatis vel ovatis, basi cordiformi semi-
amplexicaulibus; scpalis angustis, longe aristatis, circa usque tertiam partem
longitudinis tubuliformi-connatis.

Type specimen: in herbarium Academy of Natural Sciences of Philadel-
phia, collected by Edgar T. Wherry 2 miles south of Mountainburg, Crawford
G)unty, Arkansas, April 29, 1934.

The usual habitat of this plant is in humus-rich soil on a wooded hillside,
the reaction being low subacid — active acidity 10 to 30, pH 6.0 to 5.5.
Flowering begins in mid-Spring and continues into early Summer. The flowers
are often sweet-scented, with a fragrance resembling that of jasmine, and arc
apparently cross-pollinated by moths as well as butterflies.

Phlox pilosa ozarkana is known to have been collected in the following
states and counties:

Arkansas: Benton, Carroll, Crawford, Logan, Lonoke, Madison, Nevada,
Newton, Pope, Sebastian, Washington, and Yell. Louisiana: Caddo, an
extension well out into the Coastal Plain. Missouri: Barry, Carter, Iron,
Jefferson, McDonald, and St. Louis. Oklahoma: Cleveland, Lc Flore,
Mc Curtain, and Muskogee.

On the accompanying map the boundaries of the Ozark, Ouachita, and
Coastal Plain physiographic provinces are shown by heavy lines. Solid dots

AN OZARK VARIETY OF PHLOX PILOSA

415

represent county records of typical material, open circles a few localities of
intermediates between this and the widespread variety (virens) .

At first sight specimens of this Phlox look decidedly different from the
widespread phase of P. pilosa. The frequent presence of glandular pubescence
well down the stem and leaves, the thickish and often branched stem, the
thinner and broader leaves and the long narrow calyx-lobes give it a unique
aspect. These are, however, all rather superficial and non-fundamental char-
acters, and moreover both within and around the margins of the Ozark region

Fig. 2. Distribution of Phlox pilosa ozarkana.

intercradations with ordinary P. pilosa yirens arc of frequent occurr«ice.
Were an attempt made to draw up a dichotomous key for separating them,
every single characterization would have to be restticted by introducing
"normally!" "usually," "except in intermediates," etc. They may be contrasted,
in geographic relationship, as follows:

Phlox pilosa virens:
Stem tending to be slender, un-
branched, and eglandular, and upper
leaves to be eglandular, lanceolate
and truncate; sepals moderately nar-
row and long-awncd; range chiefly
eastern.

Phlox pilosa ozarkana:
Stem tending to be thickish,
branched, and glandular, and upper
leaves to be glandular, ovate and
cordate; sepals decidedly narrow and
long-awned; range chiefly Ozarkian.

Specimens of the new variety have been found in some herbaria labelled
Phlox drrartcata, and it does a^ree with that in the thickish i\^^^f^JJ'^^
but there the resemblance ends: P. dtrartcata has much better developed per-
sistent sterile shoots, fewer stem-nodes, shorter leav^ which ^^ n^lh^^J^e
markedly wider upward, broader sepals, consistently glabrous corolla-tube, and
more violet corolla-color.

The distribution of its several varieties points to the Ozark region having
been the ancestral home of Phlox piloia, and except for its narrow sepals,

416

THE AMERICAN MIDLAND NATURALIST

variety ozarkana might be taken to be the ancestral phase of the species. As
however, foliaceous sepals seem likely to be a primitive character, and several
of the outlying varieties have this character well-developed, it is considered
probable that the real ancestor of the group has become extinct.

A more northern variety, fulgida, and two more eastern ones, amplexicaults
and detonsa, have been discussed in the papers already cited. Westward from
Ozark region, there appear to be still other varieties developed. One re-
sen^ling the widespread variety virens, except in having more numerous nodes
and so looking much more leafy, barely enters the western borders of this
region, but is apparenriy more widespread in Texas, where it passes into the
plant named Phlox aspera by E. Nelson. While I have obtained a slight field
acquaintance with these more western varieties, it is not my intention to
describe them, because that can be better done by someone who can collect
them more extensively. At fMresent, Miss Eula Whitehouse is engaged in
their study.

Thanks are hereby extended to Miss Caroline Dormon for guidance to
localities of this Phlox in dddo Parish, Louisiana, and to Professor Dwight
M. Moore of the University of Arkansas for similar aid in locating a number
of stations in that State.

REFERENCES
Wherry, Edgar T. 1929— The eastern subulate-leaved Phloxes. Bartonia 11:5-35.

1931— The eastern short-styled Phloxes. Bartonia 12:24-53.

1932 — The eastern long-styled Phloxes, part 1. Bartonia 13:18-37.

1932— The eastern long-styled Phloxes, part 2. Bartonia 14:14-26.

1933 — The eastern veiny-leaved Phloxes. Bartonia 15:14-26.

1935 — Supplementary notes on the eastern Phloxes. Bartonia 16: — in press.

Dept. of Botany,
University of Pennsylvanu,
Philadelphia, Pa.

Reprinted from the Journal of the Southern Appalachian Botanical Club

1:12-15, 1936
1:32-35. 1936

•i

Polemoniaceae of the middle Appalachian region'

Kdgar T. Whkbry

Fourteen members of this plant family are known to occur na-
tive in the nudaie Appalachians, 12 belonging to the genus Phlox and
2 to Polemonium. The features of these and accounts of their dis-
tribution have already been published by the writer in Ba.lonia and
elsewhere, but it seems worth while to call attention to certain gaps
iu our knowledge concerning them, which local collectors may be

able to fill. . , . 1 u „*

1 Moss-Phlox, P. si'bulata L. Characterized by decumbent

woody stems, numerous subulate ,)ersistent leaves, and long-pedicelled
flowers with notched corolla-lobes, in early spring. Crows on rock
ledges and gravelly slopes nearly throughout this region, also further
north and west. Palls into 3 geographic varieties or subspecies:
ciliata (Brand) Wherry with pubescence dominantly eglandular
(glandular only in occasional mutants) and often deep purple eorol-
las- brittonii (Small) Wherry with pubescence dominantly gland-
ular (eglandular mutants) and small white to lavender corollas; and
var austmlis Wherry with likewise glandular pubescence, but larger
and mostly purple flowers. Along the Virginia - West \ irg.n.a
boundary and nearby these segregates seem to merge, ^nd we need
statistical .studies as to the percentage of glandular and eglandular,
large and small-flowered, and purple and lavender corolla-bearing

plants, within individual colonies. , , , , .

2 Blue-Phlox, P. mvABiCATA L. Characterized by decumbent

somewhat stolon-forming stems with persistent elliptic leaves with
tall flowering shoots bearing cymes of short-pedieelled flowers having
glabrous corolla-tubes, lavender often notched lobes, and very short

.Contribution from the Botanical Laboratory and Morris Arboretum of
the University of Pennsylvania.

14

stanions and styles. Si)i*in«^, in open woods. Only one geographic
variety, canadensis (Sweet) Wherry, occurs here. Field notes on its
variability in such featui'es as corolla lobe-shape, notching, and color-
ing would be of interest.

3. Downy Phlox, W pilosa L. Characterized by erect shoots
bearing lineai' to lanceolate deciduous leaves, and infloi*escence much
like that of P. divaricata except that corolla-tube is often pubescent,
the lobes entire, and theii- color varying from lavender to white and
to bright purple. Late spring. While common in the central states
and in the southernmost Appalachians of Alabama, this species be-
comes exceedingly rare northeastward, being almost unknown in the
mid-Appalachians. The J^irsh hei-barium assembled in the earlv
1800's includes a specimen labelled "Staunton", but it is not known
to have been found there since. Search should be made for it in
meadows and open woodlands, so that a better picture of its actual
range can be obtained.

4. Hairy Phlox, P. amokna Sims. Shoots becoming decum-
bent, their small ellijitic oblong leaves partly persistent; inflorescence
very compact, the bi'acts forming an involucre, covered with coarse
eglandular hairs, and bearing small bright i)urple flowers. Late
spring. This southern species extends up the Ai)palachians as far as
middle North Carolina, but its exact limits are unknown. Old man-
uals report it in ''Virginia" but no specimens from that state appear
to be presei-ved in any herbariuuL There is a published report from
Payette County, West Virginia, again unsupi)0]ted by specimens,
never fully identified, and considered by its collector as adventive
anyway. It should be sought in open woods.

5. Crekpin(; Phlox, P. stolonifkra Sims. Stems markedly
stoloniferous, with pei'sistcnt s])atulate leaves; inflorescence few-
rtowei-ed, the sepals well united, and corolla-tube, stamens and styles
all elongate; flowers large, bright purple, in late spring. Reaches its
maximum development in West Vii-ginia, and eft'oits might well be
made to find it in every county. (Jrows in W(K)dland litter.

6. Sw'ORDLKAF Phlox, P. BucKLKvi Wherry. This is in several
respects the most remarkable of our eastern I'hloxes. It was first
collected in 1838 by S. B. Buckley, but lay unnoticed in his herbarium
for over 75 years, and only received recognition as a species in 1930.
Its range is the most restricted of all the .species, being so far as knowTi
limited to an area less than 100 miles in diameter, centering east of
the Virginia- West Virginia boundary around Covington. And it is

■

i

i

i

15

the only species as yet found to be tetraploid, that is, to have double
the usual number of chromosomes in its cells. It can be recognized
by the basal rosettes of sword-shai)ed evergreen leaves from which in
late spring arise flowering stalks, with conspicuously glandular-
pubescent herbage and cymes of showy purple flowers; the latter
have well-united sepals, densely glandular corolla-tubes, and elongate
stamens and styles. It grows in open woods, but chiefly or wholly m
those situated at the base of a shaly slope, where the soil contains
slabs or flakes of the shale rock.

The following stations for it are known to date: VIRGINIA,
Alleghany Co.: 1 mile north of Alleghany Sta. : VA miles southeast
of same ; 1 mile southwest of Longdalc Furnace. Aiujusta Co. : Deer-
field valley. Bath Co.: 4 miles northwest of ^Mountain Grove; just
northwest of Hot Spi'ings; Flag Rock trail 1 mile southeast of Warm
Sprin-s ■ 3 miles northwest of Bacova June. Craig Co. : 4 miles south-
east of Craigs Creek. WEST VIRGINIA. Creenbrier Co.: % mile
southeast of Caldwell ; V4 mile south of White Sulphur Springs Sta-
tion, perhaps the type locality; V/. miles southeast of White Sulphur
Springs Poealimias Co. : 4 miles southeast of Driscol. Unfortunate-
ly some of these colonies have been destroyed by vandals, by grazing

ft

animals, etc.

{To he continued)

IRREGULAR PAGINATION

Polemoniaceae of the middle Appalachian region, pt. 2.

Edgar T. Whi:rry
{Continued from p. 15)

7. Mountain Phlox, P. ovata \j. Characterized by the pres-
ence of short decumbent sterile shoots with persistent elliptic leaves;
flowering shoots, produced in lale sprinj;, having very few (usually
4) nodes below inflorescence; sepals unusually long, 8 to 12 mm.;
rather large bright purple coi'ollas, with stamens and styles nearly
as long as the tube; and herbage glabrous except for minute pubes-
cence on pedicels. Best developed in North Carolina, but extend-
ing north on both sides of the Virginia-West Virginia line to central
Pennsylvania. Two varieties have recently been segregated, latifolia
(Michx.) Wherry, which is widespread, and pulclira Wherry, which
is known thus far only in Walker County, Alabama. The latter has
longer shoots and beautiful pink flowers.

8. ThiCkleaf Phlox, P. Carolina L. Distinguished from the
next-preceding species, with which it has often been confused, by hav-
ing elongate but rarely persistent sterile shoots ; flowering shoots with
more numerous nodes, — often 10 or more — and sepals shorter, —
6 to 11 mm. ; and stems often more or less pubescent. Three varieties
are distinguishable: iriflora (Michx.) Wherry, ranging from North
Carolina to northern Virginia (or possibly Maryland) and blooming
in Spring; altissima (Moench) Wherry, Georgia to North Carolina,
late summer; and heterophylla (Beauv.) W^herry, centering around
Alabama and blooming in early summer. More data are needed con-
cerning the first of these, var. iriflora. The northernmost point at
which it is known to have been collected in recent years is in Bath
County, Virginia ; it has thus far not been recorded from West Vir-
ginia. Nearly a hundred years ago, in a list of Maryland plants by
Aikin, there was included a Phlox ''revoluta" which so far as the

1

33
description goes seems most likely to have been this variety. Its
locality is supposed to iiave been the Appalachian rid(?es near Fred-
erick and it should be sou-ht .'speeialh- between that place and the
Virginia region above mentioned as its present limit. I'hc occur-
rence and significance of colonies in which the plants cond)ine the
characters of two varieties also need study.

9 Smooth Phlox, P. gi.abkrrima L. Though not known ui the
middle and northern parts of the Appalachians, this relative of the
next-preceding species does .'nter this province southward, m upper
Georgia and Alabama, bloon.ing in sunnner. Only one ot the two
recognized varieties is concerned, - var. mdampunfoha (Sabsb.)
Wherry, the other being n.ore western. 11 is to be distinguished from
P Carolina bv the rarity of .lecun.bent si, -rile sh..ots, Ihe n.ore num-
erous nodes - fre<,uently ir. - below the inHorescenc... th,. usually
narrower leaves in con.bination with a short calyx, ran'ly over 9 nun.
long (in r caroliva altixsim, espe.Mally short caly.K l.en.g cor-
relate.1 with decidedly broad leaves) . Both /'. c.roli.u, an<l /'. fjM><r-
rima are rather variable in leaf-si.... an.l son,e occurrences of one may
have been misinterprete.l as representing the other, so n.ore reco..ls
of variability within single colonics a.e needed. For cxan.ple a
PMox of this relationship with leaves as linear as n. any i • flM>n-
ri,ua ha.s been repeatedly collected in the vicinity "Y'-J'^ "^ '^•
Burke Co., North Carolina ; because of a s.i.all number ..f nodes I
have classe,! it as P. Carolina, but until this oc-nrrence receives further
studv this identification can not be regar.le.l as final.

"lO Mkapow Pi.i-ox, 1>. ^.A.M•,,^T^ 1. When typically developed
this showv Phlox has so n.any disling.iishing features that there is
no difficuify in recognizing it. The rootstocks a,e slender, and cin'v
up at the end into a stout, pnrple-spofte.l stem which may reach a
height of several feet, and the often .lense inflorescnce - ".vL-lnc
rather than eorvmhose (as in all the preceding species becoming
onicL in especi lly vigorous plants. There are two varieties, o.lorala
sTet) Wherrv which ranges from Mis.souri and the moun ains o
S,::! northward, being characterized by relatively few ai.un
15) nodes and by blooming in late spring; .uapyraMs (Smith)
Wherry, Tennessee and North Carolina to the -"'hern edg of
Pennsylvania, with up to 30 nodes, a summer-bloonu r. Fhe s ua-

^rS however, not so simple, for in --'"-'' "nn-IIitoo::
wise well-differentiated varieties appear to <^^^'^r;'J^;'^^
another but also into phases of P. carohna. While only ^ar,et>

1

i

I

34

pjfrdmidalis appears to be known in North (*aroliiia, collections macle
thus far would indicate vai'. odoraia to be the only representative of
the species in the Appalachians of southwestern Virginia, Avhile in
West Virj^inia both vai'ieties as well as intermediates are common.
Kvidently moi-e collecting is needed to work out the significance of
these complex I'elations.

11. Broad-leaf Phlox, P. amplifollv Britton. The final sec-
tion of the genus Phlox in this region comprises two species, dis-
tinguishable from all the i)receding ones by their I'clatively large
leaves with jirominent areolate veins and hispid-serrulate margins,
and i)ale oj" white-coloi'cd anthers. Both bloom in late summer. The
first of them, here taken up, has few nodes, opposite often coarse-
bristly leaves, glandular inflorescence-hei'bage and glabrous corolla-
tube. It is rather rare, having been collected in but four Ap-
palachian counties each in Tennessee and Noilh Carolina, and one
(Lee County) in Virginia; there is also a doubtful record from Hawks
Nest, Fayette County, West Vii'ginia. Beyond this region the species
I'anges to central Alabama and to Missouri, but it is apparently no-
where common. Search for new stations for it should certainly be
made on wooded slopes in the mid- Appalachians.

12. Veiny-leaf Phlox, P. panicuu\ta L. This is the ancestor
of the common perennial gai-den phlox, and some of the records of it
may represent garden escapes. It is, however, unquestionably native
at many j)laces in the Appalachians as fai" north as central New
York, also ranging west to the northeast coiiier of Kansas. From the
next-preceding sfx'cies it is distinguished by more numerous nodes
with the leaves only sub-o])positc, the leaf-surface being usually glab-
rous at least above, the inflorescence-herbage being normally egland-
ular-pubescent, and the corolla-tube pubescent. It has been suffi-
ciently collected for its range to be well understood, although further
study of occasional colonies where it has taken on one or more of the
characters which usually set off P. ampli folia (especially in the high-
er Smokies) would be worth while.

One other genus, Polrmonium itself, is represented in the middle
Appalachians, but by only two species. The more primitive of these,
known as Poij£MONium van-bruntiae Britton, is of northern rela-
tionship, ranging from southeastern West Virginia across Marj^land
into Pennsylvania, New York, Vermont, and the west border of New
Hampshire. It grows in swamps and blooms in summer. Dis-
tinguishing features are tall habit, thyrsoid inflorescence, and rather

i

35

large dark violet flowers with subi)ai'allel-declincd exsei'ted stamens
healing golden anthei's. The southernmost colony of this plant thus
far known is in the Cranberry Glades of Pocahontas (^ounty. West
Virginia; it has also been found at one or more localities in Preston
and Tucker, in that state, and (larrett in :Marylanil, but seems in gen-
ital to be decidedly rare. Perhaps systematic search will disclose

additional stations.

The 14th and last member of the Polemoniaceae on the list is
Polemonium reptans L. This blooms in si)iing, and appears in a
variety of habitats, including deep or open woods, thickets on alluvial
flats, rocky slopes, meadows, swami)s, and even around the margins
of bogs, it is dwarfer than the preceding, and bears ojx'n |)anicles of
light violet blue flowers with irregularly decumbent stamens tipped
with cream-colored anthers. The Spring Polemonium has a wide
range over the eastern half of the Tnited States, (between longitudes
75 and 100 degrees west) but apparently becomes rare in the middle
Appalachians. I have seen specimens from only about five mountain
counties each in Virginia and West Virginia, and there is but a
single record, in Haywood (^ounty, for all of North Carolina, in
this case also further collecting is obviously to be desired.

Polemonium and Polemoniella in the Eastern States

Edgar T. Wherry

In contrast with the genus PhloXy already treated in these pages, the other
genera of the family Polemoniaceae are but sparsely represented east of the Miss-
issippi River. Two species of Polemonium and one each of Collomia and of
Ipomopsis comprise the total of native species, while less than half a dozen other
members of the family appear as occasional adventives. Before passing to the
study of the far more numerous western representatives, however, the assembling
of data on these eastern ones seems worth while, and the results will be presented
in two papers, of which this is the first. Polemonium and allied genera are char-
acterized by their pinnately compound alternate leaves with relatively large
elliptic leaflets ; by their wholly herbaceous, accrescent calyx ; by their campanu-
late (or rarely funnelform) corolla, of which the hue is blue-violet or violet-blue,
(exceptionally pink or even yellow) and by their blackish obscurely polyhedral
spindle-shaped seeds. As in Phlox, the corolla is actinomorphic, but the stamens
are distinctly zygomorphic; and there are three fused carpels with the styles
united up to their stigmatic tips. The segregated minor genus Polemoniella is
distinguished by annual duration and greatly reduced inflorescence, the incon-
spicuous grayish flowers having short, practically equal stamens, and being self-
pollinated.

POLEMONIUM [TOURNEFORT] LINNfi
This genus name, the derivation of which has never been adequately explained,
was used by Tournefort ^ in 1700 for a plant which had been classed by other
writers as a Valeriana. Linne « defined it more fully in his Genera Plantarum,
being acquainted at that time with a single European species. In the first edition
of his Species Plantarum * he listed three, and in the second edition «» as many as
five, although all but the first two of these are now placed in other genera. A
key' for distinguishing the two species recognized in the eastern United States is

given on the following page. .,,.,. ixi. u

Various common names are ascribed to this genus in the literature, although
few of them are in actual use. Perhaps the most frequent is " Greek Valerian "
a translation of the term Valeriana graeca applied to some similar plant by early
herbalists. Another often seen is " Jacob's-ladder," in reference to the ladder-

1 Contribution from the Botanical Laboratory and Morris Arboretum of the University
of Pennsylvania.

2 Inst. Rei Herb. 1: 146; 2: pi. 61. 1700.
8 Genera Plantarum ; 46, 1737.
♦Species Plantarum: 162, 1753.

» Species Plaptarum, ed. 2. 1 : 230, 1762.

IRREGULAR PAGINATION

6

BARTONIA

like arrangement of the leaflets. The name most used by laymen is ' Bluebells,
but as this is also applied to species of Campanula, Mertensia, etc , it is not
sufficiently distinctive. Possible modifications would be " Fernleaf Bluebell or
" Bluebell Valerian," but, like the occasionally heard " False Forgetmenot these
are too cumbersome. The layman who already uses without hesitancy such
names as Chrysanthemum, Delphinium, and Geranium should have no difficulty
in adopting the genus name Polemonium itself for a " common name."

POLEMONIUM : KEY TO EASTERN SPECIES
^tPm relatively strong 75 to 125 cm. tall; principal leaves divided into 11 to 21 leaflets; in-
^'^rescenTe a ^^^^^^^^^^ corolla-color rather deep violet; stamens dechned,^s^bpa^^^^^^^

exserted ; anthers orange-yellow " * ■. V V '•' '/ ' V'/ i c i fl^f. . i«

Stem relatively weak, 25 to 75 cm. tall; principal leaves divided mto 7 to 15 leaflets, m-
^CescenJra'^rde'Wnicle corolla-color rather light blue-violet; stamens -rog"lariy d.-

cumbent, included; anthers cream-color ^- • *"

Barton I A, No. 17

Plate 2

Fig. 1. Distribution of Polemonium vanbruntiae.

1. Polemonium vanbruntiae Britton. Tall Polemonium. Plate 2, Fig. 1.

History— This Polemonium was apparently first observed in Schoharie
County, New York, by a Dr. E. C. Howe about 1860, and was included in the
list of addenda to the fourth edition of Gray's Manual ' under the name
P. coeruleum. In subsequent years its known range gradually increased, although
it was long referred to that European species. Its distinctness was first urged
by Britton =^ who in 1892 named it in honor of Mrs. Van Brunt. The original
form of the name was Van-Bruntiae, but the writer prefers condensation and
decapitalization of all species names.

Geography.— 'During the Glacial epoch this plant evidently survived in the
uplands of western Maryland and adjoining states, and subsequent to the melting

1 Man. Botany N. U. S., ed. 4: xcvi, 1863.

2 Bull. Torrey Bot. Club 19: 224, 1892.

Fig. 1. Polemonium vanhrunliae Britton.
Georgetown Station, Madison County, New York.

Fk; 2. P(tl( I'Kniiinii Kjilnns JAunv.
Kn«t of Monnlain Lake, Gam-tt County, Maryland.

6

BARTONIA

like arrangement of the leaflets. The name most used by laymen is " Bluebells,
but as this is also applied to species of Campanula, Mertensia, etc., it is not
sufficiently distinctive. Possible modifications would be " Fernleaf Bluebell " or
" Bluebell Valerian," but, like the occasionally heard " False Forgctmenot " these
are too cumbersome. The layman who already uses without hesitancy such
names as Chrysanthemum, Delphinium, and Geranium should have no difficulty
in adopting the genus name Polemonium itself for a " common name."

POLEMONIUM : KEY TO EASTERN SPECIES

Stem relatively strong, 75 to 125 cm. tall; principal leaves divided into 11 to 21 leaflets; in-
flore^encea'^narrow 'panicle; corolla-color rather deep violet; stamens declined subp^^^^^^^^^
exserted; anthers orange-yellow '."."-"" V*/ . r i a , • '

Stem relatively weak. 25 to 75 cm. tall; principal leaves divided mto 7 to 15 leaflets; m-
fllTrescence a"^ wide panicle; corolla-color rather light blue-violet; stamens "-cgularly de-
cumbent, included; anthers cream-color £. r. rt^j

Haktuma, No. 17

Plate 2

Fig. 1. Distribution of Polemonium vanbruntiae.

1. Polemonium vanbruntiae Britton. Tall Polemonium. Plate 2, Fig. 1.

His^ori/.— This Polemonium was apparently first observed in Schoharie
County, New York, by a Dr. E. C. Howe about 1860, and was included in the
list of addenda to the fourth edition of Gray's Manual ^ under the name
P. coerulcum. In subsequent years its known range gradually increased, although
it was long referred to that European species. Its distinctness was first urged
by Britton ^ who in 1892 named it in honor of Mrs. Van Brunt. The original
form of the name was Van-Bruntiae, but the writer prefers condensation and
decapitalization of all species names.

Geography.— During the Glacial epoch this plant evidently survived in the
uplands of western Maryland and adjoining states, and subsequent to the melting

1 Man. Botany N. U. S., cd. 4 : xcvi, 1863.

2 Bull. Torrey Bot. Club 19: 224, 1892.

Fk;. 1. Pole mount tn v(tnl)niiihiir Britton.
Georgetown Station, M.idison County. N« w ^'o^k.

Fic 2. /'"/' iiioiiKini I'l I'ltiHs l.iiinv.
Ea>i of Mount nil L;.k. . ( Ininit (*ou..iv. Mi.iyhn.l.

^4

y

i

'.■«

tOLEMONruM AND POLEMONIELIA •

away of the last ice sheet succeeding in migrating into the glaciated territory as
far as northern Vermont (or New Hampshire). The state and county list is:
Connecticut: Known only in Litchfield County (inadvertently marked too

far north on the map.) , j t-

Maryland: Occasional in Garrett' County ; reports from lowland counties are

based on misidentifications.

[New Hampshire: Reported from Grafton County, but doubtful.]
New Jersey: Specimens are widely distributed in herbaria from a swamp two
miles north of Washington, Warren County, although search for the plant there
in recent years has proved unsuccessful. Mistakenly reported from Morris Co.

New York: Reaches its maximum development in this stat«, being known
from over 20 localities distributed through 10 counties: Chenango, Delaware,
Greene Herkimer', Lewis Madison', Schoharie, Sullivan, Tioga and Ulster.

Pennsylvania: Shows a curiously interrupted distribution, havmg been col-
lected in Berks, Somerset', Sullivan, and Wayne counties only.

Vermont: Native in Addison and Windsor counties, escaped elsewhere.
West Virginu: Found at high altitudes in 3 eastern counties, Pocahontasr,
Preston, and Tucker', the first being its southernmost known station.

Ecology -This Polemonium is a swamp plant, its rootstocks creepmg through
hummocks of mosses, sedges, and shrub-bases. Even though the water beneath
the surface may be alkaline, the humus into which its roots extend is almost
always subacid or low mediacid. In successions '*«««"?'« V"- TtTflnw^
position, dying out as climax forests develop. Its dark purple-violet^ flowers
are borne chiefly in mid-summer, and are cross-pollmated by bees.

Taxonomy.-The original assignment of this plant to P. coeruleum Linne is
easily understood, for it is closely related to that European species, having evi-
dently descended from the same circumboreal ancestor Indeed, few of the
characters considered by Britton to justify its segregation as an independent
species are really distinctive. The European plant, when growing in moist seal,
may develop a horizontal rootstock much like that in the American one; the
SLtfps vary from rounded to mucronulate in both. Supposed differences in
accrescence o7calyx do not appear to be significant, as when fully fertilized both
sneles develop rather numerous seeds, leading to marked enlargement of the
ova^and coLquent distension of the calyx. So far as I can discover, from
IdT of herbarium sheets as well as living material of both the chief differences
consL in the American plant having slightly broader and fewer leaflets, and
onTr more nearTy parallel stamens. In the interest of simplicity their specific
dUtinctrss is here maintained, pending further work on related west-American

plants.

X In Mae« and Paul'^ Dictionary of Color ranging from 42 H 8 to 42 J 10 and to 43 J 9,
that ii, a moderately deep and slightly grayed violet.

sf^

8

BARTONIA

POLEMONIUM AND POLEMONIELLA

9

2. Polemonium reptan* Linne. Low Polemonium. Plate 2, Fig. 2.
History.— In the first edition of the Flora Virginica, published in 1739, Gron-
ovius 1 referred to a " Polemonium foliis pinnatis, radicibus reptatricibus. Clayt.
n. 249." According to the correspondence assembled by Darlington,^ roots of
what was evidently the same species were sent by John Bartram to Peter Collinson
in 1740. The latter at first questioned its distinctness from the European " Greek
Valerian ", but later noted differences between them in stature and blooming-time.
Linne « cited the Gronovius record under the European P. coeruleum in the first
edition of the Species Plantarum; later, however, he saw a color-plate of it
among the illustrations of plants in the Gardeners' Dictionary by Miller,* and,
realizing their distinctness, gave the American species the name it now bears, in
the 1759 edition of his System."* As the plant illustrated had been sent by
Clayton to Gronovius, the type locality of the species is to be taken as eastern

Virginia.

Not feeling bound by the principle of priority, Salisbury « proposed the appro-
priate name P. humile, but the rules of nomenclature prevent its adoption. On
the other hand, one published by Rafinesque,^ P. quadriflorum, is meaningless
and well deserves rejection. Many years later Brand® based a variety p
macrophyllum on a specimen showing abnormally large leaflets, and Wetzstein *
termed the albino mutation P. reptans album.

Geography. — In contrast to the other eastern species, Polemonium reptans is
both common and widespread, having a range nearly as vast as that of Phlox
divaricata. Though rare in the Coastal Plain, it is abundant in piedmont, mon-
tane, and interior physiographic provinces, from northwestern Georgia to
Arkansas, easternmost South Dakota, and central New York state.

Alabama: Known in 6 Appalachian counties: Colbert, Cullman, Madison,
Marshall^ Morgan^ and Tuscaloosa, the last a southern limit.

Arkansas: Probably occurs nearly throughout the northern part, having been
observed by Nuttall in 1819 along the Mississippi river, and collected in recent
years in Benton, CarrolK, Crawford, Cross, Franklin, Madison, Marion, Newton,
Polk, and Washington^ counties.

[Connecticut: Escaped from cultivation in 4 counties: Fairfield^, Hartford*,
Litchfield*, and New Haven<^.]

Delaware : Frequent on the piedmont in New Castle^ County.

1 Flora Virginica: 22, 1739; ed. 2: 29, 1762.

2 Memorials Bartram and Marshall : 138, 155, 1849.
« Species Plantarum : 162, 1753.

♦Figs. Plants Miller's Gardeners' Diet. 2: pi. 209, 1758.

»Systema Naturae, ed. 10. 2: 925, 1759.

•Prodromus Stirpium, etc.: 125, 1796.

T Atlantic Journal : 177, 1833.

8 In Engler's Pflanzenreich IV. 250: 33, 1907.

•Proc. Ohio State Acad. Sci. 4: 361, 1906.

f ift

'-■J •.

>*•

*j»%

-■i *s

S= S5r.^;^rr ' S v^t^te-SU" ana ..-

counties: Champaign, Cook, Du Page, «^"^_^^'"' ^^^ ^^,5^^^^^^ M;ntgomery,
Jefferson, Jo Daviess, Kane, Kankakee ^f «^La ^dle, McH y, ^^^.^.^^_
Ogle, Peoria, Randolph, Rock Island, St. Clair, scnuy ,
Wabash, Warren, Will, and Winnebago.

Fia. 2. 'Distribution of PoUmonium reptans.

i™.. More ^fy;ou..^rc::ztt.Si:: Detr : -^-^-

ford. Brown, Carroll Cas^ Clark, ™rd , ^.^^^ ^^^^^^

Kalb, Delaware, Dubois, ^Ikhart^ ^^^J^^^ *°"h,„, a, j^y_ j,ff,,,o„,

Greene, Hamilton, Hancock, H^"^'^^""' Jf^' ^^rtl La;rence, Madison, Marion,
Jennings, Johnson, Knox, Ko^msko ^ake.^a^rte, ^^^^^^^ ^^^^^

Marshall. Miami, Monroe, Montgome>7, Ne^"- ^ g^ j^^^p^. Scott,

Parke. Perry^ Pike, Porter, Posey »^^^ J.„ Vermilion, Vigo,

raih,' r^'^:^^^^^-^^^^ ^^^^ -' ™"' '

total of 73 counties. records: Allamakee, Benton, Black

low.: Widespread, there be ng^^^^^^^^^^ ^^^^.^_ ^^^^^ ,„,,,„„,

Hawk. Cerro Gordo. Cherokee, iJeca , . j^i^^

Mahaska, Muscatine, Poweshiek, Scott, btory, ana

10

BARtOKiA

Kansas: Barely enters the state, in Cherokee and Doniphan counties.

Kentucky: Probably present throughout, though known to have been collected
in but 18 counties: Anderson, Bath, Carter, Christian^ Edmonson, Estill, Fayette,
Franklin, Graves^ Henderson, Jefferson, Kenton^ Lyon, Morgan, Owen^ Rowan^

Shelby and Warren.

Maryland: Known in 8 counties, all above the Fall Line: Allegany, Balti-
more, Carroll, CeciK, Garrett^ Harford, Howard^ and Montgomery^

[Massachusetts: Escaped in Hampshire*^ County.]

Michigan : Reported only in Berrien and Washtenaw counties.

Minnesota: Extends into several of the southern counties: Brown, Douglas,
Fillmore, Goodhue, Hennepin, Houston, Olmsted, Scott, Wabasha, and Winona.
The second-named is the northwestemmost station for the species.

Mississippi: Rare, and limited to the northeastern counties: Lafayette, Pren-
tiss, Tippah, and Union.

Missouri: Common throughout, with 24 counties represented by collections:
Barry, Boone, Butler, Carter, Cass, Clay, Cole, Franklin, Greene, Iron, Jackson,
Jasper, Jefferson, Johnson, Lincoln, McDonald, Ozark, Phelps, Ralls, St. Charles,
St. Francois, St. Louis^ Shannon, Taney, and Washington.

[New Hampshire: Escaped in two or three counties.]

New Jersey: Barely enters along the Delaware valley, there being records for
6 counties: Burlington, Camden, Hunterdon, Mercer, Salem, and Warren.

New York: Occasional westward, in Allegany, Cattaraugus, Chautauqua,
Chemung, Erie, Livingston, Steuben, Tioga, and Wyoming counties.

North Carolina: Reported only from Haywood County.

Ohio: Occurs throughout, there being 53 counties in the list: Adams, Allen,
Athens^ Auglaize, Belmont, Butler, Clark, Clermont, Columbiana, Coshocton^
Crawford, Cuyahoga, Darke, Defiance, Delaware, Erie, Fairfield^ Franklin,
Gallia, Geauga, Greene, Guernsey, Hamilton^ Hardin, Harrison^ Highland^
Holmes, Huron, Jackson, Lawrence, Licking, Lorain, Lucas, Mahoning, Medina,
Mercer, Miami, Monroe, Montgomery, Morrow, Perry, Pike, Portage, Preble,
Richland, Ross, Seneca, Stark, Van Wert, Warren, Washington, Wayne, and

Wyandot.

Pennsylvania: Fairly common, except in central and northeast portions;
county list: Allegheny, Armstrong, Bedford, Berks, Bradford, Bucks, Butler,
Cambria, Chester^ Clarion, Crawford, Cumberland, Delaware^ Elk, Erie, Frank-
lin, Huntingdon, Lancaster, Lawrence, Luzerne, Lycoming, McKean, Montgom-
ery^ Philadelphia, Potter, Somerset, Westmoreland, and York^

[South Carolina: Elliott's report of it in " vallies " refers to Georgia.]

South Dakota: Reaches a western limit in Clay County.

Tennessee: Though stated by Gattinger to occur " over the state," specimens
have been seen from but 6 counties: Blount, Dyer, Jefferson, Knox, Madison, and
Shelby, 3 of these lying near the eastern, and 3 near the western, end.

[Vermont: Escaped in Addison*, Caledonia*', and Rutland* counties.]

polemonium and polemoniella

11

/*» •

si>

M>B m.

,V3

Virginia: Widely distributed, but known to have been collected only in 7
counties: Bath, Fairfax, Highland^ Russell, Smyth, Wythe, and York.

West Virginia: Occasional along the Ohio river and in a few upland counties:
Cabell, Gilmer, Monongalia, Ohio, Preston, Randolph, and Wood.

Wisconsin: Common in the southern half of the state, in Adams, Brown,
Buffalo, Columbia, Dane^ Dunn, Eau Claire, Grant, Green, Iowa, Jefferson, La
Crosse, Marathon, Milwaukee, Richland, Rock, Sauk, Trempealeau, Vernon,
Walworth, Waukesha, Waupaca, and Wood counties.

Ecology.— One reason for the wide dispersal of Polemonium reptans is its
extraordinary adaptability. Although reaching its maximum luxuriance in cir-
cumneutral soil on alluvial flats, it also grows in fertile loam over limestone, in
sterile, more or less acid clay, in the decidedly acid litter in upland oak woods,
and even, at least marginally, in mediacid sphagnum bogs. Moisture may be
abundant throughout the year, or for but a few weeks in spring. In its favorite
situations there is usually plenty of light early in the year, but it can adapt
itself to all variations from full sun to deep shade during the rest of the growing
season. Occasionally a pioneer in meadow lands, it becomes abundant in inter-
mediate successional stages, and often remains so into the final cliniax forest.

In a classification according to blooming season, this Polemonium is to be
considered pre-vernal. Shoots begin to arise from the crowns in the Fall, persist
even through winter temperatures below zero F., and resume growth while the
air is still rather cold. The flowers begin to open toward the end of March in
the southern part of its range, or of April northward. Anthesis lasts a few weeks,
and the seeds are soon ripened and dispersed. Robertson ^ noted that insects
land on the anthers of the lower stamens and insert their tongues between the
shorter upper ones. Visitors observed comprised 38 hymenoptera— one named
Andraena polemonii Rob.— and 12 members of other classes.

Variation and taxonomy.— Polemonium reptans is variable in many respects.
There is extreme variation in the size of the plant and of the leaves and leaflets,
while the corolla in some individuals is fully twice as large as in others. Though
it is usually described as glabrous or nearly so, there is a tendency for the stem
and even the leaf-rachises to become sparsely covered with long hairs. This is
perhaps more frequent westward, but is by no means lacking at the eastern
margin of the range. Another tendency is for the leaflets to become confluent,
at least toward the tip of the leaf. Rafinesque gave this as one of the features
of a supposedly distinct species from Arkansas, but it sometimes appears in the
east as well. The corolla is normally of a light violet hue,* although pallid,
albino, purplish and deep violet mutants are occasionally found. In the opinion
of the writer all of these variations should be regarded as of formal rank, and as
deserving no special names.

1 Trans. Acad. Sci. St. Louis 5: 578, 1891. Flowers & Insects: 152. 1928. . ^ . ,

« In Maerz and Paul's Dictionary of Color ranging from 41 B 6 to 41 G 9. or light violet-
blue to blue-violet, and often around Wistaria Violet.

I

12

BARTONIA

FORMS OF POLEMONIUM REPTANS

Stem pilose. Termed the " western form " by Coulter,^ but collected by the writer as far east

as Chester and Delaware counties, Penna. n j r»„«„.,;««oi «nrfViwP«?t-

Leaflets enlarged, up to 6 cm. long. P. r. var. macrophyllum Brand. Occasional northwest-

Lelflets terminally confluent. P. quadnflorum Rafinesque. Rare but widespread.
Corolla pale or white. P. r. album Wetzstein. May develop m any colony.

Cultivation,— Introduced into horticulture by Clayton, through Gronovius,
about 1730, and by Bartram through Collinson, in 1740. Owing to its adapt-
ability to growth under a wide variety of conditions, responds well to cultivation,
thriving alike in the border, the rock garden, and the woodland.

POLEMONIELLA HELLER

Taxonomists have often been criticized for splitting genera on inconsequential
grounds, but in the present case the segregation of Polemoniella from Polemomum
by Heller ' seems fully justified. The new genus clearly represents an evolu-
tionary development from the older one, the changes comprising perennial to
annual duration, compound to simple inflorescence, large showy violet corolla to
small inconspicuous whitish one, long declined unequal to short spreadmg equal
stamens, and style extending beyond anthers to holding stigmas in the midst of
anthers so that cross-pollination by insects has given way to self-pollination.
Three species have been recognized, one native to western North America, the
others to southern South America.

Polemoniella micrantha (Bentham) Heller

Discovered by Douglas in the Columbia valley in the 1820's, the plant was
named by Bentham » Polemonium micranthum in 1845. Peter * in 1891 divided
the genus Polemonium into three sections, placing this species in one named
Polemoniastrum. Thirteen years later Heller (loc cit.) pointed out its differences
from the typical Polemoniums, and made the combination here accepted.

A specimen of this plant collected near Milton, Norfolk County, Massachu-
setts is preserved in the New England Botanical Club collection at the Gray
Herbarium. It may turn up elsewhere, especially in fields where grass seed
from the northwest has been sown. It is, however, unlikely to become a pest,
because it is a diminutive winter annual which flowers in early spring, ripens its
seed in a few weeks, and then vanishes.

The remaining eastern representatives of the family Polemoniaceae will be
discussed in the next number of Bartonia.

1 Cat. Fig. Pits. & Ferns indig. to Indiana: 891, 1900.

2 Muhlenbergia 1 : 57, 1904.

8 In DC. Prodromus 9: 318, 1845.

*In Engler & Prantl Pflanzenfam. 48a: 62, 1891.

/

The Ranges of our Eastern Parnassias and Sedums *

Edgar T. Wherry

PARNASSIA

' Becoming interested in the soil-reaction relations of members df the genus
Pamassia, 1 have during the past few years visited a number of their localities.
On attempting to draw up a field key to the species, so as to make sure as to
which of them each test was made upon, certain discrepancies in the literature
came to light. A report on progress in straightening these out has recently been
made in Claytonia,? but as that mimeographed journal is not widely available,
and as one nomenclatorial change and certain modifications of the key there given
seem desirable, this article is being written.

In the case of one species, P. asarifolia Vent., no nomenclatorial difficulties
appear to have arisen, although its range is inadequately stated in current man-
uals. Confusion of a second, P. grandifolia DC, with other species has sometimes
occurred, but its range is given correctly in recent literature. The other two
species of the eastern United States have been completely misunderstood as to
their geography and nomenclature for over 125 years.

In 1803 Michaux^ described Pamassia caroliniana, giving as its locality " in
nndis Carolinae." Nine years later Sims * applied that species-name to a plant
shown by his colored plate to have short staminodia and greenish ovary ; this was
said, following Michaux, to be " Native of the bogs of North Carolina," although
its collector, Masson, is known to have limited his activities to Pennsylvania and
more northern states. Pursh,* on the other hand, gave P. caroliniana as growing
from " New York to Virginia," thus actually lumping two quite distinct species, —
one occurring in the former, the other, P. grandifolia, in the latter— and using
for them a name referring to a native of still a third state. This confusion has
been perpetuated in all subsequent manuals and floras down to Small's Manual
of the Southeastern Flora in 1933.

While the manual just mentioned was in active preparation, a Pamassia with
large flowers, elongate staminodia, and white ovary was received from Mr. H. A.
Rankin, who had collected it in the region termed by Michaux " Imdae Caro-
linae," namely, inland from Wilmington, North Carolina. There being no evi-
dence that the quite different plant figured by Sims grows as far south as this
state, a change of current nomenclatural usage became inevitable: Michaux's
name must be applied to the plant for which it was originally intended, — which

1 Contribution from the Botanical Laboratory and Morris Arboretum of the University
of Pennsylvania.

! 2 Glaytonia 1 : 61. 1935.
' 3 Flora Bor.-Amer. 1 : 184, 1803.

♦Curtife's Bbt. Mag. 35: pi. 1459, 1812.
i 8 Flora. Amer. Sept. 1 : 208!, 1814.

.17

IRREGULAR PAGINATION

18 BARTONIA

had meanwhile been named P. fioridana by Rydberg.* This was duly done by
Small - and by Alexander ,« the latter recommending the adoption for the northern
species — that figured by Sims — of the name P. americana Muhlenberg.* Now,
this author furnished no description, but gave as the range of his species " Virg.
Car. Pens." It therefore comprised at least two species, — for the one which
grows in Pennsylvania does not extend even as far south as Virginia, the plant
well developed in upland portions of this state and of North Carolina (and rep-
resented in his herbarium) being P. grandifolia,—&nd the rules require a name to
be rejected if undescribed or if derived from two or more discordant elements.
In 1840, however, Rafinesque ^ described several Parnassias, and the features of
his P. glauca correspond best to those of the plant in question. The four eastern
species are, then:

SYNOPTICAL KEY TO THE EASTERN PARNASSIAS

Leaf-blades cuneate, truncate, or subcordate at base; petals sessile.

Flowers 3 to 4 cm. in diameter; petals few-veined; staminodium-tips globular; ovary
conical, deep green.
Staminodia shorter than stamens; soil circumneutral ; range, N. J. and Pa. to la. and

northw. (unknown s. of lat. 40°) P. glauca.

Staminodia longer than stamens; soil circunmeutral ; range, n. Fla. to La., n. to Mo.

and to Va. mts P. grandijolm.

Flowers 4 to 5 cm. in diameter; petals many-veined; staminodium-tips lanceolate; ovary

ellipsoidal, white; soil mediacid; range, Fla. to N. C P. caroliniana.

Leaf-blades cordate; flowers about 4 cm. in diameter; petals clawed; staminodia somewhat
shorter than stamens, their tips ovoid-conical ; ovary conical, greenish white ; soil mediacid ;
range, Ga. piedm. to Tenn. & Va. mts. & coast P. asarifolia.

SEDUM

Field studies of the eastern Sedums have led to certain changes in current
views as to their classification and distribution, which have been summarized in
a recent article in a horticultural journal.® The eastern species fall into four of
the sections recognized by Praeger ^ and by Berger,* which are as follows:

KEY TO SECTIONS INCLUDING EASTERN SEDUMS
Plants perennial from stout fleshy caudices ; follicles erect, at most their beaks spreading.

Flowers chiefly perfect ; follicle-beaks slender § Telephium.

Flowers in part dioecious ; follicle-beaks stout S Rhodiola.

Plants perennial from slender stolons, or annual; follicles spreading.

Duration perennial ; leaves relatively broad; petals white § Seda genuina.

Duration annual ; leaves relatively narrow; petals pinkish S Epeteium.

1 N. Amer. Flora 22: 80, 1905.

2 Manual SE. Flora: 590, 1933.
8 Addisonian: 43, 1934.
*Catalogus: 33, 1813.

» Autikon Botanikon : 42, 1840.

•Gard. Chron. Amer. 38: 264, 1934.

» J. Royal Hort. Soc. 46: 1, 1921.

•In Engler & Prantl Nat. Pflanzenfam. ed. 2, 18* : 438, 1930.

OUR EASTERN PARNASSIAS AND SEDUMS

19

<*<^

I

i

J

Section Telephium. One species, S. telephioides Michaux.
Praeger and Berger consider this a mere variety of the Eurasian S. telephium,
but the plants differ in so many respects that I prefer to follow current American
usage and keep them distinct. They may be separated thus:

KEY TO TWO MEMBERS OF SEDUM, § TELEPHIUM
Herbage glaucous; leaves mostly opposite or subopposite, short-petioled, few-toothed; petals

white to pink; follicles pink, tapering into a spreading beak 1 to 1.5 mm. long; range, Ua.

piedm. to 8. Pa. mts S telephiotdea

Herbage green; leaves mostly alternate, scarcely petioled, often many-toothed; petals dull

purple; foUicles bronzy, tipped by an abrupt beak barely 1 mm. long; range, escaped

throughout ne. U. S., but rarely blooming s. of lat. 42° S. tekphium.

Sedum telephioides has been reported from several stations in western and
central New York, but as shown by House ^ most of these refer to S. roseum. In
the U. S. National Herbarium there is a specimen which seems to represent
S. telephioides from Clove, Dutchess County, but it may be an escape.

Section Rhodiola. One species, S. roseum (L.) Scop.
The southernmost station for this circumboreal species is usually regarded as
one in northern Bucks County, Pennsylvania, the plant of the isolated occurrence
on Roan Mountain, North Carolina— Tennessee, 500 miles away, being classed
as S. roanense Britton. While individual plants of the two may have a dissimilar
aspect, specimens can be found in either colony which match the characters of
those in the other. The real situation in respect to the supposed differentiatmg
features, leaf -margin, petal-shape, and petal-color, is, then:

Leaves toothed in most plants but entire in an occasional one; petals when carefuUy dried
oblonXceolate, but sometimes curling into linear or lanceolate shapes; groundcolor of
petals green, with bronze coloration extending from the tip Vs or sometimes ?4 the W
down ; range, s. to Fa • • u «««^

Leaves toothed in many plants, but entire in an occasional one. such as that which chanced
to be^lected asX^ Petals when carefully dried oblong-lanceolate but Bometinies

cSrlinglnto lanceolate or liAear shapes; ground color of petals green, with .bronre coloi^tion
expending from the tip V3 or sometimes only H the way down; range, isolat^ on Roan

Mt

Should anyone consider these differences of specific rank, they may use the

second name. If, however, only varietal distinctness is favored, the second plant

may bear the name S. roseum var. roanense (Britton) Berger. Perhaps they are

not separable at all.

Section Sed* genuina. Two species.

Sedum tematum Michx. is the one eastern species concerning which no tax-
onomic difficulties have arisen, its broad obovate leaves consistently whorled in
threes being highly distinctive. There is merely some question as to the northern
limit of its range as a native plant, the stations north of middle Pennsylvania
being conceivably escapes from cultivation.

1 Annotated List Pita. N. Y.: 376, 1924.

r

I

20

BARTONIA

i

The second eastern :member of this section, S. ncvii Grtty, ha«, on the other
hand, been repeatedly confused with S. pulchellum. Thus, reports of the latter
from Harpers. Ferry, [West] Virginia, are based on the occurrence there of the
former; and reports of S. nevii horn Missouri refer, as shown by, Palmer and
Steyermark,^ to juvenile S. pulchellum. The best evidence at present available
indicates the real range of &. nevii to be from- central Georgia to Alabama, south-
ernmost Illinois, easternmost West- Virginia, and central Virginia.

Section Epeteium. Two species.

The name Sedum pulchellum was assigned by Michaux ^ to a species he col-
lected in Tennessee; he gave the locality as KnoxviUe, but this was presumably
a slip for Nashville, as it is abundant around the latter place,— which his journal
«hows he visited— but unknown much farther east. This plant is a winter annual,
but Pursh » took it to be a perennial, and horticulturalists use- " pulchellum " for
a perennial species of unknown origin— not, as they suppose, " native to the
southern states." Michaux's plant, the only one to which his name properly
belongs, ranges from northwestern Georgia to Arkansas, southeastern Kansas,
and central Kentucky. It does not grow in Virginm.

Material from Georgia distributed by Curtiss has recently been n&med Sedum
vigilimoniis by SmaU* but I am unable to find any real differentiating char-
acters. When several collections are compared, one obtains these data:

T^nvpq linear-subterete when dried under considerable pressure flat-linear; cyme crowded on
pilots growing^r^^^ less so when in the shade or in moist soil; flower-size varymg with
Crtion sepals from broadly to narrowly lanceolate, and from 1.5 to 3-mmm length;
pet"ls as muc'h L outline and 3 to 6 mm. in length; and follicles 4 to 8 mm. -th b^eaks^O^^^
to 1 mm. in length ; \ A u

Tpflvps linear-subterete when dried under light pressure remammg so; cyme rather open when

growing TJm^^^^ more crowded in the sun or in dry places; habitat of type station

unfrvofable resulL^ in sepals being rather narrowly lanceolate and little exceeding 2 mm.

fSh-t petals b'eing aL rather narrow and Httje exceedmg 4 mm.; and m the foH^^^^^

being around 5 mm. long, with correspondingly short beaks ^. vigiUmontis.

In other words, plants exhibiting the features of S. vigUimontis can be found
in the midst of colonies of S. pulchellum wherever the environmental conditions
are unfavorable, so the former is but an ecological form.

Sedum pusillum Michaux needs little discussion. It has been placed m a
separate genus, Tetrorum, by Rose,^ but the slight differences in outline of flower
parts and the variation in their number from 4 to 5 are scarcely of genenc value.
There seems to be general agreement as to its range,— granite ledges from Georgia
to North Carolina, although it is rather rare.

1 Ann. Missouri Bot. Gard. 22 : 553, 1935.

2 Flora Bor.-Amer. 1 : 277, 1803.
8 Flora Amcr. Sept. 1 : 324, 1814.

• * Manual SE. Flora: 587, 1933.
6 N. Amcr. Flora 22, pt. 1 : 59, 1905.

Reprinted from Proceedings of the Pennsylvania Academy or Science

pages 12-15. Vol. X, 1936 ^ci^nce,

PRESIDENTIAL ADDRESS

REFLECTIONS ON THE ORIGIN OF LIFE

By Edgar T. Wherry

University of Pennsylvania

During the first ),alf of its span of existence, the Earth was in an azoic
or ifeless state. Then, about a billion years ago, life appeared upon it
and the living matter proceeded to evolve into a vast series of organisms
which have occupied nearly the whole of its surface. The purpose of this
address is to consider some of the details of this remarkable phenomenon
It IS difficult to formulate criteria which will distinguish livin- from
non-living matter, for exceptions always have to be admitted. AbFlity to
reproduce itself is an attribute of living matter in general; although a
gas flame is not to be classed as living merely because it can light an in-
finite number of identical flames, nor a mule as non-living merely because
this power of reproduction is lost. Responsiveness to external stimuli is
another characteristic of life; yet our non-living flame is able to move
away when we blow upon it. and our living mule may at times exhibit a
provoking failure to respond to stimuli. However, living matter is cer-
tainly characterized by its variability; for although any one „on-livin<-
molecule of a given composition is identical with innumerable others no
two living organisms seem ever to be exactly alike

The possibility that life as we know it did not originate on this Earth
but was started by a spore which came in from some other part of the solar
system is a well-known subject for speculation. However, as with those
plays which were not written by Shakespeare but by another man of the
same name, if the place of origin of life was not on our Earth, then it was
on another planet of the same makeup, and the discussion here given will
apply equally well to this other Earth.

Protoplasm, the fundamental sub.stance underlying living matter con
fains relatively large amounts of proteins, which in turn are built up of
ammo acids. These are organic molecules with an acidic radicle on one
side and a basic radicle on another, and accordingly produce, when dis-
solved m water, both positive and negative ions, in unlike amounts The
strains set up in the structure by this double and unequal ionization may
well be one .source of the responsiveness to external stimuli exhibited bv
aggregates of such molecules. The "peptide linkage" by which amino
acids are held together into proteins is perhaps another place in the struc
ture at which respon.se to external influences may occur. More important
than either of these, however, is the .spiral strain .shown by these "build-
ing-stones of the protein mass. For many of the amino acid molecules

PENNSYLVANIA ACADEMY OF SCIENCE

13

\'

I

contain one or more ''asymmetric" carbon atoms, some twisted to the
right, others to the left. And when a series of these becomes attached, an
intricate system of balancing of the strains develops, resulting in extreme
sensitivity of the system to stimuli. This subject of spirally strained
molecules requires, then, further consideration.

Pasteur is deservedly famous as the first scientist clearly to recognize
and incontestably demonstrate the role of bacteria in the causation of
disease ; but he also made discoveries in the inorganic realm, at least one
of which was of fundamental importance. Quoting from Mrs. Devon-
shire's translation of Vallery-Radot's Life of Pasteur:

Pasteur noticed that the crystals of tartaric acid and the tartrates had little faces,
which had escaped even the profound observation of Mitscherlich and La Provostaye.
These faces, which only existed on one half of the edges or similar angles, constituted
what is called a hemihedral form. When the crystal was placed before a glass [mir-
ror] the image that appeared could not be superposed to [upon] the crystal. . . .
Pasteur thought that this aspect of the crystal might be an index of what existed
within the molecules, dissymetry of form corresponding with molecular dissymetry.
. . . Therefore, reasoned Pasteur, the deviation to the right of the plane of polariza-
tion produced by tartrates and the optical neutrality of paratartrates would be ex-
plained by a structural law. The first part of these conclusions was confirmed; all
the crystals of tartrate proved to be hemihedral. But when Pasteur came to examine
the crystals of paratartrate, hoping to find none of them hemihedral, he experienced
a keen disappointment. The paratartrate also was hemihedral, but the faces of some
of the crystals were inclined to the right, and those of others to the left. It then
occurred to Pasteur to take up these crystals one by one and sort them carefully, put-
ting on one side those which turned to the left, and on the other those which turned
to the right. lie thought that by observing their respective solutions in the polarizing
apparatus, the two contrary hemihedral forms would give two contrary deviations. . . .

With anxious and beating heart he proceeded to this experiment with the polariz-
ing apparatus and exclaimed "I have it! " His excitement was such that he could
not look at the apparatus again . . . never was there greater or more exuberant joy
on a young man 's lips. lie foresaw all the consequences of his discovery. The hith-
erto incomprehensible constitution of paratartaric or racemic acid was explained; he
differentiated it into righthand . . . natural tartaric acid of grapes, and lefthand tar-
taric acid. These two distinct acids possess equal and opposite rotatory powers which
neutralize each other when ... in aqueous solution ... in equal quantities.

That carbon atoms whose bonds are attached to four different atoms
or groups are under a spiral strain is shown by the fact that they will
tw^ist the plane of a transmitted beam of polarized light. However, just
as our two hands are mirror-images of one another, but not superimposa-
ble, so every spiral has both a riglit-handed and a left-handed form.

1

Numbering the attached radicals consecutivelv, the molecule 4C2 is not

3
1
identical with 2C4; the two are related as an object to its mirror image,

14

PEXXSYLVAXIA ACADEMY OF SCIENCE

but cannot be superimposed. A solution or crystal of the onp Cm . • .

that 'i:e:r;;:;"ie rreSinriu"^ ''- ^^^''=""^"°"

selves .sliow re.s,,ectivelv rZt I7.ff^ f w substances may them-
by crystallizing and sorting tbe crystals into two piles

-.. one ma!:- bei-rt": thiTcZ nT. ib/^r "sr L^"' "

^^=:! r::: Lt^^faTt:: r.r r^' -'^; ^^^" --^^^

tion of the left-bar/Sle t^'^ort^rorrrnTS ''' '°™^-

s In , ""'''" """^ '''ft-'"'nded molecnle^ are always formed h

solntion being correspondingly optically inert. This result is due to tl.
operation of a law of f'liflnnp +i.« i , • -""^'^ ^^^^ii is due to the

will attach its" If t L "l t-Tr a t te'e'fTha':;? '^V '"* '' ="^" '•^^'^^'
carbon atom. If, „ow t reauires i Jf ""f""" "" '" ««y»metric

und!ubtedlv T 'P'"-'"'' °f """S "tatter, carbon compounds were
undoubtedij forming .spontaneously at various points on the earth's sur

luced "!"■"; "'''"""''"' «•""•" --'^^ -« - doubt oc sionalVp 0-'
ie ,v I r '■^■^"'V'^^"^ '•''-'"*- «-' "- '"ft-handed moditic^th s

other direction,re.:d;;:rtl"Xt%7r:^^^^^^^
r t,,e 0 1 ei form of a synthesized compound must be sought

n...i en^iTrut; :;i""^t""^' ""-- ""'^•" •-'" *■- -o-^ or

the surface 0 L en 7 mT' ''T"''"'"" "' ''''^''' ^^'^"^J «"-■«
in a glass vesse L cr !.t 1 .. T ""™''= '^'"""''♦^^ ^^'n «''"tion

on «:-.ean :::t of'^^ltit tXt^^t^^^^^ »'"' ^^•■'- P'^-''

in definitelv oriented positions r'.''"? "'*'"''*' "^''^'^' '^'^'^''^^
liauidcrvst«K»r 1 P""*'""'- Likewi.se, when substances vieklin-

dS b •:;::: zi^tz::'-"'-'''''' "''''''-' "--«■ '^ p-

PENNSYLVANIA ACADEMY OF SCIENCE

15

Quartz, the most abundant mineral on the earth's surface, has its sili-
con atoms so united with oxygen atoms that spiral strain is developed.
As required by the law of chance, there are in the world, to be sure, just
as many right-handed as left-handed quartz crystals; but these individ-
ually may grow to large size.

The unsatisfied attractional forces extending up in the unique or
c-axial direction from a right-handed quartz crystal must have a right-
handed twist, and those from a left-handed quartz crystal a corresponding
twist in the opposite direction. If, then, spontaneous synthesis of asym-
metric amino acids took place on the surface of a single quartz crystal
which chanced to be, say, right-handed and to have broken across the
unique axial direction, the molecules which formed may well have been all
right-handed ones.

If conditions favored the successive production of different molecules
on the same right-handed, twist-invoking quartz surface, and these were
stimulated to unite, protein aggregates capable of exhibiting the attri-
butes of living matter could ultimately have formed. The adsorptive
properties of the clay colloids produced by tlie weathering of rocks may
well have been the stimulus leading to such aggregation, and the richness
of these clay colloids in potassium rather than sodium corresponds to the
fact that plant life utilizes potassium in its physiologic processes. Ani-
mal life, being parasitic on plants, must have developed later, and the fact
that it utilized sodium indicates its origin in the sea, where this element
accumulates. However, for the reasons herein outlined, I would locate
the site of origin of the earliest life in a mud-puddle on the surface of a
broken quartz crystal.

Reprinted t'loiu Amekk ax Fkkx .JorKXAL, Vol. 26, No. 3,
l);i}4es 77-8«). Jiily-Septenibcr, 193fi.

I

Variants of Some Appalachian Aspleniums'

Edcah T. Wiii:k'i;v and William J). (Ihav

Teji years ajio it was pointed out by llie senior autlior-
that tlie "A])])alaeliian As])]eniinns" i'oriii a series sliow-
iiig intermediates between certain loii«>-r('eo«»nized sj^e-
cies. Two such intermediates were assifrned names,
.1. stotlcri and ^1. fruddli. In recordin«r the latter in
New Jersey, Kobbe and Davis' classed it as a form of
-t. piniKitifidum, but Kestner^ found its spores to be
imperfect and non-viabl(\ favorinj,^ a liybrid interpre-
tation. It has beeu rep-arded, on morpholojrical p-rounds,
as of hybrid orijrin, hy (iraves.' I]h)m(|nis1" ami Small."

A question was raised l)y CJraves^ as to the distitictness
of Asj/hiii>nif stothri fiom .1. t/raiu sii, \n\\ after e.xamin-
in«r photographs of the foi-mei- he acce|)ted their inde-
pendence. Kestnei-'' raised fiom spoi-es plants of .1.
sfoflrri Jiiatchinp- in i^xcvy i-espect the features of the
oi-ipinai colnnw

In the ])i-esent ai-tich' a scries of ilrawinps of tiiese
plants and some as ycl unreported intermediates and
variants arc presented. These were made by placin«< the
fronds in an <'nlar<iintr camei-a. ti-acin«r their outlines on
pajHM-. and reiliicinL! the drawin«rs apain in makinjr the
ruts here rcprodnccd. All arc natural size, except where
otherwise stated.

1 Contribution from the Botanical J^aboratory and Morris
Arboretum of tlie University of Pennsylvania.

-•Tliis Journal 15: 47. 11)2.1. - Ibid. IH: 22. ]928. "* Ibid.
•22: S4. 1932. ''Ibid. P3: 48. 1926. « Ferns of N. C: 91.
1934. -Ferns of X. Y. : ". ]!>35. « This Journal 1(5: 49.
192«;. V Ibid. 22: 83. 1932.

< <

1

Kf'piinted from Amkkk ax Vers Journal, Vol. 26, No. 3,
pages 77-80. .July-Scptonibor, 1930.

Variants of Some Appalachian Aspleniums'

ErXJAIi T. AViIKHKY AM) WlLLIA^f 1). (JuAY

Ten years ajio it ^\as i)()in1e(l out by llie senior author-
that the "Appalachian Aspleuiunis" t'oi-ni a series shtnv-
iiig intermediates between certain lon«»- recognized spe-
cies. Two such intermediates were assifjned nam<>s,
-1. static ri and A. trudrlli. In record injz" tlie latter in
New Jersey, Kobbe and Davis" classed it as a foi-m of
.1. piniKittfidnm, but Kestner"* found its spores to be
imperfect and non-viable, favorinji' a hybrid intei-pre-
tation. It has been rej?arded, on morpholog-ical ji-ronnds,
as of hybrid orij^in. by Graves,' ]jlom(|uist'' and Small."

A (piestion was raised by Cii-aves^ as to the distinctness
of Asplv Ilium stotlcri from .1. ffravrsii, but after examin-
injr photo«rraphs of the former he acce])ted their inde-
pendence. ICestner" raised from spor<'s plants of A.
sfotlfri matcliinji" in evei-y resjx'ct the featui-es of the
nrijiinal colon}'.

In the present article a sei-ies of drawiiij^^s of these
l)lants and some as yet unreported intermediates and
variants are presented. These were made by placinjjr the
fronds in an enlarpring eamcia. tracin<r their outlines on
|)aj)ei-. and reducing' the di-awinj^s a«2ain in makin<r the
cuts hei'e repi'oduced. All ar-e natural size, except where
otherwise stated.

1 Contribution from the Botanic.'d J^aboratorj and Morris
.Xrboretiini of tlie University of Pennsylvania.

•-' This Journal 1.1: 47. 192.J. ^ Ibid. IH: 22. 1928. ^ Ihid.
22: 84. 1932. -> Ibhl. 10: 48. 1920. e Ferns of N. C: 91.
1934. "Ferns of N. Y. : ."7. 193.";. s This Journal 10: 49.
1920. oma. 22: 83. J 932.

i I

INTENTIONAL SECOND EXPOSURE

IRRFriTIT AP PAnTMAT-T/-kM

1.

3.

78 American Fern Journal

The pinnatifidum — trudelli — montanum series.
A common phase of Asplenium pinnatifidum Xut-
tall, from the McCall's Ferry region, Lancaster
Connty, Pa. The cutting is pinnatifid throughout,
even the lowest pair of lobes being adnate to the
rachis; lobes obtuse and marginal teeth obscure.
A frequent variant, here represented by a frond
from Giles Co., Va. Lower lobes set off on short
stalks, and margins more distinctly toothed. This
may be termed A. pinnatifidum, phase approaching
A. trudelli, or, on the hybrid interpretation, A.
pinnatifidum > Y^ montanum.

Typical A. trudelli Wherry from its type locality.
Cully, Lancaster Co., Pa. Three pairs of lobes have
now become pinnae, supported on their individual
stalks, and tliese pinnae are somewhat pinnatifid.
This lies so nearly intermediate between the two
end-members of the series that an independent
name is useful for it, although there seems no doubt
that it represents A. pinnatifidum X montanum.
Intermediates between A. trudelli and A. montanum
are apparently rare, but after some years of search
a colony of such a plant was located near Ohiopyle,
Fayette Co., Pa., from which the frond sketched
was obtained. In this the pinna-stalks are so elon-
gated and the toothing so prominent that it might
be classed as A. montanum, phase approaching A.
trudelli, or, as a hybrid, A. pinnatifidum X <
mo7itanum.

A typical pliase of A. moyitanuin from Buckingham
Co., Va., bipinnate at the base with the pinnules
pinnatifid.

In these, as in most of the other drawings, the en-
larged pinna showing the venation represents a
member of the second pinna-pair.

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American Fern Journal

The pinnatifidum — gravesii — bradleyi series.

6. Aspenium pinnatifidum from the type locality along
the Schuylkill River in Philadelphia, Pa. This
shows the lobes better developed than No. 1, and in
addition the tendency toward areolate veining occa-
sional in this species.

7. This specimen, from York Furnace, York Co., Pa.,
is similar to No. 2, but shows dark coloration
throughout the stipe and dark brown sori. It is
therefore regarded as A. pinnatifidum, approaching
A. gravesii, or .1. pinnatifidum > X hradleyi.

8. A. gravesii Maxon from McCall's Ferry, Lancaster
Co., Pa., % natural size. In outline this resembles
some phases of A. trudelli, but again the dark stipe
and sori indicate a relationship to A. hradleyi
instead of montanum. Being about intermediate
betw^een the tw^o end-members of the present series,
it may be designated A. pinnatifidum X hradleyi.

9. Search for a member of the series lying nearer to
A. hradleyi was long unsuccessful, but one was
finally found in a colony discovered many years ago
by Dr. Roland M. Harper along Flint River in
northwestern Meriwether Co., Ga. This shows a
peculiar feature, upward-sloping pinnae — also pres-
ent on a plant in the U. S. National Herbarium from
Hartsville, Darlington Co., S. C.-— suggesting a rela-
tionship to the subtropical A, deiitatum (cf. fig. 20.)
As the latter seems improbable, however, it may be
termed simply ^1. hradleyi, phase approaching A,
gravesii, or A. pinnatifidum X < hradleyi.

10. A small frond of typical A. hradleyi Eaton, from
Gaston Co., N. C. The dark coloration extends up
the stipe to the fourth pinna-pair, and the margins
are much toothed and cut.

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Am ERIC AX Fern Journal

11

Presumed pinx atu idu m — plat YxVeurox crosses
Asplenium pinnafifidum, a phase with acuminate
kjbes instead of tlie obtusish ones of figs. 1 and 6;
from York County, Pa., 73 natural size. In this ease
the three presumed liybrids, wliich follow, do not
form a series from one end-member to the other.
Apparently, as in tlie analogous A. ehenoidcs Seott.
The parent sp.'cM's are too distantly related; the
liybrids combine parental characters in an erratic
way, and develop some details of lobing and cutting
not shown by either parent.

12 Tliis curious plant, here shown about half natural
size, was found by Sadie F. Price near Bowling
Green, Kentucky, many years ago, and distributed
as A. pinnatifidioit. It may well represent, liowever.
a hybrid with A. plat ^1 neuron, similar elongate lower
divisions being known in A. ehrnoides.

13. A. stotleri Wherry, from the type station in Jeffer-
son Co., AV. Va. Although the rounded lobes are
anomalous, this is still interpreted, as it was origin-
ally, as having developed by hybridizatiou of th.'
two species under consideration.
This is a frond on a plant recently collected by Mr.
Thomas X. McCoy in Boyd Co., Ky. It appears to
represent another result of the same crossing, being
intermediate in texture and sorus-color between the
two presumed parents. All three of these may be
referred to as A. pinnatifidnm y( platffncuron : or.
if a single name is desired, that first proposed for
a hybrid between these two species, A. stotleri. may
be extended to cover the others, even though their
resemblance is but slight. ^^

A typical fertile frond of .1. platyneuron (L.) Oakes.
y^ natural sizr.

10 This plant is (U'snilH-d .'i« n now spocics on page 104.

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84

American Fern Journal

Additional Aspleniums.

16. A narrow, elongate, but normally cut phase of A
pinnatifidum from York Furnace, York Co., Pa., -/s

natural size. .

17. Same locality but somewhat greater '■educt'on in
size than No. 16. Further development of the ten-
dency to narrowing and elongation has here resulted
in a frond without well-marked lobes but with
jagged margins, (cf. Camptosorus rkizophyllus
forma boycei Wilson, This Journ. 25: 18.) Such
fronds are known to attain at times a length of 30
em., the maximum for A. pinnatifidum. The succes-
sive figures 1, 6, 11, 16 and 17 bring out all the
frequent phases of this species ; although still others,
as well as various freaks, appear occasionally.

18 A giant hybrid from Muddy Creek, York Co. 1 a
V, natural size. The elongate upper part of the trond
with its simple lobing points unmistakably to A.
pinnatifid,m as one parent. The cutting of the
lower pinnae suggests A hradkyi to have been the
other parent, and this is also indicated by a rather
dark coloration of its sori. It is therefore perhaps
a phase of A. gravesii with unusually long, narrow
pinnae, although the broad base and deep cutting
are so reminiscent of A. moidanum as to lead to a
question as to whether a cross of A. gravm, with
montanum may not be represented.
19 In the York Furnace region A. monlanum some-
' times develops slender phases, a typical example of
which is here reproduced, slightly below natural

size. ^ „

90 The final dia-ram represents A. dcntatum L. trom
Dade County, Florida, natural size. It is put ni for
comparison with No. 9, the upper pinnae of which
show such a curious resemblance to those of this
presumably unrelated species.

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American Ferx Journal

The recommendation is frequently made that it' two
plants are connected by a complete series of intergrada-
tions, they should be considered as only subspecifically
or va'rietally distinct. To follow such a plan here, how-
ever, would result in confusion, for nearly all the plants
figured would then have to be classed as belonging to the
earliest recognized member of the group, A. platyneuron
(L.) Oakes, or perhaps to A. montanum Willdenow.
This would require a system of nomenclature so complex
that no one would ever use it.

In partial series such as shown in plates 1 and 2, how-
evei-, names for some of the intermediates may serve a
useful purpose. AVhile the few specimens included in
herbaria in 1875 could be adequately named as either
.1. montanum, A. pinnatifidum, or A. hradleyi, so many
ad<litional collections were made during the following
50 vears that more names were needed. Two new ones
accordingly came into use, making the two series
montanHm—tnidelli — pinnatifidum, and pinnatifidum—
gravesii—hradlfyi respectively. Possibly in another like
period names will be found desirable for the plants here
shown to bridge the gaps between these five. For the
present, however, we prefer not to propose four new
names, but to designate the intermediates by expressions
bringing out their positions in the series. Tn the prepar-
ation of dichotomous keys to the plants covered in a
manual or local flora, the existence of such intermediates,
as well as of cases of unusual cutting like those shown
in figs. 17 to 19, should certainly be allowed for, other-
wise amateurs may be misled, and perhaps overlock note-
worthy hybrids or variants.

PniLADELl'TIIA, Pa.

^14

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Eeprinted from Science, August 14, 1936, Vol. 84,
No. 2172, pages 160-161.

%

SOME RESULTS OF INVESTIGATIONS ON
POLYPORUS SCHWEINITZII FR.

In July of 1928, while forest pathologist in the
New York State Conservation Department, the senior
writer's attention was called to a resinosis disturbance
which was present in the root crown and in the roots
of northern white pine {Pinus strobiis L.) in forest
plantings at Norwich, N. Y. A year later, there wae
reported to him a similar condition in the municipal
forest plantings of the city of Rochester, N. Y., located
along H«mlock and Canadice Lakes. Preliminary
surveys of these situations showed that, although the
disease was making very rapid progress, the cause was
quite obscure. The plantings along Hemlock Lake,
where the disturbance was most severe, were estab-
lished from 1910 to 1914 on abandoned fields. All the
stock had been grown from seed in the nurseries of
the New York State Conservation Department. Im-
ported white-pine seedlings had been grown a few
years previously in some of these nurseries which
have since been abandoned. This fact suggested the
possibility that a parasitic fungus of foreign origin
might have been introduced into the nursery soil and
spread to various places in the state on stock grown
from seed in these nurseries.

Intensive investigations of the cause of this disturb-
ance were begun in the summer of 1930 by the senior
writer and have been conducted largely in the region
of Hemlock and Canadice Lakes. Reddish streaks
suggestive of incipient decay were observed occa-
sionally in the central part of some of the "resinosed'*
roots; many cultures from these discolored regions
failed to yield any organism known to cause the decay
of wood ; bacteria were the only organisms present in
the majority of the cultures.

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In August, 1932, a single sporophore which re-
sembled Polyporus Schweinitzii Fr. was found, but it
was 60 imperfectly developed that a species determina-
tion was impossible. It was attached to the base of a
living tree which showed little evidence of resinosis.
In the summer of 1933, sporophoree of P. Schweinitzii
were occasionally observed near the base of dead and
diseased white-pine trees in an eleven-acre tract along
Hemlock Lake. Sporophoree of this fungus were quite
abundant in this and two other plantings in the sum-
mer of 1935. The earliest date on which a perfect
sporophore of P. Schweinitzii has been observed in this
locality was June 6, 1935. With this discovery of
P. Schweinitzii, the senior author then sought to deter-
mine whether the resinosis was caused by this organism
or if it was due to some other pathogene. Hundreds
of cultures from the "resinoeed" lesions failed to yield
P. Schweinitzii, but a grayish-black fungus developed
in approximately 75 per cent, of these cultures; all
efforts to induce the latter to fruit have been unsuc-
cessful. Thus it would seem that no direct connection
exists between the resinosis disturbance and the injury
caused by P. Schweinitzii, since the trees attacked by
the latter may never show any, or very Uttle, evidence

of resinosis.

No sporophoree of P. Schweinitzii have thus far
been found in the plantings at Norwich, even though
hundreds of trees have died from resinosis. The roots
of trees which die from this disease may never show
any traces of the reddish streaks. Resinosis has been
found to be most severe where the pH of the soil is
6.0, or above, and the colloidal content 52 per cent, or
more. Even though P. Schweinitzii has been found to
be very abundant in such areas, it is also causing very
severe damage where the pH of the soil is around 5.5
and the colloidal content varies from 46 to 50 per cent,
or even less. The resinosis disease is of minor impor-

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tance, as contrasted with the damage which la being
caused by P. Schweinitzii,

Advanced stages of decay in the roots of living trees
of white pine due to P. Schweinitzii were first observed
in the summer of 1933. By the end of the summer of
1934, this root rot was found to be very wide-spread
in plantings totalling about 1,200 acres of white and
red pine in the Hemlock Lake region. Living trees
whose roots were badly rotted had beg^n to fall over
and have continued to fall in increasing numbers. At
the present rate of destruction, not a single tree in
this area will reach merchantable size. The losses
from the damage caused by P. Schweinitzii along Hem-
lock and Canadice Lakes are the most serious which
have thus far been reported in forest plantings in the
United States, if not in the world. P. Schweinitzii was
reported in 1925 as causing root rot in forest plantings
of Douglas fir near Biltmore, N. C.^

Chemical analyses of soil extracts show that in
plantings where P. Schweinitzii is present there is
nearly 21 per cent, more calcium in the upper four
inches of the soil than where the organism is not known
to occur. A number of chemical analyses of the ash
of the wood of infected trees show less calcium pres-
ent than in the wood of normal trees. It occurred to
the senior author that the activities of P. Schweinitzii
in the soil may render the calcium less available to the

trees.

Chemical analyses of a water extract of silica quartz
sand from around the roots of three-year-old seedlings
of eastern white pine in pot cultures, which were
inoculated with P. Schweinitzii and supplied with a
given nutrient by Wean's method,^ showed twenty-five
times more calcium where the fungus was present than
in the controls. These cultures and inoculations were

iG. G. Hedgcock, G. F. Gravatt and R. P. Marshall,
Phytopathology, 15: 568-569, 1925.

aBobert E. Wean, Sctencb, 82: 336, 1935.

IRREGULAR PAGINATION

<

Reprinted from ISIS N-^ 69 (Vol. XXV, i), May 1936.

made by Wean while a graduate student in forest
pathology at the University of Pennsylvania. He
found that P. Schweinitzii was highly parasitic on the
roots of one- and three-year-old seedlings of white
pine. The hyphae readily penetrated both the epi-
dermal and peridermal layers, especially in pot cul-
tures with nutrient solutions at a pH of 6.0 and 7.0
and under conditions of a reduced supply of phos-
phorus. Reddifih streaks were present in the central
part of infected roots which were even less than two
millimeters in diameter. These results are indicative
of the possibility that P. Schweinitzii may be distrib-
uted in nursery stock.

In view of the possibility that P. Schweinitzii as
found in the Hemlock Lake region might be of foreign
origin, Childs, while a graduate student in forest pa-
thology at the University of Pennsylvania, worked
with cultures of this organism from various parts of
the world, including many from the Hemlock Lake
area. At present, it appears that P. Schweinitzii,
which is causing root and butt rot in the forest plant-
ings in question, is purely of local origin. He has
found that within P. Schweinitzii there are a great
many individuals which, when grown under a g^ven set
of conditions, show a wide range of cultural reactions.
He has also found that a number of these individuals
are definitely homothallic and fruit readily in culture;
clamp connections have not been observed in any of
these cultures.

Harlan H. York
Robert E. Wean

Thomas W. Childs
University op Pennsylvania

^
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Animals impregnated by the wind

(0

When Camerarius first published the evidences of sexuality
in the flowering plants, he was faced with the necessity of proving
that wind could carry pollen from one flower to another. In
the course of his presentation he cited several classical examples
of the fertilizing power of air, and even quoted a passage in
Virgil's Georgics which tells how certain mares were impregnated
by the west wind. He thus called attention to a very ancient
fantasy. In his famous letter to Valentine, De Sexu Plantarum
Epistola, Tubingen, 1694, he referred to these mares and to the
** wind-eggs " of several species of birds :

" Those unfertile eggs of birds produced without the influence of the male
are generally called wind-eggs because, as Aristotle explains, in springtime
the birds seem to receive a fertilizing breath from the west wind. Virgil also
refers to this when he describes the excited condition of the mares :

" These words can be applied better to the conception of plants than to that
of animals, for the latter, although they may be moved and stimulated by the
wind, receive no germ from it, but plants owe much more to the wind because
in the spring, their organs of conception are directed to the Zephyr like so many
nostrils to inhale the rustling airs and the flower dust. Impregnated by the
wind's breath, (oh wonder to relate) they conceive without copulation. Also
the egg that is impregnated here in the plant through the medium of the wind
or the air, the carrier of the male principle is fertile and not a wind egg. Pliny
also commends the power of Favonius (the west wind)... That must now certainly
be interpreted with a grain of judgment, for apart from the old fable about the
wind-impregnated Spanish mares, how can such a fruit be other than a sterile
thing, a wind egg, a misbirth."

Strangely enough most classical and medieval philosophers
believed that certain mammals and birds could be impregnated
by wind, although they were ignorant of the wind's function
in the cross-pollination of the flowering plants. This belief in
the existence of what we might call anaemophilous animals is

(i) This investigation was aided by a University of Pennsylvania Faculty
Research Grant.

k\

96

CONWAY ZIRKLE

4

ANIMALS IMPREGNATED BY THE WIND

97

i

extremely ancient. From the earliest times it existed in Egypt,
Greece and the Orient and ultimately it became a part of the
intellectual heritage of many races. It spread along the sea routes
of the whole eastern hemisphere, and it has been found from
Portugal to Japan, and in all of the countries between. Our
first record of this belief comes from the Iliad, our last from
the folklore of the Ainu; between these extremes are records
by Greek philosophers, Roman naturalists and early Christian
saints. Medieval schoolmen copied the classical accounts while
Arab travellers, Chinese geographers and modern anthropologists
have reported numerous instances of the existence of this belief.
Indeed few scientific errors have had such an honorable historv.

Four types of females were the chief recipients of the winds'
attentions; they were mares, vultures, hens and women. This
makes it possible for us to arrange the stories into four classes,
each concerned with one of the four subjects. Surprisingly
enough, when the stories are thus grouped they do not seem
to be mere variants of some one primitive legend, but rather
four separate tales, each with its own origin but each influenced
by the others in the course of its development. The reason
why the wind was chosen as the fecundating agent in all four
cases is not at all clear. It will probably have to be sought in
some psychological twist of the not quite civilized folk among
whom the stories originated.

To the best of my knowledge the descriptions of anaemophily
in the animal kingdom have never been collected or made available
to modern students of parthenogenesis. Perhaps too much of
the history of science is concerned with the origin and development
of ideas which we now view as sound, yet any true picture of the
intellectual equipment of our predecessors must include, of course,
their errors as well as their occasional lapses into accuracy. Few-
subjects can illustrate the scientific standards of the credulous
ages as well as these stories of the wind-impregnation of animals.

Fortunately, the numerous passages which describe this
fecundation are not too long to quote, for many of them are
in works not easily obtained. As each illustrates some stage
in the development of the legend, or some practical use to which
the legend was put, they are given here in full. The passages
are arranged chronologically in four sections, each section dealing

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with one of the objects of the wind's attention. Needless to
say the following pages do not include all such records.

Mares

No one reading of the amours of Zephyrus and Boreas in
the Iliad would have cause to suspect that their philanderings
had a place in the history of science. In Book XVI (translated
by Andrew Lang) occurs the passage,

" And for him Automedon led beneath the yoke the swift horses, Xanthos
and Bahos, that fly as swift as the winds, the horses that the Harpy Podage bare
to the West Wind, as she grazed on the meadow by the stream Okeanos."

From Book XX (translated by Ernest Myers) —

" Three thousand mares had he that pastured along the marsh meadow, rejoicing
in their tender foals. Of them was Boreas enamoured as they grazed, and in
semblance of a dark-maned horse he covered them : then they having concieved
bare twelve fillies."

There is nothing in these passages about the fertilizing power
of the unpersonified wind but during the next five hundred
years the legend became full grown and we find it in the Historia
Animalium of Aristotle.

From Bk. VI; i8 p. 572a (translated by Thompson) —

" The mare is said also about this time to get wind-impregnated if not
impregnated by the stallion, and for this reason in Crete they never remove the
stallion from the mares; for when the mare gets into this condition she runs
away from all other horses. The mares under these circumstances fly invariably
either northwards or southwards, and never towards either east or west. When
this complaint is on them they allow no one to approach, until either they are
exhausted with fatigue or have reached the sea. Under either of these circum-
stances they discharge a certain substance called " hippomanes," the title given
to a growth on a new-born foal; this resembles the sow-virus, and is in great
request amongst women who deal in drugs and potions."

When Varro (118-30 B.C.) related the story he shifted the
scene of the impregnation from Crete to Spain. This may seem
to be a strange move to us, for in classical times the Cretans
were reputed to be the greatest of liars. In De re rustica (II; i),
however, Varro apparently did not need a Cretan alibi.

" Speaking of pregnancy, let me tell you some thing which happens in Spain;

J

IRREGULAR PAGINATION

9^ CONWAY ZIRKLE

no one will believe it, but it is none the less true. In Lusitania, near the ocean,
in that stretch of country where is the town Olisipo, certain mares on Mount
Tagrus conceive at a certain time of the year by means of the wind, just as hens
frequently do with us. the eggs of which we call ' wind eggs.' The foals, however,
born ot these mares do not live longer than three years."

By far the most famous description of these mares is in the
Georgics composed by Virgil in 31 B.C.
From Bk. Ill, 273 (tr. by Ogwan)—

" ^^j^/'^J^'^^'y "^^'^^^ «f ^". apparently, is the passion of the mares; even
reason did their mere passion lend them, when his Potnian team tore piecemeal
with their teeth the limbs of Glaucus. Love guides them over Gargarus. and
across the loud Ascanius; they climb the mountains and swim the torrent At
first when flame kindles in their longing vitals-in spring especially; for then the
heat revives along their bones.-facing the western breeze, all stand on the tall
cliffs, and breathe the gentle wind; and often, without copulation-strange to
tell-impregnated by the wind, they disperse among the rocks and headlands,
and deep valleys; not toward thy region. Eurus. or the sun-rise, but toward Boreas,
and Caurus. or where darkest Auster rises, and blots the heaven with a chilly
damp. Then, at this particular season, flows the hippomanes-so shepherds
truly call it-wliich jealous step-dames often gather to medicate their herbs and
deadly charms.

In the version by Columella (30-40 A.D.) the lascivious thought
of the mares aids the wind in the accomphshment of its purpose
From Bk. VI. Ch. 27. (Translated by M. C. Curtius)-

" Nor is there any doubt, but, in some countries, the mares are inflamed with
such a strong and ardent desire of coition, that, altho' they have not the male
yet. by their continual and excessive desire, raising in themselves the imagination
of venery. they (in the manner of fowls constantly kept in a barren) conceive
with the wind; which things the poet expresses with greater license...

Forasmuch as it is a thing also very well known, that on the Sacred Mountain
in Spain which extends itself toward the west, hard by the ocean, mares have
frequently been pregnant without coition, and have brought up their offspring :
which nevertheless is of no use, because it is snatched away by death when three
years old, before it comes to maturity.

•'Wherefore, as I have said, we must take care, that mares be not tormented
with their natural desires, about the time of the vernal Aequinox."

Shortly after Columella wrote his Res rustica, Pliny's famous
Natural history appeared (77 A.D.). The story of the mares
IS not one that Pliny would miss. The translation here quoted
was made by Holland and published in 1601. From Pt I
Bk. 8, p. 222-

" In Portugal, along the river Tagus, and about Lisbon, certaine it is. that

ANIMALS IMPREGNATED BY THE WIND

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when the West wind bloweth, the mares set up their tails, and turne them full
against it, and so conceive that genitall aire insteed of the naturall seed • in
such sort, as they become great withall. and quicken in their time, and bring
toorth foles as swift as the wind, but they live not above three years."

SiLius iTALicus (25-101 A.D.) was contemporary with Pliny.
In his poem, Punka, he referred twice to these wind-begotten
horses. He allowed them seven years of life.

From Bk. Ill, Hnes 378-383—

r !i ^"li ^^^^*""' ^^'*' ^^^ '"'"^^ ^'"'^^ ^^^ «P^" ^"^^^^^ of the land of the Vettones
Indeed here when the spring is mild and the breeze is temperate, the herd of horses
makes its race sure by observing secret copulations, and with the fruitful breeze
conceives a hidden love. But the span of life of the race is not great, and old

sTbles'^^' ^""^ '^''^" ^^^'' '' '^^ '''"^^'' '^""^ '*'^' *^^ ^^'"^ 'P^'^^^ '" ^hese

The following reference is to a race horse, and is reminiscent
of Homer. From Bk. XV, lines 362-366—

•; But on the inside of the left track he would just graze the turning post, distin-
guished by his great neck and also by many sportings of his mane over his neck-
his mother (Harpe) bore him in the presence of the fresh blasts of the West Winds
in the fields of the Vettones."

About the year 200 A.D., Aelian wrote De natura animalium.
He did not mention the legend of the mares of Spain but his
description of the begetting of sheep shows the influence of
the story. From Bk. II, Ch. 46—

"Indeed sheep are not ignorant of this, that the north wind and the south
wind accomplish their fertilization no less than the rams entering into them-
and moreover they know that the north wind is effective in the generation of
male offspring (2) and the south wind in the generation of female offspring -
wherefore, that sheep which is involved turns herself toward one or the other

^ *

(2) The belief that the direction of the wind at the time of copulation determined
o, '.';t "l!^"™^' "^'^^ widespread. It is recorded by Aristotle. HhtoHa Animalium,
.. ; ; '^' ^'^^''^' ^"'''"^ Naturalis, Bk. VIII. Ch. 72; Antigonus. Historia

Mtrabtltum, Ch. CXI; Columella. De Re Rustica, Bk. VII. Ch. 3; Albertus
Magnus. De Animal. Tract. III. Bk. 6, Ch. 2; etc. A Middle English translation
ot Palladius' De Re Rustica expresses the belief rather quaintly. Bk. VIII
Lines 95-99 — *

" And toward that wynd yf the tuppis ofre,
With litel malis fillith they the cofre,
And toward southwynd getith they femalis,
Yf hit be sooth, right notabul this tale is."

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CONWAY ZIRKLE

accordmg to wh.ch sex is desired... But sheep easily and without any ceremonies
have the wmds ass.stance in their conception, spontaneously with neither entreat^
nor boldness^ Even the shepherds are observers of this practice, for when the

hev iJ'l T' """ *' ""'"''"■ "^ f'™^'' °*P""« '' 'hereby increased,

they let loose the rams among the sheep."

Caius Julius Solinus, who lived toward the latter end of
the third century, repeated the story of the mares in his famous
work which was later known as the Polyhistor seu de mirabilibus
mundt. His version reads —

" In the neighborhood of Olissippo the mares indulge in sexual play and bear
■ssue w,th marvelous fertility, for when the west wind blows upon th J thTy

At about the time of Solinus or perhaps a little later, Lactantius
(264-340 A.D.) found a practical application for the story
Lactantius was a respected Christian writer who influenced greatly
the formation of the doctrine of the early Church although he
was never raised to the rank of a real authority. He attempted
to make the virgin birth of Christ more plausible to the Pagans
by reminding them of one of their own beliefs, i.e., the partheno-
genesis of certain foals. It is true that this was not the first
use that was made of such an argument, as many years earlier
Origen and Clement had cited the popular belief concerning
vultures and hens in their defense of their theological concep
tions. From Lactantius' account—

" Chap. XII. Of the birth 0/ Jesus from the Virgin ; of his life, death ami resurrec
turn, and testimonies of the prophets respecting these thirds

ViZ'TafH ""' T"'" ^'""' f °°''' <•'=-'"'''"« f™™ heaven, chose the holy

If The' Svine%""* t ■"'?" ™""'- ^"' '"'• *«'"« «"'" hy .he possession

of the D,v.ne Sp.r.t, conceived; and without any intercourse with a man her

vrgm womb was suddenly impregnated. But if it is known to all thaT ce'ttaL
an,mals are accustomed to conceive by the wind and the bree«, why should

Spint of God, to whom whatever He may wish is easy ? And this might have

z^nce'"" ^^' •"" ""' *' "^'"'"^ "^y "«- p«™->y " <::

Nearly a hundred years after Lactantius. Claudianus
(397 A.D.) referred to the wind impregnation of the tigress
This passage is evidently but a poetic variation of the same general

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ANIMALS IMPREGNATED BY THE WIND loi

theme (3). From the Rape of Proserpine, Bk. Ill, hne 265—

" Speedier than the West Wind that is her paramour, rushes the tigress, anger
blazing from her stripes, but just as she is about to engulf the terrified hunter
in her rapacious maw, she is checked by the mirrored image of her own form."

Justin, a contemporary of Claudianus, introduced the first
note of real skepticism into the story. He even went so far
as to term it a fable. From his History of the World, Bk. 44,
Ch. 3—

" Several Authors have affirmed that in Lusitania, near the Banks of the River
Tagus, the Mares conceive by the wind. What gave occasion to this fable, is
the great Fecundity of the Mares; and the vast Numbers of Horses that are' to
be seen in Gallicia and Lusitania, where the Jennets are so prodigiously Swift,
that 'tis not without some Reason they are said to be begotten by the Wind."

Justin's doubts did not extend to St. Augustine. Indeed
the Bishop of Hippo was anything but a skeptic. The following
extract from De Civitate Dei (413-426 A.D.) illustrates perfectly
the intellectual standards which made the Middle Ages possible.
Needless to say the books of St. Augustine influenced profoundly
the development of medieval Christianity. From Bk XXI
Ch. 5-

(<

Chap. 5.— That there are many things zvhich reason cannot account for, and
which are nevertheless true.

" Nevertheless, when we declare the miracles which God hath wrought, or
will yet work, and which we cannot bring under the very eyes of men, skeptics
keep demanding that we shall explain these marvels to reason. And because
we cannot do so, inasmuch as they are above human comprehension, they suppose
we are speaking falsely. These persons themselves therefore, ought to account
for all these marvels which we either can or do see. And if they perceive that
this is impossible for man to do, they should acknowledge that it cannot be
concluded that a thing has not been or shall not be because it cannot be reconciled
to reason, since there are things now in existence of which the same is true. I
will not, then, detail the multitude of marvels which are related in books, and
which refer not to things that happened once and passed away, but that are per-
manent, in certain places where, if any one has the desire and opportunity, he
may ascertain truth; but few only I recount. The following are some of the

(3) More than two centuries before Claudianus, Oppian denied this story.
From Cynegetica III : 355 (tr. by Mair) written in the last half of the second
century :— " Tiger, swifter is it than all wild beasts that are : for it runs with
the speed of its sire, the West wind himself, yet the West Wind is not its sire;
who would believe that wild beasts mated with an airy Bridegroom? For that
also is an empty tale, that all of this tribe is female and mates not with a male :..."

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CONWAY ZIRKLE

marvels men tell us :-The salt of Agrigentum in Sicily, when thrown into the
fire becomes fluid as if it were in water, but in the water it crackles as if it were
in the fire. The Garamantae have a fountain so cold by day that no one can
dnnk it, so hot by night no one can touch it.

"In Epirus. too, there is a fountain which, like all others, quenches lighted
torches but, unlike all others, lights quenched torches. There is a stone found
in Arcadia, and called asbestos, because once lit it cannot be put out The wood
of a certam kind of Egyptian fig-tree sinks in water, and does not float like other
wood; and, stranger still, when it has been sunk to the bottom for some time
it rises again to the surface, though nature requires that when soaked in water
It should be heavier than ever. Then there are the apples of Sodom, which
grow indeed to an appearance of ripeness, but. when you touch them with hand
or tooth, the peel cracks, and they crumble into dust and ashes. The Persian
stone pyrites burns the hand when it is tightly held in it, and so gets its name
from fire. In Persia, too. there is found another stone called selenite, because
Its interior brilliancy waxes and wanes with the moon. Then in Cappadocia
rtie mares are impregnated by the wind, and their foals live only three years
1 lion, an Indian island, has this advantage over all other lands, that no tree which
grows in It ever loses its foliage.

" These and numberless other marvels..."

Certainly any belief endorsed by such a galaxy of authorities
as Aristotle, Virgil, Pliny and St. Augustine would not lapse
during the Middle Ages. Shortly before the year 1200, Alexander
Neckam, whose mother was the wet-nurse of Richard Cceur-de-
LiON, wrote De naturis rerum. This book, which contains the
earliest European mention of the magnetic compass, also tells
of the mares.

"They say that the female horse whose opportunity with a male is lacking
conceives by the snorting blast of the North Wind, but its offspring will hve onlv
a few days. And so the hen when it takes its bath with frequent flutterings and
uith applications of dust, may lay soft eggs without coition, but does not obtain
the joy of offspring. The eggs, however, are called soft as if they were without
a skin, and whenever they are such that they may be supped up thev receive
this name. Those especially, however, are called soft which are found in the
DeiJy ot hens and are without an outer skin."

Walter of Metz included the story of the mares and the
wind in his U image du monde written in 1246. This was one
of the first books ever to be printed in English as it was translated
by William Caxton himself who issued an edition in 1480
From Caxton's translation —

" Ther IS a maner of Mares that conceyve of the wynde, and ben in a contre
that IS named Capadoce; but they endure not but iii yere."

Just twenty-two years after Walter of Metz wrote Uimage

animals impregnated by the wind

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du Monde, Roger Bacon (1268) completed his Opus Majus. Here
the story of the mares is related to a rather vague allusion to
the wind pollination of the dioecious date-palm.

" The human mind can be influenced to accept the truth of the virgin birth,
because certain animals remaining in a state of virginity conceive and bear young,
as, for example, vultures and apes, as Ambrose states in the Hexaemeron. More-
over, mares in many regions conceive by virtue of the winds alone, when they
desire the male, as Pliny states in the fifth book of the Natural History, and
SoLiNUS tells us in his book on the Wonders of the World. Aristotle maintains
in the second book on Vegetation that the fruits of the female palms mature from
an odor coming from the male trees."

Nicolaus Perottus (1430- 1480), Archbishop of Siponto,
included the story in his Cornucopia. From page 41 1 of the edition
of 1521 : —

... They say that in Lusitania by the river Tagus, the mares conceive foetuses by
the wind, facing it when Favonius is blowing, and offspring is born through
that (nicissimus), but it does not live more than three years. I believe that this
fable has arisen because of the fertility of the horses and the immense size of
the herd, for they are very abundant in Lusitania and are observed to be so fleet
that the wind itself seems not unworthy to share in tl eir conception."

Shortly after Perottus, Johannes Ravisius Textorus wrote
Epitheta, Parrhisiis, 15 19. He also repeated the story of the
mares. (From p. 141b, edition of 1524) —

" ... It is known that when the females assemble they are mostly seized with
such a great ardor that if they do not have a male they imagine a lover for
themselves and in too great a love desire, they conceive by the wind. For Varro
says that in further Spain in the spring time, the horses, incited by too much heat,
throw open their mouths against the cooler airs and winds to relieve their heat;
and thereafter they conceive and bring forth young, but these are not allowed
much time for their surpassing fleetness. For before they reach their third
year they are removed by death."

In 1570, Heresbach published his Res rustica. This was
translated in 1586 by Barnaby Googe as Four Books of Husbandrie.
The story of the mares occurs on page 1 1 7b :

" ..., neither is to be doubted, but that the mares in some Countries so burne
with luste, as though they have not the horse, with their oune fervent desyre they
conceave and bring forth after the manner of Byrdes, as the Poet noteth.

" In furious lust the Mare exceeds all other beasts that be. It hath been said
that in Spaine Mares have conceaved with the winde, and brought up their Colts,
but the Colts have not lived above three yeeres."

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CONWAY ZIRKLE

At last, in the seventeenth century, real doubt was felt as to
the ability of the wind to replace the male. Johannes Johnston
referred to the matter very briefly in Thaumatographia Naturalis,
Amsterdam, ,632. The English edition of this work appeared
in ,657 ^ An History of the Wonderful Things of Nature, translated
by A Person of Quality." From page 218—

" In Portugal they say, the mares conceived by the wind Varro Pi iw ,„h

fr- tr 'tht:: i^ti^- - ^ -- - ^^^^^::^.

Just three years after Johnston's Thaumatographia, the Historia
Naturae of Joannis E. Nieremberg appeared. Nieremberg seems
to have been imbued with the scientific spirit and he undertook
to 'nvest^ate the legend. From Bk. I, Ch. 66, p. 4,0a (edition

fe«iii'n,^l "° '"^''' '"'I °^ *'' *"« °f f"™" ""•«. th" « of unmarried

statLTn'^sfr."-^??* ^"'''' °'''°^''' '^95. W. D. Thompson
stated (p. 48) that the legend of the vulture's being impregnated

with Maut, the goddess of maternity. This would indicate that

(4) Gahca de Resende (.470-1536), a Portuguese poet.

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ANIMALS IMPREGNATED BY THE WIND

105

the legend is very old, perhaps older than the story of the mares
being fertilized by the west wind. The classical records are
not as ancient, however, and the earliest of these do not show
the myth in its fully developed form. Aristotle described the
production of vultures in De Mirabilibus but his account is notable
for its description of degeneration in the animal kingdom rather
than for its statements concerning vultures. He evidently felt
that the vulture was no ordinary bird but eagles and hawks seemed
to him just as exceptional. If the beliefs recorded by Aristotle
were held at all generally, it is obvious that the ground was well
prepared for the spread of the Egyptian legend. From 831a,
I, (tr. by Dowdall) —

" Of the offspring of a pair of eagles, so long as they pair together, every second
one is a sea eagle. Now from the sea eagle springs an osprey, and from these
spring black eagles and vultures : yet, on the other hand, these do not bring
the breed of vultures to a close, but produce the great vultures, and these are
barren. And a proof is this ; That no one has ever seen the nest of a great vulture."

If no one could find the nest of the vultures the question naturally
arose,— How could they breed ? Pliny (Bk. X, Ch. 6) recorded
this problem three hundred years later.

" The black Vultures are the best of that kind. No man ever could meet
their nest; whereupon some have thought (but untruly) that they fly unto us
out of another worid, even from the Antipodes, who are opposite unto us."

Plutarch (46?- 125? A.D.) referred directly to the Egyptian
tale in the Quaestiones Romanae. He seemed to be very skeptical,
yet he could not help revealing the fact that he really desired
to believe the story. In Question 93 he discussed auguries.
Now, if the victory or defeat of a Roman army depended upon
a correct prognostication by the augurs, it is obvious that reliable
birds were of the utmost importance. Plutarch stated the case
for the vultures. At this time the legend was fully grown and
in this form it remained for the next twelve hundred years.
From Bk. II, Ch. 46 (tr. by Chauncey)—

** Concerning Vultures.

" ... They say indeed that male vultures are never found, indeed, all are female;
for which reason these winged female creatures, not ignorant, and fearing scarcity
of offspring contrive to beget offspring in this way : they fly before the South
wmd : if perhaps the South wind is not blowing, they spread themselves toward
Ihe East wind by opening the mouth and after three years they give birth. They

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refuse to shelter any vultures in the nest; indeed I hear that Aegypios, which
are half way between vultures and eagles, and are male and black in color, are
discovered in their nests. I understand that vultures do not even hatch out
eggs, but produce offspring which are winged instantly at birth ; all of this I have
heard."

In 248 A.D., the story got into patristic literature as an aid
to the dogma of the virgin birth of Christ. No less an authority
than Origen used it in his refutation of the errors of Celsos.
From I : 37 —

"... But as a further answer to the Greeks, who do not believe in the birth
ot Jesus from a \ irgin, we have to say that the creator has shown, by the generation
of several kinds of animals, that what He has done in the instance of one animal,
He could do, if it pleased Him, in that of others, and also of man himself. For
It is ascertained that there is a certain female animal which has no intercourse
with the male (as writers on animals say is the case with vultures), and that this
animal, without sexual intercourse, preserves the succession of race. What
incredibility, therefore, is there in supposing that, if God wished to send a divine
teacher to the human race. He caused Him to be bom in some manner different
from the common ! Nay, according to the Greeks themselves, all men were
not born of a man and woman."

The great Christian historian, EusEBios (260-340 A.D.) described
the belief m the Praeparationes evangelicae written shortly before
318 A.D. From Bk. Ill, ch. 12—

" Finally they describe the third celebrity in the town in the devout veneration
of Diana whose effigy is like a female vulture in flight, to whom are adjusted wings
constructed from selected and choice gems. And they show that the winds
have power from Diana of creating and producing by this image of a vulture,
but vultures which they assume to be all female are thought to conceive from
the wind."

HoRAPOLLON NiLOUs, who lived in the fourth century, pretended
to mterpret the heiroglyphics on the Egyptian monuments. The
following is his rendition of the vulture character.

•• To denote a mother... they delineate a Vulture. They signify by it a mother,
because in this race of creatures there is no male. They are produced, however,
m this way. The vulture is kindled with a desire to conceive, opening her womb
to the North Wind, she is, as it were embraced by him for five days, during which
time she partakes neither of food nor drink, being intent upon procreation. There
are also other kinds of birds which conceive by the wind, but their eggs are of
use only for food and not for procreation, but the eggs of the vultures that are
impregnated by the wind possess a vital principle."

Ammianus Marcellinus, the historian, referred casually to

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ANIMALS IMPREGNATED BY THE WIND

107

the lack of male vultures, in Res gestae^ a work completed
probably in 378 A.D. From Bk. 17 —

" Meanwhile there may be an example of the knowledge of this fact in these
two things, they disclose the nature of birth through the vulture : for the reports
of natural science mention that no male can be found among these birds. And
they show the Queen bee among the species of bees which produce honey."

The story of the vultures got back into patristic literature
with St. Basil the Great who wrote The Hexaemeron, nine sermons
on the Creation as described in Genesis. The exact date of
this work is not known, but it was probably written sometime
between 330 and 379 A.D. From Homily VIII, Sec. 6 —

" Many birds have no need of union with males to conceive. But their eggs
are unfruitful, except those of vultures, who more often, it is said, being forth
without coupling and this although they have a very long life, which often reaches
its hundredth year. Note and retain, I pray you, this point in the history of
birds ; and if ever you see any one laugh at our mystery, as if it were impossible
and contrary to nature that a virgin should become a mother without losing the
purity of her virginity, bethink you that He who would save the faithful by the
foolishness of preaching, has given us beforehand in nature a thousand reasons
for believing in the marvellous."

St. Ambrose of Milan (334-397 A.D.) wrote his Hexaemeron
at about the time that St. Basil was writing. Like St. Basil
he mentioned the vultures only as evidence of the reasonableness
of the virgin birth of Christ. From V : 20 —

" We have spoken about the widowhood of birds and that virtue arose from
them first; now let us speak of chastity which also is proved definitely to dwell
in many birds, that can be perceived in vultures. Indeed vultures are denied
to indulge in coition, and conjugals by a certain practice and nuptial bonds engaged
in by chance, and thus without any mate they conceive by seed and generate
without conjuction, and the offsprings of these because of their longevity reach
a great age so that up to a hundred years of life a succession of them is produced,
and they do not die easily of needy old age. What say those who are accustomed
to smile at our mysteries when they hear that a virgin may generate and do they
esteem impossible bearing by an unmarried girl whose modesty no custom of
man violates ? Is that thing thought impossible in the Mother of God which
is not denied to be possible in vultures ? A bird bears without a mate and none
confutes it, and because Mar>' bore when betrothed they question her chastity.
Do we not perceive that the Lord sent beforehand many examples from nature
itself by which incarnations he proved the virtue of the suspected one and added
truth (to the story)."

In the last half of the sixth century, Cassianos Bassos mentions

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the wind-impregnation of vultures in his Geoponica (Blc XIV
Ch. 26). '

Concerning Vultures. Aristotle.

Aristotle relates that vultures are destroyed by the odor of unguents just as
beetles are by the odor of roses. For a rank odor is good for their hel^Jth. More!
over vultures do not unite sexually but fly headed into the South Wind and thus
become pregnant, and after three years they produce young.

St. Isidore of Seville (560-636 A.D.) in his Etymologiae sive
Urtgtnes, cited the wind-impregnation of vultures merely as a
natural curiosity. This work was written between the years 622
and 633 From Bk. XII, Ch. VII, 12-13—

"... The vulture is thought to be famous for slow flying; certainly because
of he greatness of its body it does no, have a quick flight They say thatTome

l^Z ""k"™"*': '". '°"'°"' ""* =°"«'^= """l «-""« without copulatio"
and offspring bom of these live almost a hundred years. And vultures ius
as eagles, perceive dead bodies beyond the seas; truly flying so high hey see
from above many things which are hidden in the obscurity of mountains.'^

A contemporary of St. Isidore, Theophylactos Simocattes
who lived 3t the other end of the Mediterranean Sea. wrote a
book on natural curiosities about 629 A.D. According to
Theophylactos vultures would copulate at certain times and
the women of Lemnos knew how to receive sailors. The following
extract is from his Dialogos, Chapter 8—

" Antisthenes —

" Many genera among birds do not have need of males in conulation for
conception. But others, indeed, which are full of wind do not bear VuUurt
however, have such a nature that they may bear without conjunction wirtimae'

ct'ofJTh" " r ™"r" '^ "™"'"'^ ""' •" "^ '«"• '<" ">e "ature of the fTmaU
cuts off he wh<Je sex from them. But a. times, when vultures fear the losTof
their children, the whole race devoted itself to copulation jus, af he Lemno,

7121^' T^ T "r '"■"• ^"^ **"« vu,.ures'crbi:e manyZd
Of cunning. They fly about against the south wind. But if there is J

south wind, to the neighboring southeas, wind they expand Their wi^; Then
they open and thus become filled with the inflowing wind- and thev rive hi«h
no. to wind-eggs but to living young. Therefore many iimes tfs nIL a'
for nature to complete the animal, Polycrates." necessary

During the next five hundred years the legend of the vultures
naturally underwent some changes. These are shown clearly in
the Htstonarwm variorum chiliades of John Tzetzes, a Byzantine
writer born m 1110 A.D. From Chap. XII—

animals impregnated by the wind

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" Some people, without investigation, say that vultures produce living things
and that they have both milk and breasts and other such things. As I have found
that tigers are all male so also I have found that the entire tribe of vultures are
female. And flying for five days with the wind against their rumps, they receive
properly the gonad from the wind. In 120 days they produce the wind eggs
and in as many days they hatch out (the young) from the eggs and in another
120 days they rear them up to the time of their flight. You have the fertilization
of the vultures, that is to say of the female generations, by the wind."

The Annales of Michael Glycas appeared sometime between
the years 1143 and 1156. From Pt. I, p. 42 (pp. 81-82, ed. of
1836)

" Now they say, among other things which differ even more from the beliefs
of men, that vultures produce young without the union of male and female, even
at an extreme age for they live to be a hundred years old. Why this ? Doubtless
to prevent them- (unbelievers) from mocking at our secret rite, just as if it were
impossible for a virgin to bear young and remain none the less a virgin. From
this birth God merely provided us with complete explanations for building our
faith in things otherwise absurd and contrary to our beliefs. Thus Basil discusses
these matters. Others, indeed, who have seen animal births, say that vultures
carry the foetus unnaturally for fully three years. Why this ? Listen ! No
male vulture is seen. Then without the union of two, they are accustomed to
produce according to the regulations of nature. As a result the complete breeding
of vultures exists in the females. Since they are not ignorant of this, in dread
of barrenness, they all simultaneosuly undertake something like a campaign
for impregnation. Thus if no South Wind presents himself to them, they spread
their wings toward the known East Wind. Thence they are filled with the wind
blowing in through their open mouths, and they bring forth animals, not wind
eggs. This is the reason why nature requires a very long time for the completion
of this animal. For it is difficult and laborious to draw a certain essence of birth
from the wind and much more delicate and exacting to form an entire animal
from it. Now, therefore, when you hear that a most pure virgin was made
pregnant without commerce with any man since the Holy Spirit had previously
overshadowed her and had dominion over her, as the prophet said who had come
from Thaemane, it by no means diminishes the faith in this but seems actually
to augment it notwithstanding its divergence from the beliefs of men. For
in the case of the vulture also, conception from God is very admirably established.
It is said in written records from India, however, that the vulture considers how
she should become pregnant, and finding a stone, which they call Eutocium because
it aids in the ease of delivery, places herself on it, shaking violently in a desire
for young, and thus accomplishing her purpose. This stone is round like a ripe
nut and, shaken like a bell, it gives forth a noise, another stone moving within.
This too is said about the vulture, that, when it is anointed thoroughly with
rose-perfume, it dies."

A hundred years later, the story appeared in Li Livres dou tresor,
written about 1266 by Ser Brunetto Latini, the friend and
perhaps the teacher of Dante. From Bk. I, pt. 5, Ch. 172 —

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ANIMALS IMPREGNATED BY THE WIND

III

" The Vulture.

" And they conceive without the union of male and female and bear offspring
which live more than a hundred years. And know that they do not peck at any
carrion without lifting it off the ground first. And they willingly go into the earth
with the great claws that they have."

The records of the wind-impregnation of vultures are brought
down to the fourteenth century by the work of Manuel Philes
(1295- 1 346), a Byzantine naturalist, who wrote De animalibus
proprietate. From (p. 21-23) Sec. 3 —

" Vultures.
" This genus produces not masculine but feminine offspring which are fecun-
dated by the South wind flowing into their body openings. Having received
this breeze they actually give birth to young after the third month, although
they have not been impregnated by a mate. Their nests, being without walls,
offer success to a forcible seduction, but this is never seen, according to the
experiences in Egypt. Here the vultures produced eggs fertilized by the seed
of the male, by which means vultures are created who are like their ancestors
neither in strength nor covering, since they are black just like the people dwelling
near Pharos."

LuDOVicus Caelus Rhodiginus (15 16) relates the story in
Lectionum antiquariim commentarii. From Bk. VIII, Ch. 18 —

" ... Or because, as the Egyptians relate, and as Ambrosius seems to prove
in the Hexaemeron, all vultures are female, and as trees are made pregnant by
the zephyr (W. wind), so vultures are made pregnant by Eurus (E. wind or S. E.)
or by Boreus (N. wind) as (Horapollon) reports. In reality it is very true that
powerful and sure signs are made therefrom. (Horapollon) reports that the
word " mother ", is portrayed by a hieroglyph in the image of a vulture.

Joannes Ravisius Textor(i52o) quoted the story of the vulture
as well as that of the mares. From page 445b (Ed. of 1524)

"... They say that this bird conceives without seed of the males and generates
without copulation, and their offspring acquire a great age up to one hundred
years. They are renowned for their odor. Black vultures are stronger, they
build their nests on high cliffs, no one lays hands upon their nests. They are
usually observed to have two offspring."

Ulisse Aldrovandi (1522- 1 605) devoted an entire chapter to
vultures in his Ornithology (Bk. Ill, Ch. I) first published in
1599. He described their wind foecundation in detail and reprinted
a number of the earlier descriptions. In the next century
NiEREMBERG in his Historia naturae ^ Antwerp, 1635, mentioned
the wind fertilization of vultures. He quoted the passage in

Theophylactos already cited and added (edition of 1675, Bk. VI,
Ch. 7, p. 956)—

" ... Antigonus writes, if they (the vultures) face the wind from the seas the
females conceive. The regeneration of the Phoenix, a virgin bird, must be con-
ceded to spontaneous origin. But since this is fabulous, so the other accounts
are not certain enough."

Hens

Pullets normally lay eggs after they reach a certain age even
if they have never been trodden by a cock, and thus the stories
of the impregnation of hens by the wind are on a somewhat different
plane from those of the mares and vultures. These sterile un-
fertilized eggs are sometimes referred to as virgin-eggs, but for
centuries they were known as wind-eggs and it was supposed
that they were produced by the fecundating power of the air.
Thus hens have to be included in the list of the wind's consorts.
Aristophanes used the term, wind-egg, in his play The Birds,
written in 414 B.C., (line 695) "In the beginning night laid
a wind-egg," and Aristotle described the production of such
eggs in considerable detail both in the De Generatione Anitnalium
(Bk. I, Ch. 21; Bk. II, Ch. 3; Bk. Ill, Ch. i) and in the Historia
Animalium (Bk V, Ch. I; Bk. VI, Ch. 2). These passages are
much too long to quote here. The following two short excerpts
are suflScient, however, to illustrate an extremely early stage in
the legend of the wind's activity. From Gen. Animal., Bk. Ill :
Ch. I, p. 749b.

" Some embryos are formed in birds spontaneously, which are called wind-eggs
and * zephyria * by some."

From Hist. Animal. , Bk. VI, Ch. 2.

" Such as affirm that wind eggs are the residue of eggs previously begotten
from copulation are mistaken in their assertion, for we have cases well authenticated
where chickens of the conmion hen and goose have laid wind eggs without ever
having been subjected to copulation... wind eggs are laid by a number of birds;
as for instance by the common hen, the hen partridge, the hen pigeon, the peahen,
the goose and the vulpanser."

Varro and Columella mentioned wind-eggs in their description
of the wind-impregnation of mares. On the other hand, Pliny

112

CONWAY ZIRKLE

gave the hens the dignity of a separate treatment. From Pt. I,
Bk. X, Ch. 60.

" Wind egs, which we call Hypenemia, come either by the mutuall treading
of hens one another, by an imaginarie conceit of the male, or els by dust. And
such egs not only Doves do bring, but house hens also, Partridges, Peahens,
Geese, and Brants, or the female Barganders. Now these egs are barren as one
would say, and never proove birds, lesse than others, not so pleasant in tast, and
besides more moist. Some are of opinion that the wind will engender them :
for which cause also they are called Zephyria (i.e. West-wind-egs :) and verily
such egs are seen only in spring, when that wind bloweth. Addle egges, which
some call Cynosura, are they that chill upon the nest,... »

Wind-eggs are mentioned by Athenaeos in his Deipnosophistae
(192-200 A.D.), perhaps the most famous cook-book in history,
and the wind-impregnation of hens is cited in the so-called
Recognitions of Clement (fourth quarter of the third century)
as evdience of the reasonableness of the virgin birth of Christ
just as Lactantius and St. Augustine had cited the story of
the mares and as Origen, Eusebius, St. Basil and St. Ambrose
used that of the vulture. From the Recognitions^ Bk. VIII, Ch. 25—

"... And not only males are produced, but females also, that by means of both
the race may be perpetuated. But lest this should seem, as some think, to be
done by a certain order of nature, and not by the appointment of the Creator,
He has, as a proof and indication of His providence, ordained a few animals
to preserve their stock on the earth in an exceptional way : for example, the
crow conceives through the mouth, and the weasel brings forth through the ear;
and some birds, such as hens, sonxe times produce eggs conceived of wind or dust;
other animals convert the male into female, and change their sex every year, as
hares and hyaenas, which they call monsters; others spring from the earth, and get
their bodies from it, as moles; others from ashes, as vipers; others from putrifying
flesh, as wasps from horse-flesh, bees from ox-flesh; others from cow-dung as
beetles; others from herbs, as the scorpion from the basil; and again, herbs from
animals, as parsley and asparagus from the horn of the stag or the she-goat."

Alexander Neckam referred casually to wind-eggs in his account
of the parthenogenetic foals. Bartholomew, the Englishman
(Bartholomeus Anglicus), also described the eggs but did not
mention the wind-impregnation of mares. The account is in
De proprietatibus rerum, Bk. 19, Ch. 79, written sometime between
1230 and 1240. The following quotation is taken from the
translation of John of Trevisa, printed at Westminster about
the year 1495 :

" Some eggs ben conceyved in every wynde : but they ben bareyn but they

<^ 4

animals impregnated by the wind

"3

ben conceyved of tredynge other by werkyng of the male and thyrlyd wyth Seminall
spyrj'te as he sayeth."

Albertus Magnus (1193-1280) completed De vegetabilibus
sometime before 1256. In his discussion of sex in plants he
mentioned wind-eggs. From I, I, 12 —

" Consequently, he says that a wind egg is not alive except potentially with
a plant's life, but not an animal's; by a ' wind egg ' he means one not impregnated
by the cock."

Two hundred years after Albertus Magnus, Archbishop Perot-
Tus mentioned wind-eggs. From the Cornucopiae (P. 365 edition
of 1521)—

•* ... a third which is worthless he calls urinum, as if full of the excrement of
urine. Indeed the Greeks call urine oZpov. Eggs from which nothing is born
are called worthless (Irrita) by us, Hypenemia by the Greeks, as if {,n6 rod dv^nov,
that is conceived by the wind, and on this account they are called Zephyria by
the Latins. These eggs are conceived by the imagination of desire of the females
within themselves wholly without intercourse with a male : some people think
that they are generated by the wind. In this they differ from the (Urina) because
the (Urina) are deserted at incubation, and others call these addle eggs (Cynosura)."

In 1570 Heresbach mentioned wind-eggs as well as the story
of the mares. From Foure Bookes of Husbandrie (tr. by Barnabe
Googe, Esq.), London, 1586, Ch. 169—

" The Hennes wyl treade one the others, but theyr Eggs never come to good,
but are wind eggs."

Harde in 161 1 changed the meaning of the term somewhat:

" an egge laied with a soft skin or filme instead of a shell : a soft sheld egge :
a wind egge."

One of the most amusing figurative uses of wind-egg occurs
in Wit without Money, a play by Beaumont and Fletcher
produced in 1614. From Act I, Scene I —

" Val. Who bid you get 'em ? Have you not threshing work enough, but
children Must be bang'd out o' th' Sheaf too ? Other Men With all their Delicates,
and healthful Diets, Can get but wind Eggs : You wi' a Clove of Gariick, A piece
of Cheese would break a Saw, and sowr Milk, Can mount like Stallions; and
must I maintain These Tumblers ?

Lance. You ought to Maintain us we."

This series of references to wind-eggs is brought to an adequate

8

r h «

114

CONWAY ZIRKLE

conclusion by the scientific discussion of William Harvey, famed
as the discoverer of the circulation of the blood. He wrote
De Generatione Animaliutn in 1666. From page 219 —

" The principal difference between eggs, however, is their fecundity or
barrenness — the distinction of fruitful eggs from Hypenemic, adventitious, or
wind eggs. Those eggs are called hypenemic, (as if the progeny of the wind,)
that are produced without the concourse of the male, and are unfit for setting;
although Varro declares that the mares, in Lusitania, conceive by the wind.
For Zephyrus was held a fertilizing wind, whence its name, as if it were Zu}r)<ft6po'i,
or life bringing. So that Virgil says :

" And Zephyrus, with warming breath resolves

The bosom of the ground, and melting rains

Are poured o'er all, and every field brings forth,
" Hence the ancients, when with this wind blowing in the spring season, they
saw their hens begin laying, without the concurrence of the cock, conceived
that zephyrus, or the west wind, was the author of their fecundity. There are
also what are called addle and dog-day eggs, produced by interrupted incubation,
and so called because eggs often rot in the dog-days, being deserted by the hens
in consequence of the excessive heat; and also becasue at this season of the year
thunder is frequent; and Aristotle asserts that eggs die if it thunders whilst
the hen is sitting,"

Women

When cause and effect are separated by a prolonged gestation
period, the connection between them is easily lost, even by peoples
whom we consider to be no longer primitive. We do not know
just when the parental function of the male was first discovered.
It could easily have been overlooked for a long while, for even
to-day it is not recognized by some savage tribes. The mysteries
of parturition were the subject of many primeval legends, however,
and consequently the literature of many races is filled with accounts
of miraculous births. The myths of the polytheistic Greeks and
Romans are particularly rich in these instances and in stories
of the philanderings of the Gods and Goddesses. It would have
been remarkable if, alone among the minor deities, the personified
winds had remained strictly celibate. Needless to say the winds
often aspired above mares, vultures and hens and, according
to Ovid (43-17 B.C.) two of them, Boreas and Zephyrus, mated
with nymphs. Significantly enough, the bride of the West Wind
was called Flora and she it was who controlled all flowers. She
describes her happy state in Ovid's Fasti V (tr. by Frazer) :
195, as follows —

f ^
t

1^

1. .

<- -

:(.

{- V

- 1

^■*N %

I.

i • y

V

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T

ANIMALS IMPREGNATED BY THE WIND 1 1 5

" A Grecian nymph was I, Chloris my name —

Change but some letters, it becomes the same

As Flora, mark you — and among the Good

Within the Happy Fields was my abode.

How fair I was it scarce beseems to tell :

But to a god I deemed desirable.

For as I roamed abroad, one April day

The West Wind saw me, and, though I said nay,

Took me by force, as Boreas before

Took Orthyia on the Attic shore.

Still, for that wrong by wedlock he atoned

Nor have I cause to grieve, as consort owned.

Eternal spring is mine, trees ever green

Earth clothed in herbage, azure skies serene;

And as my bridal gift and marriage dower

He gave me governance of every flower."

If we except the tales of the personified winds and nymphs,
we must look to China for the first stories of women being
impregnated by inanimate objects, and we must consult particularly
the works of the Chinese geographers. According to these savants,
three diff"erent countries of women were located among the outer
barbarians. The one they placed in the far west near Ta'ts'in
or Fu-lin (the Eastern Roman Empire) was doubtless the oriental
version of the country of the Amazons. The second country
of women supposedly somewhere to the south of China may
have been the land of the polyandrous hill tribes of India. The
third country of women, located on an island to the east of China,
is harder to identify. Schlegel was of the opinion that it was
purely mythological and originated from stories told by sailors
of the islands visited by seals and sea-lions. This is a most
plausible explanation of the peculiar inhabitants of this country
and of their anatomical characteristics, viz., females without
breasts but with the heads of Chinese women, males with the
heads of dogs, a language which sounded like barking, etc.

Of course these countries became confused in the minds of
many writers and in time the characteristics of each were transferred
to the others. Originally the country of the Amazons was merely
ruled by women; in the country to the south the women were
not remarkable except for the fact that they had many husbands,
but in the eastern island they employed novel means to perpetuate
the race. From the stories of the eastern kingdom of women
grew the legend of their wind-impregnation. The curious,

ii6

CONWAY ZIRKLE

muddled growth of this legend is best shown by a chronological
arrangement of the various records.

Hui-SHEN, a Buddist monk, seems to have been the first to
describe the eastern country of women. In the year 499 he
returned from a long journey to the east. Among the marvelous
stories he told of the lands he had visited was one of this country.
As translated by G. Schlegel the story reads—

" Nui Kouo : The Isle of Women.

" Hui-SHEN in the year 499 A.D. came to Ktng-tcheou, at that time the capital
ot the country, of Fousang, and related that one thousand miles east of Fousang
was the Country of women. That these women had a decent and correct ai^
and that they were very white, although their bodies were hairy and their hair
was so long that it trailed on the ground. That at the second or third
month they hasten to enter the water, and become pregnant in this way. That
they give birth the sixth or seventh month.

"That these women have no breasts at all on their chests, but that, behind
their neck, spnng out hairs; that from one of these white hairs a liquid comes out
that serves to suckle their children. These children, continues Hui-shen, can
walk a hundred days after their birth, from which I conclude that they are adults
trom their 3rd or 4th year (5) "

This fecundation by means of water instead of wind is a variation
of the theme which reappeared at intervals for nearly a thousand
years.

Li-Yen-shou wrote Nan-shih in the seventh century and in
It he noted a peculiar race living to the east. As translated by
Schlegel, —

"In the sixth year of the period Tien-kten of the Liang (507 of our era), a man
of Tsm-ngan (province of Fou-kien) who was crossing the sea, was cast by the
wmds upon an island; he set foot on land and found there inhabitants of which
U^ women resembled those of China, but whose language he could not understand.
1 he men had human bodies but heads like dogs (6), their voices resembled barking

(5) It is interesting to note that in the national epic of Finland, the Kalevala
the Virgin of the Air was impregnated by both wind and water. From Runo l'

mes 131-136 (trans, by Kirby)—

" Thus the tempest rocked the virgin
And the billows drove the maiden.
O'er the ocean's azure surface.
On the crest of foaming billows.
Till the wind that blew around her.
And the sea woke life within her."

(6) More than eight-hundred years later these people were described by a
Christian traveller, Friar Jordanus who wrote Mirabila descripta, 1 324- 1330;—

4

ANIMALS IMPREGNATED BY THE WIND

117

«

#■■ '

■I

* ■ ♦

* *

*ih

A • »

Their food consisted of little beans and their clothes were as if made of cloth.
They constructed walls of clay of a round form with an entrance like a cave."

Chang Wen-ming (634-713 A.D.) wrote Ta-fang hsi-yii
chHu-fa hao-seng ch'uan probably in 648. He confused thoroughly
the eastern with the western country of women. The Roman
Emperor (king of Fo-lin), is depicted as being moved by enlightened
self-interest, as follows —

•' On an isle situated to the southwest of the realm of Fo-lin, is found the realm
of eastern women. One sees only women, not a single man. This country
contains a great number of rare and precious things which are sold in the realm
of Fo-lin. That is why the king of Fo-lin every year sends men (7) to unite with
them ; but if they give birth to boys, the custom of the country does not permit
them to be raised."

During the T'ang Dynasty (618-907 A.D.) appeared Ta-huan-
hsing'ching-chi. As quoted by Ma Tuan-lin —

" There is also a report that in the west there is the country of women who,
being affected by the influence of water, give birth to children."

By the tenth century the tale of the Island of Women reached
the Mohammedan world. The Arabian geographers located this
land in some of the most inaccessable regions, generally far to
the east. Buzurg-ibn-Shakriyar (912-1000 A.D.) described this
country in his collection of sailors' stories, the Marvels of India,
written shortly after 954. The following excerpt shows the legend
in a most primitive form. It is taken from the French translations
of L. Marcel Devic. Pt. I, Ch. XX—

" There are many other different islands in which are men having the heads
of dogs, but their women are said to be beautiful. I cease not to marvel at the
great variety of islands that there be."

(7) From Marco Polo (Bk. IV, Ch. 3) written in 1299 A.D. —
" Distant from Kesmacoran about five hundred miles toward the south, in
the ocean, there are two islands within about thirty miles from each other. One
of these is inhabited by men without the company of women, which is called
the island of Males : and the other by women, which is called the Island of
Females... The men visit the Island of F'emales and remain with them for three
successive months, namely March, April, and May, each man occupying a separate
habitation along with his wife. They then return to the male island, where
they live the rest of the year, without the society of any female... This mode
of living is occasioned by the peculiar nature of the climate, which does not allow
of their remaining all the year with their wives unless at the risk of sacrifice."

n •* ►

■j

ii8

CONWAY ZIRKLE

ANIMALS IMPREGNATED BY THE WIND

119

" During these transports (of joy), suddenly from the interior of the island
arrived a mob of women whose number God alone could count. They fell
upon the men, a thousand or more women to each man. They dragged them
toward the mountains and forced them to grow their instruments for their pleasure.
There was a continuous struggle bet\^'een them and a man belonged to the strongest
The men died of exhaustion one after another; and each time that one died, they
still fell upon him without being bothered by the stinking odor of the corpse.
A single man survived, he was a Spaniard whom one woman had carried off
She visited him at night, and at dawn she hid him near the sea and carried him
food. Finally the wind changed and commenced to blow in the direction of
the land of India whence the fleet had come. The man took a small boat called
Felou and supplied it during the night with water and food. The woman, seeing
his plan, led him to a place where, having pushed aside the earth, she revealed
a mine of gold dust. They filled the boat with as much as it could hold Then
they both set out and after ten days of sailing reached the port whence the vessel
came. There, he told of his adventure.

" The woman lived with the Spaniard, learned his language, became a Moham-
medan and bore him several children. Questioned about that island and those
women who lived away from the society of men, she spoke thus :

" • ... One should know that, by the will of God, the women of this country
bear first a boy. the second time two girls, and continue thus alternately the rest
of their lives. Thus it happens that men are scarce in our country, and the women
becoming more numerous, would dominate them. So the men would equip
ships, embark with thousands of the women and throw them on that island saying
to their God, the Sun : " To you belongs the right to what you have created;
as for us, we have more of them than we can use."

" ' The women were thus left on the island, where they die one after another
No man had passed among us before you came. Never had one approached
Because our island is situated in a vast sea, under Canopus ; and no traveller
could visit there and return; none would dare leave the shore and solid dry land
for fear of being drowned in the ocean. Thus has willed the all Powerful. Blessed
be God, the best of creators.' "

The Moslem, Ibrahim ibn Wasif-shah, described the island
of women. His version appeared in 1031 A.D. and in this version
the mhabitants are impregnated by the wind. This tale appears
to be only a variant of the older Chinese legend where water is
the fecundating agent. As translated by Farrand—

" The hie of Women. This is an island situated at the limits of the China
Sea. It IS said that it is inhabited only by women who are fertilized by the wind
and who bear only females; it is said that they are fertilized bv a tree whose fruit
they eat (8). Gold, it is claimed, shoots up, with them, in caves, like bamboo
and they are nourished with the gold. Once a man fell among them ; they wished

(8) This method of impregnation is found in many other legends. According
to Ovid, Juno became pregnant by eating an herb given her by Flora and in due
time she g^ve birth to Mars. In the Kalevala Marjatta finds all of her precautions
to no avail and she becomes enceinte from eating a cranberry.

'I I*

K

^

.1.

I

to kill him; but one of them had pity on him, placed him on a log and committed
him to the sea. The waves and the winds bore him to the country of China.
He went to find the king of China and told him of the isle. The king sent vessels
to search for it but after three years of effort, they found no news nor traces of it.
... Others resemble women having hair and breasts; there is no male in this race;
these females are made pregnant by the wind, and they bear only individuals
which resemble them ; they have ravishing voices and they attract many individuals
of other races by the charm of their voices."

Ou-yang Hsiu (1007-1072) was a contemporary of Ibrahim.
In his Hsin T'ang shu he repeated the earlier story of Chang
Wen-ming. As translated by Julien —

** According to the Hstn-T'ang-shu an island in the south-west of Fu-lin is
inhabited by a tribe called " western women " who are all females. The country
contains many precious articles and is a dependency of Fu-lin. The rulers
of Fu-lin send males to them every year to couple with them. It is their custom
not to bring up the male children they have borne. On the eastern sea there
are likewise women with a female government, which is the cause of these being
called eastern women."

Approximately a hundred years after Ou-yang Hsiu, al-Mazini
(1080-1170) wrote Tuhfat al-albah, etc. (1062). His story of the
women is obviously from Chinese sources. As translated by
Farrand —

" It is said that in the deserts of Magrib there is a tribe of descendants of Adam
all of which are women. There are no males among them, and none are seen in
their country. These women enter into a water (p. 47) which is found among
them and they become pregnant from this contact with the water. Each female
bears a daughter and never bears a boy... But Allah is the most wise."

While the Moslem, al-Mazini, had the women fertilized by
water, the Chinaman, Ch'ou Ch'u-fei, had them impregnated
by the wind. From the Ling-wai'tai-ta written in 11 78 (tr. by
Farrand) —

•' The Isle of Women.

" More to the southeast there is the realm of Women. The water flows
unceasingly toward the east and overflows once in many years. On the water
float grains of lotus more than a foot long and peach stones two feet long. When
the inhabitants gather them they offer them to the queen. In days of old there
was a vessel of commerce which was cast upon this kingdom. The women seized
(the sailors) and took them away. At the end of several days they all died. One
alone (more) discreet (than the others) got possession of a boat during the night
and at the risk of his life succeeded in escaping. Afterward he related the affair.
The women of this country, when the wind blows from the south, undress and
conceive by the wind, all give birth to girls."

}

I20

CONWAY ZIRKLE

Ibn Tufail (1150-1185) explains the miraculous births by a
new method, (tr. by Farrand) —

" Our virtuous predecessors report— May God be satisfied with them !— that
there is an isle of India, situated under the equator, in which man is born without
father nor mother. There is a tree there which in the guise of fruit, produces
women ; it is of these that Masudi speaks under the name of daughters of Wakwak."

Chao Ju-Kua described the land of the women in 1225 by
quoting the passage in the Ling-wai-tai-ta of Ch'ou Ch'u-fei.

Al Qazwini's (1203-1285) revised geographical writings date
from 1276. In the following passages he describes two of the
countries of women (tr. by Farrand).

*' The Isle of Women in the China Sea. Women are found there and not a
single man. They conceive by the wind and bring into the world females like
themselves. It is also said that they conceive by eating the fruit of a tree which
grows in their isle. They conceive and bring females into the world. A merchant
relates that the wind bore him towards this isle. ' I saw here he said, women
but not a single man with them. I saw in this isle gold as abundant as the ground
and reeds of gold like bamboo. They wished to kill me, but one of them preserved
me, put me on a plank and abandoned me to my fate on the sea. The wind
brought me to China and I informed the sovereign of China of the nature of this
island and the gold there was there. He sent out people in exploration who
were absent three years, they never reached the isle and they came back again."

In the next passage Qazwini describes women who kill all
of their male offspring but keep male slaves.

" THE CITY OF WOMEN ! This is a large city situated on an island of
the sea of Maghrib. Turtusi says that its population is composed of women,
over whom men have no power. They mount a horse and make war themselves;
they are full of ardor and attack. They have male slaves; each of them comes
in turn to pass the night with his mistress. He stays near her all night, and
rises very eariy to leave without being seen before dawn breaks. When one of
them bears a male child, she kills it immediately, but if it is a giri it is allowed
to live. And Turtusi adds; the city of women certainly exists; there is no doubt
about that."

Ma Tuan-lin completed Wen hsien fung k'ao in 1280 but
did not publish it until 13 19. Schlegel describes his references
to the country of women.,

" ... That these people had the practice of plunging the virgins into the sea on
the seventh month. They relate again that there was a country in the sea inhabited
entirely by women, without there being a single male there, etc., etc...

" The Ainos have a like tradition, with this difference that the women of this
isle are said to be cannibals and kill men who have been shipwrecked on the

ANIMALS IMPREGNATED BY THE WIND

121

.

island after having abused them. Others say that these women become pregnant
on leaving the bath, and facing the South wind, or, according to the Ainos, the East
wind."

Twenty years after Ma Tuan lin, Hamduallah Mustafai
(1339) related the story of the women with a number of variations
(tr. by Farrand).

" ... In the middle of this desert of sand rises a town which is inhabited only
by women. If a man goes to establish himself there he loses all his virile qualities,
due to the effect of the climate, and he soon dies. The propagation of the race
is assured by means of a spring where the women go to bath, and which makes
them pregnant, and they give birth to giris ; if a boy is bom unexpectedly from
time to time he always dies young. When these women are purified of their
menses, they reappear the second day after they have bathed in this spring, and
so much blood flows that they are in danger of death.

" By an effect of the whole power of God most High, these women never have
any passions; this is to such a degree that if one of them arrives at this province
(le Maghrib), when a man has commerce with her, she becomes seriously sick;
but when she stayed here a long time and has become used to this climate, the
desire of passion takes her also."

About 1340, Ibn al-Wardi described the Island of Women
(tr. by Farrand) —

" THE ISLE OF WOMEN is large. They say that it has never had men
on it. They relate that the women conceive by the wind. When they have
become pregnant they give birth to (females) like themselves. Some people claim
that in this isle there is a kind of tree whose (fruit) makes (the women) conceive
when they eat it. Gold (springs up) in this country; its stems are like those
of the bamboo. The earth also contains (mineral) gold, but the women (who
inhabit this isle) pay no heed to it.

" It is related that a man was cast upon this isle by God. The women wished
to kill him, but one of them had pity on him, put him on a small boat and
abandoned him to the sea. He became the sport of the waves which bore him
in to an isle of China. He related to the king (of this isle) what he had seen
in the Isle of Women, in what concerned its inhabitants and the abundance of gold.
The king sent ships and men with (the individual in question). They sailed
for a long time on the sea, in search of the isle (of Women) but they found no
trace of it."

al-Bakuwi abstracted the geographical work of al-Qazwini
in 1403- 1404. His short account of the island of women follows —

"THE ISLE OF WOMEN. An island in the neighborhood of China; it
is inhabited only by women. It is said that they become pregnant by the wind,
or by eating the fruit of a certain tree which is in the country."

In the fifteenth century the Portuguese made the Arabia-India-

122

CONWAY ZIRKLE

China trade very precarious and as a result the contacts of the
Mediterranean Moslems with the Far East were very effectually
broken. The story of the wind-impregnated women, however,
continued to reach Europe. Magellan's sailors heard the story
in the East Indies on the first circumnavigation of the earth.
The scribe of the expedition, Antonio Pigafetta, included the
tale in the First Voyage round the World written in 1522. From
the Hakluyt Soc. Pub. No. 154; Vol. II, p. 169—

" Our oldest pilot told us that in an island called Acaloro which is below Java
Major, there are found no persons but Women, and that they become pregnant
from the wind. When they bring forth, if the offspring is a male, they kill it,
but if a female they rear it. If men go to that island of theirs, they kill them if
they are able to do so."

Ultimately, of course, this story was not taken at its face value.
John Milton in U allegro (1640) found the legend very useful
for its poetic effects.

" Or whether (as some sages sing)

The frolic wind that breathes the spring

Zephyr, with Aurora playing,

As he met her once a-maying.

There, on beds of violets blue,

And fresh- blown roses washed in dew,

Filled her with thee, a daughter fair.

So buxom, blithe and debonaire."

We would expect the stories of wind-impregnated females to
die a natural death sometime within the sixteenth century, at
least among the European scientists, even if the stories should
remain alive in the orient. It is certain that the intellectual leaders
no longer believed in the fecundating power of air. On the
somewhat lower level of Natural Theology, however, the legend
had a strange rebirth, surprisingly enough in the early eighteenth
century. William Wollaston was a London clergyman, who
published for his friends in 1722 The Religion of Nature
Delineated (9). This work became so popular that it was re-issued
publicly in 1724, 1726, 1731, 1750, etc. Wollaston was
committed to the theory that the souls of children were not
generated by the souls of their parents and in order to support

(9) The type of this book was set in part by Benj. Franklin when he wat
a printer in London.

animals impregnated by the wind

123

this view, he postulated an aerial origin for spermatozoa. Section
five of his book was called '* Truths relating to the Deity." The
following extract is from truth number fifteen. From p. 89.—

"... If then the semina, out of which animals are produced, are (as I doubt
not) animalcula already formed; which, being distributed about, especially in some
opportune places, are taken in with aliment, or perhaps thru the very air; being
separated in the bodies of the males by strainers proper to every Kind, and then
lodged in their seminal vessels,do there receive some kind of addition and influence;
and being thence transferred into the wombs of the females, are there nourished
more plentifully, and grow, till they become too big to be longer confined : I
say, if this be the case, why may not those additions and the nutriment received
from the parents, being prepared by their vessels, and of the same kind with
which they themselves are nourished, be the same in great measure to the
animalcula and embrya that it is to them, and consequently very much assimilate
their young, without the derivation of anything else from them? Many
impressions may be made upon the foetus, and tinctures given to the fluids
communicated to it from the parents; and yet it, the animal itself, may not be
originally raised in them, or traduced from them. This hypothesis (which has
long been mine) suggests a reason, why the child is sometimes more like the father,
sometimes the mother : viz. because the vessels of the animalculum are disposed
to receive a greater proportion of aliment sometimes from the one, sometimes
from the other : or the fluids and spirits in one may ferment and operate more
strongly than in the other and so have a greater and more signal effect. (Here
it ought to be observed, that tho what the animalculum receives from the father,
is in quantity little in respect of all that nutriment, which it receives by the mother;
yet the former, being the first accretion to the original Stamina, adhering
immediately, and being early interwoven with them, may affect it more.) " (lo)

(10) Strange as it may seem to us, this passage greatly influenced James Logan
(1674-1751) a Colonial Governor of Pennsylvania. In 1735 Logan proved
that the pollen of Zea Mays was the male element and that it was necessary for
the production of viable seed (Phil. Trans. Roy. Soc. London 39 : 192-195^.
His results were republished four years later, Experimenta et Maletemata de
Plantarum Generatione, Leyden, 1739, and translated into English in 1747. Logan
apparently thought that no really frugal Deity would have invented a method
as wasteful as wind-pollination without some hidden reason. The following
passage is from the English translation. Experiments and Considerations on the
Generation of Plants, London, 1747.

" As therefore the Article of Generation has to this Time lain under insuperable
Difficulties, and as all the methods of accounting for it, hitherto invented, have
at last been found defective, why may we not suppose, that Nature herself points
out the whole Process of the Affair; especially in forming the seminal Substance,
distinct from the rest of the Seed ? namely, that the Farina is for this Purpose
committed to the Air, that it may receive out of the Air this little Seed or Plant,
praeexistent and completely formed, tho' in Stamina inconceivably minute and
invisible; and thus become pregnant hereby? It is drawn by an inherent
attractive Force (like as the Males and Females of Animals we see are mutually,
and even at a Distance, affected) first into the Style, and thru' that it slides by

124

CONWAY ZIRKLE

To the best of my knowledge, this contribution of Wollaston's
has had no direct effect upon biological thought, (ii) Indirectly,
however, it has exerted a beneficient influence upon all scientific
development, for a copy of the 1750 edition of Wollaston's
book fell into the hands of Sir John Hill, M.D. At that time
Sir John was greatly chagrined for he had just failed to be
elected to the Royal Society and he was inspired by Wollaston
to attack the Society, an attack which was so devastating that it
altered the whole tone of the Society's publications.

proper Canals to the Ova: And from this Farina, nourished by the Juices
of the Plant for the Purposes above described, the Bulk of the Seed is formed.
Lastly, the little Plant hid in the Seed, and cloathed with a terrestrial Matter,
which it borrows from the Farina, exerts itself; and, increasing by proper Nutri-
ment, which it draws from the Earth, at length springs up.

Nor perhaps is this Method of accounting for Generation to be confin'd to
Vegetables only; but it may, with equal Reason, be applied to the Production
of every living Creature that makes a Part of this Universe of Things. ' Whatever
in this World have Life (Plants as well as Animals) first of all exist completely
formed in Miniature, or in their first Rudiments, and are taken from the Air
or Aether, then, being cloathed with an earthly Substance, they increase and
grow larger. Nor is Generation any thing else than the placing this Principle
in the male Seed, where it acquires an earthly Nature, tho' not without some
Portion of an Aura coelestis, and a proper Conveyance of it into a suitable Matrix,
where it may grow and increase.' This was the Opinion of the ingenious Claude
Perrault long ago, as well as of the celebrated Wollaston our Countryman, as
appears from the Writings of these great Men."

<ii) BENOtT DE Maillet (1656-1738) gave in Telliamed : or the World Explained,
Amsterdam, 1748, an account of fertilization, which is almost identical with
Wollaston's. The following quotation is from the Baltimore edition of 1707
pp. 264, 265 : '^'*

" In order to understand this oeconomy of nature, imagine to yourself. Sir,
that the whole extent of the air... is full of the seeds of everything which can
live on earth...

When the male has arrived at a certain age, the seeds of his species re-unite
m him, by the air which he respires, and the aliments with which he is nourished,
according to a general law of nature, which wills that every thing should tend
to be attached to its own species. Then these seeds are prepared for fertility
m the vessels of the male, by the dispositions which puberty has put into them.
If with a good microscope you examine the seed when warm, you will see it
composed of small animals, like fish, which move up and down, but after the
seed IS grown cold they lose the motion, and, no doubt, the life which they had
acquired m these vessels. Hence it is evident, that these seeds receive in the
vessels of the male, a disposition to life, and to augmentation, which they had
not when they were introduced thither.

These vessels therefore are a kind of first uterus, where the seeds are prepared
for a greater growth, which they are to receive in the uterus of the female."

ANIMALS IMPREGNATED BY THE WIND

125

Sir John began the fight with Lucina sine Concubitu, a letter
humbly addressed to the Royal Society, London, 1750, purporting
to be written by one Abraham Johnson, a provincial doctor
of medicine. In this essay, Wollaston is quoted and the
quotation followed by an ironical passage, an excellent parody
of many of the papers in the Philosophical Transactions. The
collection of the floating spermatozoa is described as follows.

" Accordingly, after much Exercise of my Invention, I contrived a wonderful
cylindrical catoptrical, rotundo-concavo-convex Machine (whereof a very exact
Print will speedily be published for the Satisfaction of the Curious, designed
by Mr. H-y-n, and engraved by Mr. V—stu) which being hermetically sealed
at one end, and electrified according to the nicest laws of Electricity, I erected
it in a convenient Attitude to the West, as a kind of Trap to intercept the floating
Animalcula in that prolific Quarter of the Heavens. The Event answered my
Expectation; and when I had caught a sufficient number of these small, original,
unexpanded minims of Existence, I spread them out carefully like Silkworm
Eggs, upon white Paper; and then applying my best Microscope, plainly discerned
them to be little Men and Women exact in all their Limbs and Lineaments,
and ready to offer themselves little Candidates for Life, whenever they should
happen to be imbibed with Air or Nutriment, and conveyed down into the Vessels
of Generation."

These animalculae were fed to the doctor's Chambermaid in a
"Chymical" preparation. She later became astonished at her
condition and the experiment was reported a success.

Although wind fertilization was now a joke in Europe it was
still believed in the East Indies. In 1783, William Marsden
published his History of Sumatra, where, in reference to the
Natives, he wrote (p. 297, Ed. of 1811) —

" Thus they believed that the inhabitants of the island Engano to be all females,
who were impregnated by the wind; like the mares in Virgils Georgics."

Bronislaw Pilsudski, an anthropologist, found this same belief
still existing among the Ainu in the twentieth century, and
described it in his Materials for the Study of the Ainu language
and folklore, 191 2. From p. 91. —

" In Yeso, an old man assured me that there was a whole island inhabited
by women like the one in this tale. They were, however, able to bring forth
children by exposing themselves to the East Wind, by which they became pregnant.
They used to kill all their male children, and kept only their daugthers. "

126

CONWAY ZIRKLE

Conclusions

An examination of these descriptions of wind-impregnation
reveals a number of interesting problems. The questions which
naturally present themselves are — how did the stories originate,
what was their raison d'etre? Of course no definite answer can
be given, for the tales are extremely old. Homer knew of the
penchant of the west wind for the mares and Aristophanes
casually referred to wind-eggs. The fecundation of vultures by
the wind was an ancient Egyptian myth, we do not know how
ancient, but it has the earmarks of extreme antiquity. These
stories may even be relics of neolithic folk-lore, although it must
be granted that the mere substitution of some inanimate impregna-
ting object for the male is no proof of their barbaric origin, for,
even in the early civilizations, the role of the father was difficult
to establish empirically. Experimentation rarely gives un-
ambiguous results in the absence of adequate controls, and con-
sequently, the necessary prerequisites for parturition were not
always recognized. The ancient myths of many peoples contain
accounts of virgin births.

The real proof of the primitive character of the legends is
shown by the choice of the wind as the fertilizing agent. To
early man there was something immaterial about the wind. It
could not be grasped, held or weighed. It blew where it listed
and he could hear the sound thereof but could not tell whence
it came or whither it went. Moreover in times of storms it
was a power worth propitiating, but the breeze which blew in
and out of the lungs was beneficial and even necessary for life.
Thus the *' breath of life " seemed a part of a great immaterial
force. We have only to note the origin of our words for the
spiritual to realize how the peculiar properties of moving air
influenced the early interpretations of the universe. The Greeks
used the word pneuma for both air and spirit. The Latin spiritus
was a breath as well as a ghost and the Germanic ghost or geist
originally meant a breeze (gust). Even as recently as a hundred
years ago, it would have taken a very bold philosopher to deny
that a pneuma, a spirit or a ghost could cause a virgin to bear
off"spring.

If

■I-

T

■1-

L

ANIMALS IMPREGNATED BY THE WIND

127

%

I

-*** •*

The actual transmission of the stories of mares, vultures and
hens offers no serious problems. They passed from the Greeks
to the Romans and from the Romans to the medieval Europeans.
The only noteworthy aspect of their transmission is the credulity
with which they were received. With a few minor exceptions,
they were repeated without critical or skeptical comment. Indeed,
during most of the two thousand years that they were accepted
at face value, skepticism itself was not altogether reputable :
it was certainly not a habit of mind. We will probably never
know just how much intellectual standards delayed the growth
of science.

The use of the stories of the mares, vultures and hens to support
the dogma of the Virgin Birth gives us an interesting side light
on the beliefs of the classical Pagans, and on the methods used
by the Christians to secure converts. No less than eight of
the early Fathers cited the wind's fecundating power to show
that the Virgin Birth of Christ was not intrinsically unreasonable.
The wind-eggs of hens are referred to in the Recognitions of
Clement. Origen, Eusebius, St. Basil, St. Ambrose and
St. Isidore described the wind-impregnation of vultures while
Lactantius and St. Augustine referred to a like fertilization
of mares. Their arguments must have been considered very
effective for they were in use for several hundred years.

The stories of wind-impregnated women are not as easy to
trace. Originally there seem to have been several different legends,
which became so confused in the course of time that their individual
elements were hopelessly mixed. These stories probably origin-
ated in China. In the earlier versions the women could be
impregnated either by men or by bathing in certain waters, and
it was only in the later form of the story that they were wind-
impregnated. Our first record of this latter method is Arabic.
This is no proof, of course, that the wind-impregnation represents
an Arabic contribution to the legend, for our present sources
of information are so incomplete that we cannot be positive
that no Chinese geographer described this means of fecundating
women before the time of Ibrahim ibn Wasif-Shah. Indeed
the wind-impregnation may have been in the stories from the very
beginning. It certainly appears in the earliest Ainu accounts.
Just how it managed to supplant the other version is uncertain.

< »,

128

CONWAY ZIRKLE

Whatever the method, it remained in both the Chinese and Arabian
accounts for several centuries. When these stories reached Europe
their resemblance to the stories of the mares and vultures was
noted and commented upon, and finally, by the middle of the
eighteenth century, they were no longer accepted as true. As
recently as 1912, however, the Ainu tribes in Northern Japan
believed that there was a country where there are no men and
where the women were impregnated by the wind.

Chronological List of the Descriptions of Wind-Impregnation.

^1

Date

800 -f B.C.

414

384-321
118-30

31

43-17
30-40 A.D.

77

25-101

46-125

156-200?

192-200

200 -f

248

200-280

275-300?

260-340

318

300-400 ?

330-379

334-397
400

400

413-426

499

550-600

622-633

629

618-907?

1031

mo

H43-1156
1162
1 178
H05-1185

Author

Homer

Aristophanes

Aristotle

Varro

Virgil

Ovid

Columella

Pliny

Silius Italicus

Plutarch

Oppian

Athenaos

Aelian

Origen

The Recognitions of Clement

Soltnus

Lactantius

Eusebius

Hora polio

St. Basil

St. Ambrose

Claudian

Justin

St. Augustine

Hui-shen

Cassianos Bassos

St. Isidore

Theophylactos

Tu-huan-hsing-ching-cki

Ibrahim ibm Wasif Sah

Tzetzes

Michael Glycas

Al-Mazine

Ch'ou ch'ii-fir

Ibn Tufail

Animal Impregnated

Mares

Hens

Mares, Vultures, Hens

Mares, Hens

Mares

Nymphs

Mares, Hens

Mares, Hens

Mares

Vultures

Tigers

Hens

Ewes, Vultures

Vultures

Hens

Mares

Mares

Vultures

Vultures

Vultures

Vultures

Tigers

Mares

Mares

Women •

Vultures

Vultures

Vultures

Women •

Women

Vultures

Vultures

Women •

Women

Women

ANIMALS IMPREGNATED BY THE WIND

129

vb

r I •

fl^

I

•• !►

^>

f.

-I-
X

4

I

1200

1225

I 230- I 240

1246

1256

I 260- I 266

1268

1276

1295-1346

1319

1339

1340

I 400- I 450

1430- 1480

1516

1520

1522

1570

1599
1611
1614
1632

1635
1640
1651
1691
1722
1750

1783
1912

Alexander Neckam

Chao Ju-Kua

Bartholomew the Englishman

Walter of Metz

Albert the Great

Ser Brunetti Latini

Roger Bacon

Qazwini

Philes

Ma-tuan-lin

Hamdalla Mustawfi

Ibn-al-Wardi

Bakuwi

Perottus

Rhodiginus

Ravisius

Pigafetta

Heresbach

Aldrovandi

Harde

Johnston

Beaumont & Fletcher

Nieremberg

Milton

Harvey

Camerarius

Wollaston

Hill

Marsden

Pilsudsky

Mares, Hens

Women

Hens

Mares

Hens

Vultures

Mares, Vultures

Women

Vultures

Women •

Women *

Women

Women

Mares, Hens

Vultures

Mares, Vultures

Women

Mares, Hens

Vultures

Hens

Mares

Hens

Mares, Vultures

Women

Hens

Mares, Hens

Women

Women

Women

Women

• Impregnated by water.

The following bibliography lists only those works which are not cited adequately
in the body of the paper.

Bibliography

BuzuRG IBN Shahriyar. — Marvels of India (tr. by L. Marcel Devic), Leiden, 1886.
Ferrand, Gabriel. — Relations de Voyages et textes g6ographiques arabes, persans

et turcs. Paris, 191 3.
d'Hervey de Saint-Denys, M. J. L.— Ethnographic des peuples Grangers k

la Chine. 2 Vol. Paris, 1876- 1883.
Hirth, Friedrich. — China and Roman Orient. Leipzig & Munich, 1885.
HiRTH, Friedrich and Rockhill, W. W. — Chau-Ju-kua. St. Petersburg, 1911.
Marsden, William. — History of Sumatra, etc. London, 1783.
al-Mazini. — Le Tuhfat al-Albab (tr. by Farrand). Jfour. Asiatique ser. 12,

Vol. 6, p. 47, 1925.
Pilsudski, Bronislaw. — Materials for the study of the Ainu language and folklore.

Cracow, 19 12.

130

CONWAY ZIRKLE

RocKHiLL, W. W. — Land of the Lamas. New York, 1891.

ScHLEGEL, GusTAV. — Problemes Geographiques IIL Le pays des Femmes. T*oun(>

Pao 3:495-510, 1892.
Thompson, D. W. — Glossary of Greek Birds. Oxford, 1895.

Philadelphitty Pa.

Conway Zirkle.

'7

fi»

LA

I

J.
i

I

Reprinted from The American Naturalist, Vol. LXX,
pages 529-540, Noveinber-Deeeinl)er, 1930.

FURTHEE NOTES ON PANGENESIS AND THE

INHERITANCE OF ACQUIRED

CHARACTERS

PROFESSOE CONWAY ZIRKLE

Department of Botany and Morris Arboretum,
■ University of Pennsylvania

In a recent issue of the American Naturalist (Septem-
ber-October, 1935), the writer reprinted a number of
ancient and medieval accounts of pangenesis and the in-
heritance of acquired characters. Indeed, the classical
and medieval naturalists, almost without exception, be-
lieved that acquired characters were heritable, and from
400 B.C. until the nineteenth century the doctrine was
accepted as a matter of course. The hypothesis of pan-
genesis can also be traced to the fourth century before
Christ and, although it was not accepted unanimously, it
was believed by the majority of the medieval philosophers
and physicians. It is interesting, thus, for us to note that
both the doctrine of inheritance that we call Lamarckian
and the pangenesis speculations of Darwin were but
restatements of accepted classical views.

In the following pages the writer has collected from
the older literature a number of passages, not included
in the earlier paper, which describe pangenesis and the
inheritance of acquired characters. While these excerpts
are not very important individually, their number and
distribution through the centuries are such that thev
enable us to view more accurately the historical back-
ground of the nineteenth century naturalists. Of course
no definitive collection of these passages can be made at
present, nor perhaps would such a collection be desirable
if enough records could be assembled to show how com-
pletely these hypotheses were incorporated in medieval
philosophy.

Hippocrates was probably the first to describe pan-
genesis. He stated, **For the seed comes from all parts

529

13©

CONWAY ZIRKLE

RocKHiLL, W. W. — Land of the Lamas. New York, 1891.

ScHLEGEL, GusTAV. — Problcmes Geographiques IIL Le pays des Femmes. T*oung

Pao 3:495-510, 1892.
Thompson, D. W. — Glossary of Greek Birds. Oxford, 1895.

Philadelphiay Pa.

Conway Zirkle.

<3

I

^

>

«j»!

Reprinted from The American Naturalist, Vol. LXX,
pages 529-546, Noveiiiber-December, 1936.

FURTHEE NOTES ON PANGENESIS AND THE

INHERITANCE OF ACQUIRED

CHARACTERS

PROFESSOR CONWAY ZIRKLE
Department of Botany and Morris Arboretum,
• University of Pennsylvania

In a recent issue of the American Naturalist (Septem-
ber-October, 1935), the writer reprinted a number of
ancient and medieval accounts of pangenesis and the in-
heritance of acquired characters. Indeed, the classical
and medieval naturalists, almost without exception, be-
lieved that acquired characters were heritable, and from
400 B.C. until the nineteenth century the doctrine was
accepted as a matter of course. The hypothesis of pan-
genesis can also be traced to the fourth century before
Christ and, although it was not accepted unanimously, it
was believed by the majority of the medieval philosophers
and physicians. It is interesting, thus, for us to note that
both the doctrine of inheritance that w^e call Lamarckian
and the pangenesis speculations of Darwin were but
restatements of accepted classical views.

In the follow^ing pages the writer has collected from
the older literature a number of passages, not included
in the earlier paper, which describe pangenesis and the
inheritance of acquired characters. While these excerpts
are not very important individually, their number and
distribution through the centuries are such that they
enable us to view more accurately the historical back-
ground of the nineteenth century naturalists. Of course
no definitive collection of these passages can be made at
present, nor perhaps would such a collection be desirable
if enough records could be assembled to show how com-
pletely these hypotheses were incorporated in medieval
philosophy.

Hippocrates was probably the first to describe pan-
genesis. He stated, **For the seed comes from all parts

529

INTENTIONAL SECOND EXPOSURE

Tr>r>

TT A T^ r^ A

530

THE AMERICAN NATURALIST [Vol. LXX

No. 731]

ACQUIRED CHARACTERS

531

of the body, healthy seed from healthy parts, diseased
seed from diseased parts,'' and he used this concept to
explain how somatic modifications could be inherited.
Democritos held practically the same views, but Aristotle
attacked the doctrine and as a result the classical authori-
ties were divided into two distinct schools. The Galenic
literature supported Hippocrates, as did some of the early
Christian fathers, although others were undecided. It
may be noted parenthetically that patristic literature dealt
with the nature and origin of semen in some detail because
of certain theological problems concerning the inheritance
of original sin and the traduction of the soul of the child
from the souls of its parents.

St. Augustine (345-430 A.D.) almost certainly believed
in pangenesis, for he described the transfusion (not trans-
mission) of the flesh of the father into that of the son
by way of the semen ;^ and two hundred years later, St.
Isidore of Seville (622-633 A.D.) stated that semen came
from a mixture of the food and the body, a statement which
reminds us of Buff on.

During the thirteenth century several philosophers and
theologians discussed the mechanism of heredity and the
origin of semen. Roger Bacon (1268) explained the
progressive shortening of human life by assuming the in-
heritance of acquired characters, while Bartholomew the
Englishman (1240), St. Albert the Great (1254-1256) and
St. Thomas Aquinas (1256) indorsed pangenesis. St.
Thomas's contribution was a 4,000-word article in liis
commentary on the sentences of Peter the Lombard. In
his *'Summa Theologica" (II; Quaest. 81; Art. 1) he de-
scribed the inheritance of acquired characters as fol-
lows :

iFroin "De Genese ad litterarum," Bk. 10, Ch. 1: "Orderliness now
seems to demand that we discourse regarding the sin of the first man; but
since the Scripture relates about the flesh of the first woman, how it was
made, but is silent regarding her soul — it has made us much more intent on
inquiring rather diligently as to how those men can or cannot be deceived
who believe that just as the flesh is from the flesh, the soul is made from the
soul of man by means of the transfusion of the seeds (seminibus) of both
these things from the parents into the children."

i

, . . thus a leper may beget a leper, or a gouty man may be the father
of a gouty son, on account of some seminal corruption, although this cor-
ruption is not leprosy or gout. . . .

But all these explanations are insufficient. Because, granted that some
bodily defects are transmitted by way of origin from parent to child, and
granted that even some defects of the soul are transmitted in consequence, on
account of a defect in the bodily habit, as in the case of idiots begetting
idiots; nevertheless the fact of having a defect, by way of origin seems to
exclude the notion of guilt, which is essentially something voluntary.

A late thirteenth century Provengal MS. (D. II, 11; in
the Basle University Library) contains two drawings, one
of the male, the other of the female generative system,
which show the vascular ducts through which the semen
was supposedly collected from all the parts of the body.
Sudhof, who discovered this MS., printed these illustra-
tions in his ''Geschichte der Anatomic im Mittelalter'^
(Leipsi^, 1909), and they were copied by Choulant in his
''History and Bibliogrp.phy of Anatomic Illustration '^
(Chicago, 1920). They illustrate a conception of ''the
seminal veins'^ as old as Hippocrates.

Needham (1934) has recently called the attention of the
biologists to a passage in Dante's "Purgatoria^' (1318
A.D.), which derives the semen from the excess blood
which had acquired its "formative virtue '^ in the heart.
There are many points of interest in this passage (Canto
XXV), and it is possible that Dante really believed in a
form of pangenesis, although on the surface his der-
ivation of semen from the perfect blood of the heart
seems to be inconsistent with even a most liberal inter-
pretation of pangenesis. Three hundred years later,
however. Sir Kenelm Digby (1645) stated that the circula-
tory blood, by coming into contact with all parts of the
body, brought back to the heart all the "specific virtues''
of the several parts, which were thus made available for
the semen. Dante, himself, did not tell how the heart
received its "informing" power from all the parts of the
body and we can not tell definitely what his ideas were on
the subject.

Pierre Bercheur (Petrus Bechorius d. 1362), the noted

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ACQUIRED CHARACTERS

533

theologian, traced the semen no further than to the loins
in his well-known work, ^'Reductorum Moralis'' (Bk. 10;
ch. 1), but in his account of the famous eel-viper hybridiza-
tion^ (Bk. 9; ch. 70) he described the inheritance of a
specific somatic modification. Thus :

According to lorath and Ambrosius the eel never conceives from its
own kind, but from a serpent, which it arouses and leads on by a hissing.
Even though the serpent is poisonous in itself, nevertheless, it belches forth
its poison before a union of this Mud, so that the eel conceives a non-poison-
ous offspring,^ in nowise similar to the serpent but like unto itself. But when
the union has been completed the serpent returns to his poison again. The
eel is the Blessed Virgin who does not conceive from her own kind, that i3
to say from man, but rather from God himself and the Holy Ghost, who then
because of his venom of justice and severity could be called the serpent; but,
to be sure, in this blessed union the serpent (i.e. God) has put aside his
venom (i.e. all justice and severity) and so the eel (i.e. the Blessed Virgin)
in nowise conceived a venomous or wicked offspring but rather a very kind
and gentle one. Moreover, this serpent (i.e. God the Father) will on the
day of Judgement take up the venom of his severity again, when he shall
hand over the sinneri to the eternal fires, so that He who, in overlooking
men 's vices, now seems to lack the venom of severity, shall at that time appear
poisonous to them when he punishes their vices.

The fifteenth century physicians followed Hippocrates
and Galen rather than Aristotle and consequently the
medical incunabula contain a number of descriptions of
pangenesis. Two examples are cited. The first is in
the work of Antonius Zeno (Policola) who published **De
natura humana,'' Venetiis, 1491, in which he discussed the
nature and origin of semen in detail (pp. 30-36). He
described pangenesis as follows :

There are four factors of the sperm: — the matter-factor, the form-factor,
the production-factor, and the end-factor. The matter-factor is the fluid of

2 There are many descriptions of this cross. The story probably arose
through a misreading of a passage in Aristotle (Hist. An. 5: 540b). The
writer has recently reprinted the accounts written by Pliny, Oppian, Athenaos,
Aelian, Horapollo Nilous, St. Basil, Achilles Tatios, Philes and Alciati
(Zirkle, 1935b). There are many other records, Michael Glycas (c. 1118),
Konrad of Megenberg (1350), Raphael Maffejus (1506), "Ortus Sanitatis'*
(1517), etc.

3 This is in sharp contrast to the statement of Athenaos ("Deipnos. " Bk.
8, ch. 12) who, quoting Andreas, wrote, "Only those lampreys have a fatal
bite which come from a viper. ' '

4!a

blood which is brought down through the veins to the testes from the whole
body and from every member in the manner of an exudation, especially from
the principle members (particularly from the brain— according to Hippoc-
rates), and though it is not entirely transformed, yet it is suitable to being
transformed.

The second example is furnished by Johannes Peyligk
(Peylick) who published ^'Philosophiae naturalis com-
pendium, ' ' Leipsig, 1499. Peyligk denied the possibility
of preformationism and described pangenesis as follows
(p. 29) :

The material portion is the matter of the semen which has flowed down
from all the members of the parent himself, from whom are produced the
spermatic members. The formative portion is the productive spirit which is
formed from three principle spirits ; namely, the vital, the natural and the
animal: and from other special spirits which are separated in the individual
members. And this receives its final form in the testes as the sperm.

Louis Bonaccioli, a French physician, who flourished in
the beginning of the sixteenth century, wrote ^^Enneas
muliebris,'' which probably was first published in 1502.
In the passage to be quoted, he does not state that the
semen comes from all the parts of the body, but he enum-
erates so many that there is little doubt that he at least
believed in a modified pangenesis. From chapter 3 :

Obviously (inasmuch as it [the semen] would have to do with food) it
starts out chiefly from that part which is the source of food (such as the
liver), and especially from its vena cava (which, to be sure obtains a far
greater supply of this matter) ; if for any reason it [the matter] should be
lacking in this [part], it is drawn from the other parts which lead from
it and tardiness of emission is an indication [of this]. From the heart [the
matter comes] through the arteries; from the liver, through the veins; from
the brain (although more copiously) through the veins and arteries; but
above all from the region of the eyes, which is the most seminal of all the
places of the head ; you may learn this fact when through venereal union they
(the eyes) are changed in exact measure and when through immoderate use
of the sexual member the eyes evidently grow weak and feeble, when the
strength, of the other parts are not yet failing. From all these [parts], I say,
it is at length carried down to the seminal passages and to the testicles in
which it receives that divine power of generation. For the vital spirit of be-
getting, coming down from the heart, becomes fructifying by virtue of the
testes, from which it slips down in men to the tube of the penis, in women to
the folds of the vulva which have been explained above.

534

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ACQUIRED CHARACTERS

535

Raphael Maffejus (Volaterranus) published his **Com-
mentiorum urbanorum'' in 1506. He described pangene-
sis in book twenty-four. From p. 746 (Ed. of 1552) :

Semen, which the Greeks call "gouem" as tho author Aristotle says, is
sent forth among men in a more copious proportion than among the other
animals. Hence after coitus a weakness sets in. Moreover, it [the semen]
comes forth from the whole body. The similarity of children who resemble
their parents in even small markings of the body is proof. In woman, to be
sure, as it pleases certain men to state, no semen is given forth, but in its
place there is a monthly flow of blood ; or if semen is given forth, it is scant
(parvum) and is not necessary for conception.

Both Rhodiginus (1516) and Paracelsus (c. 1526) en-
dorsed pangenesis. Jacques Du Bois (Jacobus Sylvius,
1478-1555) also described it in his *^De hominis genera-
tione, sive foecunditatis et sterilitatis causis'^ (1530) as
follows :

Moreover, these differences of semen in quality and quantity, depend upon
the organization (or other conditions) of either the whole body or of the
chief parts, especially the testes. Having first treated of this [the semen]
and of the menstrual blood, we have abundantly demonstrated that the semen
is derived from the individual parts, especially from the principle ones.
Indeed inasmuch as the testes strain the semen and provide it with its final
form, they also must needs affect it according to their own organization.

Nicolas de LaRoche published **De morbis mulierum
curandis ' ' in 1542. He describes pangenesis in the female.
The female testes are of course ovaries. From chapter 1,
section headed ** Testes foeminarum'';

For the vasa spermatica attract the matter of the female semen from all
parts of the female body and then the testes receive [it] and after it has
been received and prepared (cooked), they transmit it to the parastatae
which are vessels embracing the womb (matrix) which as they recede from
the testicles, start to become a little wider until they enter the womb
(matrix).

Thomas Vicary, chief surgeon of St. Bartholomew's
Hospital, was surgeon to Henry VIII, Edward VI, Queen
Mary and Queen Elizabeth. He published **The Anatomic
of the Bodie of Man'' in 1548. Pangenesis is described in
the ninth chapter entitled **The Anatomic of the Handles
and their Parts."

*'i i f-

: the which seede of generation commeth from al partes of the body,
both of the man and woman, with consent and wyl of al members, and is shed
into the place of conceyving, where, through the vertue of Nature it is gath-
ered together in the selles of the matrix of the mother, in whom— by the
way of the working of the mans seede, and by the way of suffering of womans
seede mixte together, so that eehe of them worketli in other, and suffereth in
other — there is ingendered Embreon.

Jerome Cardan described pangenesis in 1550, and four
years later Jakob Rueff (Ruff us) discussed the origin of
semen in some detail. He concluded that it was produced
in all the bodily parts and that acquired characters were
inherited. From **De conceptu & genera tione Hominis,"
Tiguri, 1554, Chapter 1 :

Wherefore it remains to be recognized in the matter of the begetting of
humans as in the case of the origin of plants, that since we observe bodies
different in respect to their members being produced from one [and the same]
semen, we also believe that it [the semen] arises from different parts of the
body. Whence have they seen that which they expound? — they who claim that
the genital semen is produced only from the brain; since this is less in
agreement both with the ratio of mixtures [concoctionum rationi — lit. ratio,
reason, manner of digestions] and with the constitution of the body. Indeed,
it is certain, that some (and not a small portion) of it is derived from the
brain, but the chief portion is gathered from the most important parts of the
whole body. For, if we should say that the semen is produced in only one
or another part [of the body], anyone will see that this follows by correct
reasoning — [namely] that only those same parts should be reproduced. And
80, we can rightly say, that in addition to what originates in the brain the
semen is produced from the whole body and from all the most important parts
thereof; indeed, its effect instructs us as to its cause (origin), especially since
in the offspring we see the distinct members perfectly completed to the exact
form of the body. Also, against the opinions of others, we have on our side
Hippocrates himself, easily the greatest of all physicians; who himself as-
serted that the seed was gathered from the whole body, and so I say that what
is begotten corresponds to the constitution of the begetter — a weak man
being born from weak semen and a strong man from strong seed. In addi-
tion to these arguments comes the fact that we often observe in children
those diseases or defective marks of the body which are present in their
parents — things which we entirely believe to have passed into them [the chil-
dren] through a defect of the seed (genitura). And so, having certainly
established these facts regarding the origin and material constitution of the
genital semen, these things suffice as a preface.

Pierre Belon used pangenesis to explain the generation
of birds in his *'L 'Historic de la natur des oyseaux."
Paris, 1555.

536

THE AMERICAN NATURALIST [Vol. LXX

Just as seeds produce such plants as those from which they have been
gathered, so animals starting their growth f ) om the seed of their sex, become
at length like to those from which they have originated. The seeds are
excrements of the bodies, which have the potentialities of those substances
from which they have come, and which proceed from the last digestion of
the body's food. . . . But the seed of the female being an excrement also,
has as a property all the parts of the body, which are engendered from it —
not in present action but merely in matter and potienally does it have those
parts whereby nature has made the female to be different from the male
and hence it happens that sometimes deformed animals engender deformed
offspring — at one time, male and another time female.

Levinus Lemnius, a Dutch physician, explained the in-
heritance of disease by pangenesis, and Ambrose Pare
(Ambrosius Paraeus, 1517-1590), although his identifica-
tion of fertile and sterile semen were directly opposed to
those of Lemnius, described pangenesis and the inheri-
tance of disease just as Lemnius did. From the preface to
his **De hominis generatione^' (1573) :

. . . Good semen ought to be white, shiny, sticky, globular with the odor
of the elder or the palm-tree, liked by bees, and in water it sinks to the bottom
(literally — sinkable to the bottom of water) ; for that which floats, is con-
sidered unfruitful.4 Indeed, a great portion of it (comes) from the brain,
but as a whole it is derived from the entire body and from its individual
particles, both solid and soft. For, unless in its entirety, it flowed down from
the whole body, all the individual parts of the foetus would hardly be formed
therefrom; since, indeed, like things are made from like. From this it fol-
lows that the foetus resembles its parents not only in form and appearance;
but also in the formation of the limbs and in the inter-connection of the
internal parts-; to such an extent that diseases become hereditary when a
weakness of this or that organ (e.-viscus) is transferred to the offspring.

Yet there are those who think that this derivation of semen from the entire
body ought not to be understood fas consisting of] bulk and material since
the part of all the blood which is weakened is only a certain portion; but
rather as consisting of energy and the formative power ; to be sure only the
natural and vital animal spirits from the individual parts are the artizans
of formation and of life ; and these and the formative force flow down into the
semen which is produced by the testicles. (They say) that their argument
is the fact that several foetuses whole and complete in the number of all
their parts are produced from maimed parents.

4 Many philosophers believed the exact opposite of this. Hippocrates
thought that semen was fertile only when it was foamy. Aristotle held that
semen was discharged by means of a blast of air, and that the air gave it
fertility. Lemnius stated that coition was a churning of air into the semen.
Air itself was supposedly sufficient to fertilize the females of a number of
species.

No. 731]

ACQUIRED CHARACTERS

537

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Girolamo Mercuriale (1530-1606) published **De mor-
bis muliebribus ' ' in 1582. He quotes and endorses Hip-
pocrates on the origin of semen in the section headed
^'Signa sterilitatis."

If by a defect of the semen, then, there is no other means of knowing,
except by a knowledge of the constitution of the entire body : because it is
the opinion of Hippocrates, in the book on air, water and places, likewise in
his book on generation and elsewhere that: "The Semen separates from the
entire body''; and so, as the body is constituted, so is the semen; thus, by
whatever symptoms an abnormality of the body shall be recognized, by those
very same symptoms shall abnormality and defect of the sperm be dis-
covered. . . .

Albertini Bottoni held that semen was an excretion, but
admitted that it had its uses. Li ' ' De morbis muliebribus ' '
(1585) he stated that it was apt to produce structures
similar to those in its place of origin. From Ch. 41 :

For although the semen is an excretory fluid, it is nevertheless a useful one
and in addition it is fruitful by its nature, and copiously filled with the
spirit of life and when taken up in a suitable matrix, it is apt to produce
(something) similar to that from which it had been derived. The menstrual
blood likewise, although it is held to be the excretion of the whole mass of
blood nevertheless exerts no influence on the whole body but customarily brings
on a vexation in non-pregnant and non-lactating women.

Mauricius Cordae in a somewhat obscure passage in his
*'Commentarii in librum priorem Hippocratis Coi de
mulieribus'' (1585) supports pangenesis. From com-
mentary number 6 :

When this happens, it is now rightly asked whether the semen ought to be
considered as derived from every part or whether it (arises) rather from a
perfusion of spirit and heat. Just as the chief parts are all joined to one an-
other by communicating vessels and all the veins, nerves and arteries are con-
tinuous down to the testes and even penetrate and permeate them ; so there is
a meeting of spirits of every kind especially from these sources, together with
heat and there is a coming together not without a violent impulse and a spasm
as of heat ( ?), in this manner from the remaining parts, which seize and carry
with them an image of some one of them although not without other things.

Archangelo Piccolhomini described pangenesis in *' Ana-
tomicae praelectiones explicantes, ' ' Rome, 1586. The
following quotation was incorporated in a footnote in

538

THE AMERICAN NATURALIST [Vol. LXX

No. 731]

ACQUIRED CHARACTERS

539

Caspar Bauhin's ' ' Tlieatriim anatomicum, ' ' Francofurti,
1635, (p. 567):

The next matter of the semen goes forth from all the parts of the body and
especially from the principle viscera: it is nothing else but food which has
overflowed from the optimal nutrition of the parts; for nature prefers that
there be too much nourishment rather than too little. This [nourishment] is
then changed by the parts and when abandoned by assimilation [i. e. — when
the process of assimilation has finished working on it] it has received the
strength [power] of the parts. Wherefore, the nourishment which overflows
from the nutrition and assimilation of the brain is the nearest (proxima)
matter from which will bo developed that portion of the fecund semen, from
which the brain of the foetus itself will be produced. Now, when the nourish-
ment overflowing in the brain cannot reach the testes because of the cutting
of those veins, then they [the testes] become unfruitful. In the same manner,
men will become sterile if the vein which leads from the heart or from the
liver is cut; since it is especially the overflowing nourishment, or seminal mate-
rial of these parts, which is the cause of the semen becoming fertile.

Seventeenth century records of pangenesis and of the
inheritance of acquired characters are doubtless quite
numerous and new ones will probably be discovered by
any one who examines the medical literature of the time.
The writer (1935a) quoted two such records (Venette,
1687, and Ray, 1691) and Needham (1934) has called at-
tention to the fact that both Sir Kenelm Digby (1645) and
Nathaniel Ilighmore (1651) explained inheritance by
means of pangenesis.

Sir Kenelm Digby 's contribution appeared in his '*Im-
mortality of Reasonable Souls.'' He did not believe that
the semen itself came from all parts of the body, and he
stated the hypothesis in this form only to refute it. From
Ch. 23 ; p. 265 :

To deduce this from its origine, we may remember how our Masters tell us,
that when any living creature is passed the heat of its augmentation or grow-
ing; the superfluous nourishment settleth itself in some appointed place of
the body to serve for the production of some other. Now it is evident that
this superfluity cometh from all parts of the body, and may be said to con-
tain in it after some sort the perfection of whole living creature. Be it how
it will, it is manifest that the living creature is made of this superfluous
moysture of the parent : which, according to the opinion of some, being com-
pounded of severall parts derived from the several limbs of the parent;
those parts when they come to be fermented in convenient heat and moisture,
do tJiko their ]K)sturt' and sitn.-ition, according to the ]K)sturo and disiM)sition

■* -^

/

of parts that the living creature had from whence they issued : and then
they growing daily greater and solider, (the effects of moysture and of heat;)
do at the length become such a creature as that was, from whence they had
their origine.

Which an accident that I remember, seenieth much to confirm. It was of a
cat that had its tail cut off when it was very young: which cat happening
afterwards to have young ones, half the kitlings proved without tails, and
the other half had them in an ordinary manner;''^ as if nature could supply
but one partner 's side, not on both. And another particular that I saw when
I was at Algiers, maketh to this purpose, which was of a woman that having
two thumbs upon the left hand ; f oure daughters that she had did all resemble
her in the same accident, and so did a little child, a girl of her eldest daugh-
ters; but none of her sonnes. Whiles I was there I had a particular curiosity
to see them all; and though it be not easily permitted unto Christians to
speak familiarly with Mohametan women; yet the conditions I was in there,
and the civility of the Bassha, gave me the opportunity of full view and dis-
course with them: and the old woman told me, that her mother and grand-
mother had been in the same manner. But for them it resteth upon her
credit : the others I saw myself.

The arguments used by Digby to prove that the semen
itself did not come from the entire body are not important
in themselves. He really accepted pangenesis, however,
and he looked upon the blood as the vehicle for the collec-
tion and transmission of the specific formative virtue.

But we should be to blame to leave our Reader without clearing that diffi-
culty, which cannot choose but have sprung up in his thoughts, by occasion
of the relations we made at the entrance into this point concerning the cat
whose kitlings were half with tails, and half without : and the womans daugh-
ters at Algiers, that had as well as their mother excrescences upon their left
thumbs, imitating another lesser thumb : and the like effects whensoever they
happen, which they do frequently enough.

5 The kittens appeared in the fundamental Mendelian one-to-one ratio.
Mendelian segregation had already been described a number of times under
the heading "degeneration" (Zirkle, 1935 b), but no previous recording
of ratios has yet been found. Six years later Highmore (1651) also described
a one-to-one ratio, this time in puppies. These meaningless observations
preceded by just over two hundred years the first really authentic record of a
Mendelian ratio which has come to light. Whiting (1935) has pointed out
that Dzierzon (1854) in the * * Bienenf reund aus Schlesien" described a one-
to-one ratio in bees, fourteen years before Mendel published. The excerpt
from Dzierzon : * * One must be completely certain that the queen belongs to
the genuine species by birth. If she herself has come from the hybrid-brood
(Bastardbrut), then it is impossible for her to produce drones, unless she
produces half Italian and half German drones — but what is noteworthy —
this is not qualitative but quantitative ; as if it were hard for nature to com-
bine the two types into a middle species."

V

540

THE AMERICAN NATURALIST [Vol. LXX

Let him therefore remember, how we have determined that generation is
made of tlie bloud, which being dispersed into all parts of the body to irri-
gate every one of them ; and to convey fitting spirits into them from their
source or shop where they are forged ; so much of it as is superabundant to
the nourishing of these parts is sent back again to the heart to recover the
warmth and spirits it hath lost by so long a journey. By which perpetual
course of a continued circulation, it is evident that the bloud in running thus
through all the parts of the body must needs receive some particular concoc-
tion or impression from every one of them. And by consequence, if there be
any speciall virtue in one part which is not in another then the bloud return-
ing from thence must be endued with the virtue of that part. And the purest
part of this bloud, being extracted like a quintessence out of the whole
masse, is reserved in convenient receptacles or vessels till there be use of it:
and is the matter or seed, of which a new animall is to be made; in whom, will
appear the effect of all the specificall virtues drawn by the blood in its
iterated courses, by its circular motion, through all the severall parts of the
parents body.

Whence it followeth, that if any part be wanting in the body whereof
this seed is made, or be superabundant in it; whose virtue is not in the rest
of tlie body, or whose superabundance is not allaid by the rest of the body
the virtue of that part, cannot be in the bloud, or will be too strong in the
bloud, and by consequence, it can not be at all, or it will be too much in the
seed. And the effect proceeding from the seed, that is, the young animall
will come into the world savouring of that origine; unlesse the mothers seed
do supply or temper what the fathers was defective or superabundant in; or
contrariwise the fathers do correct the errours of the mothers. ...

... But to go on with our intended discourse. The seed, thus imbued with
the specificall virtues of all the severall parts of the parents body, meet-
ing in a fit receptacle the other parents seed; and being there duly con-
cocted, becometh first a heart: which heart in this tender beginning of a new
animal containeth the severall virtues of all the parts that aferwards will grow
out of It, and be in the future animal ; in the same manner as the heart of a
complete animal containeth in it the specifike virtues of all the severall parts
of Its own body, by reason of the blouds continuall resorting to it in a circle
from all parts of its body, and its being nourished by that juice to supply the
continual consumption which the extreme heat of it must needs continually
occasion in its own substance; whereby the heart becometh in a manner the
compendium or abridgement of the whole animal.

Nathaniel Highmore gave his conception of the mecha-
nism of heredity in his ''History of Generation/' Lon-
don, 1651. He disagreed with Digby in most details, yet
he advocated pangenesis. His conception of the forma-
tion of semen was essentially that of Buffon. He held
that our food consists of a great variety of elements be-
cause every part of the body requires a different type of

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No. 731]

ACQUIRED CHARACTERS

541

element for its repair and reconstruction. Highmore
comments upon the views of Digby as follows (from p.
29):

For the material part of this seed, there is a large dispute whether it be a
toto vel a parte decisum. I shall not stand to tell you the names of those who
are Patrons of the one, and of the other ; nor rehearse their arguments. If you
examine them, you shall finde theirs most rational, that aflfirm the decision
from the whole body; what we finde more particularly discours't of our
forementioned author, in his 24th chapter [Kenelm Digby-Discourse of Bod-
ies], we shall take up and a little review. Where he hath truly and fully
evicted the wand 'ring phancies of some, that would have this compound of
severall parts, to be collected from every particle, so as passing by or through
every little atome of the Parents body, in its passage ; should be impregnated,
and imbued with the nature of it, and so retire to the reserve where it is kept
for generation. And afterwards these particles being fermented by con-
venient heat, do take their posture and scituation ; according to the posture
and dispositions of those atonies they visited in their passage, and from whom
they received those imbibed natures. But this circulating our Author tells us,
is impossible. I will not wrong him so much as to rank his more solid reasons
with my own. Could we finde these chanels and conveighances in the Body,
by which this matter should pass; yet I might doubt of the unquestionable
verity of this doctrine. For what should hinder this matter circulating about
the Body, from receiving qualities, and so likewise the nature of every part
it passeth by; and so every particle of this matter, should be impregnated
with the natures of the whole; and every small Atome should become a living
Creature, or else the Subsequent should blot out the Antecedent Character and
the Impression should be only from the last Part. We may likewise as
truly, as safely conclude with our Author, that it is impossible for every little
part to remit some parts impregnated with the nature of that whole part
from which it fell. This by some is thought to be done by that Quasi Epilepsia
in Coitu, that kindo of convulsion or concussion of the parts, by which is
shook off from them somewhat retaining the nature, and property of every
part, and these being joyned make up the seed. This seems to be very much
befriended by our Authors relation of the cats kitned without tails; and of
the Womans daughters with six fingers upon a hand. Myself also have seen
a kinde of Poultry without rumps: which breeding with their own kind still
brought forth Chicken wanting that part: if with others, sometimes they
had rumps, sometimes but part of a rump. And not long since I saw a
Mungril Bitch, that had her tail cut close to her body almost, whose Whelps
were half without tails, and half with tails: the next year following, she
brought them forth all with long tails, as she had before the cutting off.
Which though it seems to favor (as I said) this opinion, it; as may appear
by the frequent perfect generations of mutilated creatures; which beget
children or issue with two legs or arms though they had but one ; Spaigniels,
whose tails are always cut, bring forth Whelps whose tails need as much
cutting, as their Dams or Sires did. Wee must therefore look out some other
way, how this may be done without the parts themselves.

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i

Aiitoine Legrand, the Augustinian missionary to Eng-
land, published ^^Institutio philosophica secundum prin-
cipici D. Renati Descartes'* in 1672 and in the next year
his ''Historia naturae." In both of these works he de-
scribed liow the semen collected from all parts of the
body and thus how mutilations could be inherited. From
''Institutio philosophica.'' (Pt. 8; Cli. 4; § 2) :

However, in order that it might be understood by what device an Animal
can bo sliaped from such a formless fluid so that it should have any resem-
blance to the element (principium) from wliich it was produced, it is neces-
sary to suppose that the Semen of the Male as well as that of the Female
flows down from all the parts of their bodies, so that there is no member
from which something of the seed of generation is not separated. For just
as a serous fluid is separated from tlie whole body through the Veins and is
stored through ' ' emulgentes " [probably— drain-ducts] in the Kidneys and
the Bladder, where at a given moment an ejection occurs; so why not, since
double Veins and a like number of Arteries go to the Testes — why should not
the Seminal parts flow into these (Veins and Arteries) from the whole body
and then from them into appointed vessels?

And this does not occur gradually and with any preparation of the
" materia, *' but it happens in a rather short space of time, in which the
whole body is so aroused to excretion, that whatever there is in the (several)
parts that is very life-giving (spirituosum) is moved, squeezed forth and
then hastens off.

The inheritance of mutilations is described in the fol-
lowing (^'Historia naturae," Pt. 8j Art. 2; Ch. 2) :

And it is this Semen which the Animal uses for the procreation of those like
unto himself. But if the animal from which the Materia Spermatica is pro-
duced, happens to lack any member (or to be overburdened by the super-
fluity of any part) then the virtue of that member will not be in the blood, and
therefore the offspring will be born maimed and lacking some part. If, in
truth, parts (of the body) of the animal are multiplied as Dighaeus mentions
in his African Women, or, if a superfluity accreses, it will effect a deformity
in the offspring also ; unless this superfluity is balanced in the semen of the
other parent, or the over-abundance of the blood (i.e. the over-abundance in
the blood of the virtue of the excrescent part) is corrected.

Legrand showed that the evidence used by certain
philosophers to prove that the semen was produced in
the brain could be given another interpretation.

Excessive effusion of Semen is most dangerous to the Brain and the Nerves
and produces a noticeable weakness in these parts; as (may be seen in) those

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543

M.

who indulge too much in Venery or befoul themselves with filthy contamina-
tions. Hippocrates once thought, and it is even now believed by many
physicans that the genital fluid flows off into vasa spermatica from the brain,
and that when an emptying of this fluid occurs, the strength of the brain, is
necessarily weakened. Indeed, since there are no special ducts from the brain
which might bring the afore-said fluid down to the spermatic parts, it is
reasonable to admit that this noble fluid is produced from the Massa
Sanguinaria, some part of which is sent off to the genitalia no less than to the
brain itself. As for the fact that excessive loss of genital fluid has a de-
structive effect upon the brain and brings about a weakness of that (organ),
that arises from the fact that the Blood restores what is lost through the
effusion of the semen that has been released, and supplies the brain with a
thinner portion of itself; since the greatest supply of the animal spirits is
[then] assigned to the spermatic parts. To be sure, since the Blood from its
own substance cannot lavish sufiicient upon the Genitals, it absorbs additional
matter from the brain and demands back what has previously been assigned
[to the brain] in order to recompense here.

Marcello Malpighi, who, together with Nehemiah Grew,
founded plant anatomy, described pangenesis in his essay
on the kidneys. The passage is in his ** Opera Omnia"
(London, 1687). From Pt. II, p. 289 :

But, it is probable, moreover, that it [the semen] is formed from the re-
tention of these particles, inasmuch as the substance proceeding either from
the whole [body] or from the blood is a deflux (decidua) of tiny particles of
semen ; wherefore the mass of blood, wheresoever driven from the structure
of the glands which are in the kidneys, is freed from the "exalted" salts
and impurities (filth) : the fermentation is kept up more easily, the first
beginnings [rudimenta] of similar and dissimilar parts are brought down and
separated in the work-shop of the testes and then when the semen is
regathered it is more fecund.

Martin Schurig wrote an excellent review of the vari-
ous theories concerning the origin of semen in his * * Sper-
matologia historico-medica" (Frankfort a. M., 1720).
His own views (Ch. 1 ; Part 11) are not definitely in favor
of pangenesis, although he conceded the probability that
the particles of semen are derived from the blood stream
and are filtered out and collected in the testes.

Accordingly, I believe that in tlie testicles, that is to say, in these vasculo-
glandular parts — because of their peculiar conformation, there occurs the
secretion or translocation of a peculiar fluid, different from all others (i.e. the
seminal fluid). On the other hand, I am quite willing to concede that such
spermatic particles just like others, e.g. the bilious particles, are hiding in
the blood and with the same, by means of the circulation (of the blood) are

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544 THE AMERICAN NATURALIST [Vol. LXX

brought through the arteries to the testicles, and then through the afore-men-
tioned peculiar vessels are separated into a fluid impregnated with the various
qualities of its kind, and finally it is conserved according to the manner
recounted by other Authors.

Conclusions
Belief in the inheritance of acquired characters can be
traced back to the time of Hippocrates. From the fourth
century before Christ until the nineteenth century it was
accepted as a matter of course; indeed, Brock noted but
two individuals during this period who definitely did
not believe that such characters were heritable, although
there were doubtless others. On the other hand, there
are many who described the inheritance of such charac-
ters. Among the latter are Hippocrates (c. 400 b c )
Aristotle (384-321 b. c), Galen (?) (130-220 a. d.), St!
Thomas Aquinas (1256), Roger Bacon (1268), Pierre
Bercheur (d. 1362), Sylvius (cit. Osborne), Jerome Car-
dan (1550), Jakob Rueff (1554), Levinus Lemnius (1561)
Ambrose Pare (1573), Sir Kenelm Digby (1645), Na-
thaniel Highmore (1651), Antoine Legrand (1672), John
Kay (1691), Michael Adanson (1763), Erasmus Darwin
(1794) and Johann Friedrich Blumenbach (1795). La-
marck's contribution to biological theory was to assume
that the inheritance of somatic modifications was cumu-
lative from generation to generation so that ultimately
new species were formed from the older ones.

Belief in pangenesis can also be traced to the fourth
century before Christ. The classical and medieval phi-
losophers, however, were not unanimous in support of
the doctrine, probably because of Aristotle's opposition
In many cases it is very difficult for us to determine their
attitude toward the hypothesis, for many who traced the
semen to some particular organ, such as the cerebrum,
liver, heart, kidneys, testes, etc., need not have been op-
posed to pangenesis. Often those who supported the
view that the semen was derived from all parts of the
body traced it to these same organs, where, they thought,
it was only stored temporarily. Among those who defi-

No. 731]

ACQUIRED CHARACTERS

545

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nitely supported pangenesis are Hippocrates (c. 400 b. c),
Democritos (c. 400 b. c), Galen (?) (130-200 a. d.), Clem-
ent of Alexandria (193-211), St. Isidore of Seville (622-
633), Bartholomew, the Englishman (1230-1240), St. Al-
bert the Great (1254-1256), St. Thomas Aquinas (1256),
Peter of Crescentius (?)(1305), Antonius Zeno (1491),
Johannes Peyligk (1499), Louis Bonaccioli (1502),
Raphael Maffejus (1506), Ludovicus Rhodiginus (1516),
Paracelsus (c. 1526), Jacques DuBois (1530), Nicolas de
La Roche (1542), Thomas Vicary (1548), Jerome Cardan
(1550), Jakob Rueff (1554), Pierre Belon (1555), Le-
vinus Lemnius (1561), Ambrose Pare (1573), Girolamo
Mercurale (1582), Mauricius Cordae (1585), Albertinus
Bottoni (1585), Archangelo Piccolohomini (1586), Sir
Kenelm Digby (1645), Nathaniel Highmore (1651), An-
toine Legrand (1672), Nicolas Venette (1683), IMarcello
Malpighi (1687), John Ray (1691), de Buff on (1750), de
Maupertius (1756), Albrect von Haller (1781), Charles
Bonnet (1781) and Herbert Spencer (1864). These names
probably represent but a small fraction of the total.

The writer is greatly indebted to Mr. David Goodman,
who translated the many Latin passages quoted.

literature cited

St. Augustine

1894. "De Genese ad litterarum, " Lipsiae.
Belon, Pierre

1555. * ' L 'Historic de la nature des Oyseaux, ' * Paris.
Bercheur, Pierre

1575. "Reductorii moralis, " Venetus.
Du Bois, Jacques

1530. *'De hominis gcneratione, sive foecunditatis et sterilitatis causis."
Geneve.
Bonaccioli, Louis

1502 (?).*' Enneas muliebris, ' ' Fcrrara.
Bottoni, Albertini

1585. "De morbis muliebribus, " Patavii.
Choulant, Ludwig

1920. "History and Bibliography of Anatomic Illustration/' Chicago.
Cordae, Mauricius

1585. " Commentarii in librum priorem Hippocratis Coi de mulidribus,"
Paris.

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THE AMEBIC AN NATURALIST [Vol. LXX

Dante Alighieri

1880. " Purgatorio, " London.
Dzierzon, Johannes

1854. " Bienenf round aus Schlesien, " Brieg.
Digby, Sir Kenelm

1645. "Immortality of Reasonable Souls," London.
Highmore, Nathaniel

1651. "The History of Generation," London.
LaRoche, Nicolas de

1542. * ' De morbis mulierum curandis, ' ' Paris.
Le Grand, Antonii

1680. "Historia naturae, variis experimentis et ratiociniis elucidata,"
ed. 2, Londini.

1683. * ' Institutio Philosophiae secundum Principia D. Renati Deeartes, ' '
Nurimbergae.
Maffejus, Raphael (Volaterranus)

1552. "Commentiorum urbanorum," Lugdun:.
Malpighi, Marcello

1686. "Opera omnia," Londini.
Mercuriale, Girolamo

1582. * * De morbis muliebribus, ' ' Venetus.
Needham, Joseph

1934. "A History of Embryology," Cambridge.
Pare, Ambrose

1573. "De hominis generatione, " Paris.
Piccolhomini, Archangelo

1586. "Anatomicao praelectiones explicantes, " Rome.
Rueff, Jakob

1554. ' ' De Conceptu et Generatione Hominis, ' ' Tiguri.
Schurig, M.

1720. " Spermatologia historia-medica, " Frankfurt.
Sudhof, Karl

1909. "Geschichte der Anatomie im Mittelalter, " Leipsig,
St. Thomas Aquinas

1912. ' ' Summa Theologica, * ' London.
Vicary, Thomas

1548. ' ' The Anatomie of the Bodie of Man, ' ' London.
Whiting, P. W.

1935. Jour. Hered., 26: 263-278.
Zeno, Antonuis (Policola)

1491. ' * De natura humana, ' ' Venetus.
Zirkle, Conway

1935. Amer. Nat., 69: 417-445.

1935. ' ' The Beginnings of Plant Hybridization, ' ' Philadelphia.

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