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Ohio State Academy of Science

Volume IV, Part 6

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_ THE WILLOWS OF OHIO

A MONOGRAPH

ROBERT F. GRIGGS

SPECIAL PAPERS, No. 11

ex Ck, DLNiGs

OF THE

Ohio State Academy of Science
Volume IV, Part 6

SPECIAL PAPERS, No. I

fale WILLOWS OF OHIO

A MONOGRAPH

BY
ROBERT, Fe. -GRIGGS

PUBLICATION COMMITTEE:
JOHN H. SCHAFFNER JAS. S. HINE BE EaRICE

Date of Publication, November 25th, 1905

PUBLISHED BY THE ACADEMY
COLUMBUS, OHIO

COLUMBUS:
SPAHR & GLENN, PRINTERS,

1905

NOTE.

The expense of publication of this monograph is covered by
a special grant from the Emerson McMillin Research Fund and
a small grant was previously made to assist in meeting the ex-
pense incurred during the progress of the investigation on which

the paper is based.
WILLIAM R. LAZENBY,

Joun H. SCHAFFNER,
C. J. HERRIcK,
Trustees.

INTRODUCTION.

The purpose of this paper is to make the willows of Ohio
knowable to persons of moderate skill in the determination of
plants. Perhaps it would be too much even to hope that it will
enable beginners in Botany to deal with the willows. But if it
helps those who have already some knowledge of the native flora
to extend their acquaintance to this very common genius, it will
have justified its preparation. For Salix, like Carex and Cra-
tegus is considered by many Botanists too difficult for any but
the specialist. Many competent workers seem to be unable to
cope with Salix and there are few even of the larger herbaria in
which the willows are correctly determined. The reason for this
the writer believes to be not in any inherent difficulty of the group
for it is not so difficult as many better known genera, but in the
fact that an adequate description of it has never been presented.
It is toward filling this need that the present effort is directed.

The species are all rather similar and variable but the dif-
ferences between them are not as inconstant as has been supposed.
But the space that has been devoted to them in the manuals is
entirely too small for their accommodation even when treated
by such a master as Bebb, whose account published many years
ago in Gray’s manual remains the best treatment of the species
within our area. Nor can their character be represented by line
drawings such as appear in Britton and Brown’s Illustrated
Flora; even the lithographs of Sargent’s Silva are but little better.
The character of a willow leaf is too subtle a thing for the ordinary
scientific artist to portray; for that, the camera is necessary.
Another fault of most of the descriptions and keys hitherto pub-
lished is that they have been written with a complete specimen
in mind, as it might be assembled on the herbarium sheet with
both kinds of flowers and leaves. But a collector never has a
complete specimen and is sure to be balked by lack of the mis-
sing parts. It must be admitted, to be sure, that there are some
stages of some species which are almost indeterminable. But
they are not so numerous as to make it inadvisable to construct
keys for the others.

Any successful treatment of the genus must for the present
be local in its scope for Salix is subject to very great geographical
variation and a treatment of a given species which would be en-
tirely accurate in a given area might be entirely inadequate for
the same species if observed a thousand miles from the first lo-
cality. The description of Salix nigra given below for example
would not cover at all satisfactorily the southern and western
plants which go by that name. And yet though the characters

258 OHIO STATE ACADEMY OF SCIENCE.

which we use in the descriptions vary enormously, one would still
recognize without much difficulty that the southern plant was
Salix nigra in a metamorphosed condition. Consequently, as
far as the native-species are concerned, at least, though it is hoped
that this paper will be of service all over the northeastern portion
of the United States, one must expect to find it more and more
at fault as one recedes from Columbus. Some day when there
is a monograph like the present covering every state in the coun-
try, some genius will write a general treatment that will effect-
ively handle all the species.

One’ reason why it is necessary at present to limit the treat-
ment to a small area is that no one has yet succeeded in describ-
ing the qualities by which we recognize a willow. We define the
species by external, artificial characters such as the shape and
hairness of the leaves, the length of the pedicels and so on; but

all such characters may vary enormously and still leave the qual-.-

ities by which the expert determines the species, constant. We
cannot study willows profitably by tables of dimensions and
geometrical descriptions of the leaves any more than we can
study the faces of our friends by measuring the length of their
noses. As there is a certain almost indefinable individuality in
a human face, there is in a species of plants an individuality no
less indefinable but no less important.

I here desire to extend my thanks to friends who have aided
me very greatly in the preparation of this paper: to Dr. W. A.
Kellerman of the Ohio State University, who at one time planned
to be joint author with me, for innumerable kindnesses done and
suggestions given during the whole course of the work; to Prof.
John H. Schaffner of the same institution for help especially in
verifying and testing the keys and to Mr. Otto E. Jennings of the
Carnegie Museum for the loan of the collections of that insti-
tution.

The present paper is based on studies and collections of wil-
lows begun in the spring of 1898 and carried on as opportunity
afforded ever since. During that time the author has been
enabled to study the plants in the field in various parts of Ohio,
in the vicinity of Washington, D. C., and in the Red River Valley
of North Dakota and Minnesota. The herbarium on which he has
mostly depended is that of the Ohio State University at Colum-
bus. The United States National Herbarium and the Herba-
rium of the Carnegie Museum at Pittsburg have also contributed
considerably to whatever value the work may possess.

TERMS.

Most of the terms used are those common to Botanical des-
criptions, but those referring to the venation may be so unfa-
miliar as to require definition. They are based on a most excel-
lent paper by Dr. Glatfelter (Rep. Mo. Bot. Gard 5:46-60. 1894.)
on the venation of the willows.

Primary. A main vein branching directly from the midrib.

Costal. A smaller vein from the midrib, of considerable
size but not reaching as far as the primaries.

Secondary. A vein of the second order given off from a pri-
mary.

Tertiary. A vein given off from a secondary.

Arch. The joining near the margin of a primary with a
fork sent off from the one next above, the two branches together
forming the arch which rests upon the two primaries.

Loop. Formed by a primary near the margin curving
forward and inward and merging into the next above, similar
to an arch but lacking the angle at the apex.

Regular. Veins parallel.

Bract. \ In his treatment of the genus in Britton’s Manual

Scale. Ie P. A. Rydberg applies the term bract to the
rudimentary perianth of the flower, a small leaf subtending the
essential organs, whereas previous writers have termed this a
scale and reserved the former term for the bracts which support
the aments. This departure from the accepted usage seems to
me almost as confusing and unjustifiable as the departures from
the rules of priority in nomenclature which the New York Botan-
ists condemn so strongly. Further the meaning given the terms
in the glossary at the end of the book, accords better with the
older usage than with Rydberg’s application of them. Those
definitions are here followed. According to Britton, a “Bract”’
is ‘A leaf, usually small, subtending a flower or a flower cluster

or a sporange.’’ That is, in this case one of the leaves which ap-
pear at the bases of the aments. And a “Scale” is “‘“A minute
rudimentary or vestigial leaf.’’-—In this case one of the small

leaves in the axils of which the flowers are borne. To make the
matter worse, Rydberg uses bract in both senses thus introducing
ambiguity also.

The Willows of Ohio.

THE GENUS SALIX AND ITS RELATIONSHIPS.

The willow family contains but two genera, Salix and
Populus. These, in most cases distinct enough in their leaves
are separated by the following floral differences. Populus has
fimbriate scales, cup shaped discs, elongated stigmas and many
stamens, while Salix has entire or only slightly crenate scales,
mostly glandular clavate discs, short stigmas and few stamens,
two in the majority of the species. In the polyandrous willows
which are treated first in the present paper, there is a distinct
approach from the diandrous or prevailing willow type toward
the poplars, most conspicuously shown in the increased number
of stamens but also evidenced in the disc which becomes com-
pound with several glands and sometimes almost forms a lop-
sided cup. It is, too, in the polyandrous willows and in those
diandrous species manifestly most closely related to them that
we find most of the arborescent species—a habit prevailing
among the poplars—while most of the willows are shrubs.

There are recognized in all something like two hundred
species of willows. They are mostly natives of the north tem-
perate zone but are not entirely absent from the torrid and south
temperate zones.

The willow may be counted one of the most successful of
present day plants. In few other genera, so compact and homo-
geneous in respect to their floral characters, is there shown so
great a variety of adaptations to varied conditions of life. With-
in this one genus may be found plants all the way from large
trees to dwarf herbs, affecting habitats from the fertile alluvial
plain to the barren mountain peak. They grow almost every-
where but yet there are certain limitations in their habitat.
Varying from hydrophytes to xerophytes, they are uniformly
lovers of the sun and never found to any extent in deep forests
or other shaded situations. In their various habitats their
vegetative structures undergo wide modifications to accommo-
date them to their environments. Some species like Salix lucida
have broad thick leaves, protected from too severe conditions by
the hard glossy surface.. In others like Salix adenophylla the
same protection is gained by a thick coating of wool on one or
both surfaces. Ora heavy coat of glaucescence may be devel-
oped probably to a degree at least for the same purpose. In
other cases the leaves are so small as to enable them to endure
the most severe conditions, especially when, as is often the case,

262 OHIO STATE ACADEMY OF SCIENCE.

there is coupled with the small surface exposed a prostrate habit
which shelters them from the wind and allows them any ad-
vantage to be gained from warmth radiated from.the ground.
In great contrast to these dwarf creeping almost herbaceous
forms are the tree willows so well known to every one. These,
by their rapid growth are enabled to succeed well in their
favorite habitat along streams and lakes because they can
quickly repair the damage done by floods and storms, while
more slowly growing trees would be almost hopelessly beaten
down and destroyed before they could rear their trunks to a
sufficient size to enable them to withstand the elements. Like-
wise their well known ability to grow from cuttings and broken
branches renders them able to use the very storms which break
them in pieces as the means of their further dispersal.

Their well known variability under different conditions is
an evidence of plasticity of constitution and adaptability to
various environments which is as important a factor in the success
of a plant in meeting the competition to which it is subjected as
is the same quality in the success of aman. From their adapta-
bility, variability and the large number of intermediates between
very divergent forms we may safely infer that we have here a
group of species with a very large amount of ‘“‘ Vital Motion” in
rapid course of evolution.

ON USING THE KEYS.

To construct a key which will hold for all individual willows
is well nigh impossible, because in the first place, there is a greater
or less number of hybrids for which no pretense at a key is made;
and secondly, there is so much variation in some of the species
that even after including them in two or three different places,
the writer is not so sanguine as to suppose that he has covered
all the variant forms. Bearing this probable defect of the key
in mind, it will be understood that the greater the diligence with
which each clue given is followed out, the greater will be the
probability of a correct determination. If the plant you have
seems to fit in both divisions of the key by all means run it
through both and after doing so refer to the descriptions and
plates as well.

The synoptic table is intended primarily to give some idea
of the relationships of the different species and consequently the
whole anatomy of the plant is used while all the parts are never
present in a single specimen. . But in the keys proper every-
thing is subordinated to the end in view, namely the identifica-
tion of the plant; and only such characters are used as are actually
present on a single specimen. In the foot-notes under the keys
their especial uses and limitations are discussed.

THE WILLOWS OF OHIO. 263

Synoptic TABLE OF OHIO WILLOWS.

Stamens 3 or more, filaments pubescent at base, catkins appear-
ing on leafy branches. Polyandrae.
Trees with rough, flaky, brown bark and brown twigs.

Amygdaleneae.
Capsules short pedicelled, short globose conic, forming dense
thin catkins; leaves green on both sides, glabrous, pri-
maries distant, very fine net veined, with a marginal
running nearly to the base of the leaf, linear-lanceolate
when mature, often falcate; of very scragly growth,
seldom upright; frequenting streams. S. nigra.
Capsules long pedicelled, short conic, in lax aments; leaves
sometimes 15 cm. long, typically oblong with straight
sides, glaucous beneath, generally hairy, primaries
ascending, but scarcely forming a marginal; sprawling

shrub or straggling tree; southern, river banks.
S. Longtpes.
Capsules long pedicelled, long conic, forming large loose
catkins; leaves glaucous beneath, glabrous, primaries
close, reticulation not so fine, marginal seldom running
below the middle of the blade, broadly lanceolate,
petiole often red; an elegant shapely tree; northern,
prefers swamps. S. amygdaloides.
Shrub, twigs shining brown, scales often dentate, capsules
large and glossy, aments thick; leaves ovate-lanceolate,
often very long attenuate, sub-coriaceous, glossy above,
light green, glands very prominent, especially on stipules;

mostly in rocky wet places, northern. S. lucida.
Stamens less than 3.
Stamens 2. Diandrae.

Filaments pubescent, catkins appearing with the leaves,
on leafy branches, except early flowers of S. interior,
scales nearly as long as the ovularies at anthesis.

Large trees with bark rough but not flaky, leaves glaucous

below, capsules glabrous, short pedicelled. Fragzles.

Not weeping, primaries and secondaries close and
regular.

Bark greenish, leaves glabrous, rather coarsely ser-
rate, and strongly glandular; capsules long
conic, remaining green. S. fragilts.

Bark yellowish, leaves pubescent at least when
young, fine serrate; capsules short conic, turning
yellow in fruit. S. alba.

Bark golden yellow, leaves glabrous in age
S. alba vitellina.

264 OHIO STATE ACADEMY OF SCIENCE.

Branches pendent, leaves small, very fine net veined,
primaries and secondaries irregular; capsules short
globose conic, remaining green in fruit, staminate
plants practically unknown. S. babylonica.

Shrub, often growing in dense clumps with many slender
stems; leaves often very long, linear-oblong, with a
strong marginal vein and distant primaries, smaller
veins except a few costals vanishing; catkins often
in cymose clusters which continue flowering all sum-
mer, or the earliest only leafy bracted. S. znterior.

Leaves short and broad (more than 1 cm.) inclined to
be very wooly; cymose clusters of catkins very pro-
nounced, carpellate plant rare. S. interior wheelert.

Filaments glabrous, catkins coming before or with the
leaves but not on leafy branches unless in fruit.

Capsules pubescent at least in flower.

Styles short and inconspicuous, less than half as long
as ovulary.

Leaves mostly broad, coarsely serrate or entire; cap-
sules long conic, very hairy; upright, many
stemmed shrubs preferring lowland swamps.

Capreae.

Catkins at anthesis seldom 2 cm. long, not very
wooly, bracts conspicuous, scales light brown,
persistent, pedicel very long, filaments slender;
venation prominent below, leaves dirty white
tomentose, especially on the veins. S. bebbiana.

Catkins seldom less than 2 cm. long, wooly pussies,
bracts small, scales almost black, filaments
thick, pedicel medium; leaves mostly glabres-
cent, veins not very prominent below.

S. discolor.
Leaves narrow.

Leaves undulate-revolute to entire, primaries prom-
inent on the under surface, distant, coming
out at right angles and arching or looping
regularly to the base of the leaf; aments small
short pussies without leafy bracts, capsules
long conic, very hairy; upland swamps and
hillsides. S. humilis.

As above but smaller throughout; leaves up to
5 cm. long, aments less than 1 cm. long,

shrub less than 1 m. tall; prairies especially.
S. humilis tristts.

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THE WILLOWS OF OHIO. 265

Leaves sharply serrate, showing a decided tendency
to blacken in drying, aments from pussies,
bracts small or none, capsules when long
conic, thinly pubescent; lowland shrubs.

Sericeae.

Capsules blunt, short pedicelled, short conic,
densely silvery silky; leaves dull above,
lustrous sericeate below, at least till old.

S. sericea.

Capsule long conic, long pedicelled, sparsely pu-
bescent; leaves shining above, glaucous be-
low, glabrous (or rusty sericeate when
young); northern. S. petiolarts.

Style very long, capsule short conic, silky, white; leaves
long and narrow, revolute with veins prominent
below and depressed above, not distinctly arching,
snowy tomentose below; wet prairie shrub not
more than a meter tall; northern. S. candida.

Capsules glabrous, styles short, filaments glabrous.

Filaments distinct to the base; leaves sharply serrate,
good sized shrubs. Cordatae.

Leaves dull on both sides, only a little paler beneath

Leaves thickly pubescent on both sides alike, short
and broad, bracts broad, remaining green; north-
ern. S. adenophylla.

Leaves thinly pubescent with most of the hair on
the veins beneath, or glabrous, green on both
sides or slightly glaucous beneath, lanceolate,
bracts narrow, remaining green. S. cordata.

Leaves glossy above, very glaucous below, glabrous,
generally broad, bracts narrow; turning black;
northern. S. glaucophylla

Filaments frequently united at the base, capsule long
pedicelled in fruit, leaves small, entire, purplish
green, conspicuously reticulated; low shrub in
sphagnum bogs. S. pedicellaris.

Stamen 1, formed by the coalescence of 2, anthers 4, scales
black, reflexed in staminate flower, capsules rarely seen,
short, thick, hairy ; leaves oblanceolate, opposite or scattered,
purplish; shrub with long slender branches.

Synandrae. S. purpurea.

266 OHIO STATE ACADEMY OF SCIENCE.

Key BaAseD ON CARPELLATE AMENTS.*

1. Flowers appearing in pussies before the leaves, bracts small
or absent. 2.
1. Aments and leaves unfolding together, bracts mostly
large, sulle
2. Capsules glabrous from a wooly rachis, inner membrane
of bud scale growing out beyond the outer. 3.
2. Capsules silky till after breaking open, bud scale not as
above. 9.
3. Capsules turning brown when ripe, 10 mm. long, aments
lax, bracts glabrous and glaucous in fruit.
S. glaucophylla.

3. Capsules remaining green, less than 10 mm. long, bracts
green on both sides. 4.

4. Bracts narrow, obscurely serrate or entire, aments be-

coming rather lax, leaves becoming glabrous above.

S. cordata.

4. Bracts broad, prominently glandular, aments dense,

‘leaves becoming densely tomentose above; north-

ern. S. adenophylla.

5. Style very long and slender, rose red at anthesis, conspicu-

ous, capsule snowy white; leaves tomentose; dwarf

shrub; northern. S. candida.
5. Style less than half as long as ovulary, leaves glabrous
above. 6.

6. Capsules short conic. 7.
6. Capsules long conic. 8.
7. Aments long, 35 mm. or more, very dense, often opposite,
capsules large, strictly sessile, carpellate plant rare.

S. purpurea.
7. Aments short, 30 mm. or less, moderately dense, not oppo-
site, capsules small, short pedicelled. S. sericea.

8. Capsules short pedicelled. 9.
8. Capsules long pedicelled; northern. 10.

*The fruiting aments of our willows are quite-distinctive and though
the differences between the species are not always easy to describe, they
are constant and easily learned, so that with fruiting specimens we should
have a minimum difficulty in determination. But in flower they are by no
means so easy to determine. The ovularies are all very much alike and
seem not to develop character until filled out. In fruit too, the leaves
of most specimens are present and help greatly.

The greatest need of caution in using the key is to be sure that the
descriptions of capsules are not applied to younger stages which are more
slender with shorter pedicels. It will also be found difficult to use the key
after the capsules have burst. The leaves referred to are those pres-
ent at flowering and fruiting time. They mav or mav not be similar to
the mature leaves of the species.

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THE WILLOWS OF OHIO. 267

Aments long in fruit with many capsules; ees greenish

white when fresh. Si; discolor.
Aments short and thick, few flowered, stigmas red when
fresh. S. humilis.

10. Aments short, scales frequently darkened at the tip,
leaves narrow, turning black. S. petiolarts.

10. Aments long, scales yellow, leaves broader, strongly
veined, remaining green. ' ~ S. bebbrana.

Capsules hairy. 12.
Capsules glabrous. 16. . :
12. Style more than 1 mm. long. S. candida.
12> style less than 1mm. long. ~ 13.
Capsule subsessile, blunt pointed, stigmas sessile. S. imtertor.
Capsule with a distinct pedicel, acute, narrow conic, stigmas
rarely sessile. 14.
14. Capsule thinly hairy, almost glabrescent, aments short,
leaves narrow, silvery, blackening; northern.
S. petiolaris.
14. Capsule persistently hairy, aments long. 15.
Scales persistent, light colored, leaves broad, wooly below
with prominent veins, pedicels very long; northern.
S. bebbiana.
Scales usually deciduous at length, almost black, leaves
without prominent veins, pedicel short to medium.
S. discolor.

- 16. Capsules long pedicelled. 17.

16. Capsules short pedicelled or sessile. 19.
Aments short, bracts obtusish, small, capsules generally
redening; small shrub in sphagnum bogs only.
S. pedicellaris.
Aments long, bracts large; good-sized shrubs or trees. 18.

18. Capsules narrow conic; northern. S. amygdaloides.
18. Capsules mostly globose conic; extreme south.
S. longipes.

Capsules short conic, aments nearly glabrous. 20.
Capsules rather long conic. 22.
20. Aments dense, short (25-mm. or less) of small calibre,
with a few small bracts. S. babylonica.
20. Aments longer, bracts larger. 21.
Scales as long as the ovulary, capsules yellowing eo fruit,
aments rather lax; leaves pubescent, glaucescent. S S.alba.
Scales shorter than the ovulary at anthesis, capsules green,
aments dense; leaves glabrous, green. S. nigra.
22. Bracts, especially stipules, very glandular, coriaceous,
bracts and leaves broad, green, capsules when ripe
more than 6 mm. long; northern. S. lucida.

268 OHIO STATE ACADEMY OF SCIENCE.

22. Bracts not especially glandular, stipules seldom present,
capsules less than 6 mm. long. 23.

23. Stigmas sessile, capsule blunt, flowers often fascicled on the

rhachis with a distinct interval between the fascicles,

bracts green; shrub in dense clumps. S. interior.

23. Stigmas on a short style, capsules pointed, flowers not fas-
cicled. 24.

24. Sprawling shrub. S. cordata.

24. Trees in clumps with few stems or single. 25.
25. Leaves green, reticulate venation very fine, bark brown,

undeveloped stages of S. nigra.

25. Leaves glaucescent, venation coarse, bark yellow or
green. 26.

26. Leaves glabrescent. S. fragilis.

26. Leaves pubescent, young stages of S. alba.

FIELD Kry BASED ON MaTuRE LEAVES AND Hapsits.*

Leaves alternate. 2.

1. Leaves opposite, oblanceolate, nearly sessile, stiff ascend-

ing, purplish green; shrub with long slender branches.
S. purpurea.

Secondary and tertiary veins prominent by transmitted

light, unless concealed by tomentum. 3.

2. Secondaries and tertiaries almost obliterated with age,
primaries distant, running into a strong marginal;
leaves narrowly oblong, serrate with distant spin-
ulose teeth or entire; often forming dense slender
stemmed clumps. S. interior.

3. Leaves persistently pubescent. 4.

3. Leaves glabrous unless very young. 18.

4. Pubescence not concealing the upper surface. 6.

4. Both surfaces hidden, at least till mature, by a thick
coat of tomentum; shrubs growing in exposed
places iano: .

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* Note. This key is constructed for mature well developed leaves.
It will not hold for leaves which have not acquired their full size or ven-
ation, nor for those of water-shoots and suckers. In immature leaves the
primaries are usually more ascending, the other veins not well developed;
and the leaf is commonly relatively broader and blunter than when mature.
Frequently the two or three terminal leaves of a twig do not assume the
character of the species so that one must be careful not to be misled
by them when they are different from those lower down on the twig. The
leaves of water-shoots are so characterless that it is often difficult for
an expert to determine them certainly. They are all thin, green on both
sides, lacking both the pubescence and glaucescence which may render the
normal leaves distinguishable.

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THE WILLOWS OF OHIO. 269

Leaves oblong, hair long and straight. S. interior.
Leaves ovate, hair shorter, matted. S. adenophylla.
6. Pubescence not lustrous. 7.

6. Leaves clothed below, sometimes only thinly, with sil-
very short hair having a lustrous appearance al-
most like changeable silk, glabrous above, narrow,
narrowed to both ends, sharply serrate, generally
blackening in drving, venation irregular; spraw-
ling shrub. S. sericea.

Leaves narrow, oblong or spatulate, broadest above the

middle, margins mostly revolute, venation very . prom-
inent below. 8.
Not as above, margins serrate (or rarely entire but not
revolute.) 10.
8. Hair dull red brown, leaves dull above, not revolute.
S. discolor.
8. Pubescence not ferruginous, leaves revolute. 9.
Secondaries depressed above, leaves very thickly snowy to-
mentose, especially on the veins, margins mostly entire;
dwarf prairie shrub, northern. S. candida.
Secondaries not depressed above, leaves gray tomentose,
margins mostly undulate; growing larger, in hillside
swamps and uplands. S. humilis.

10. Leaves coarsely serrate with relatively distant, blunt
teeth incurved except when young, or entire;
swamp shrubs with ascending stems. 11.

10. Leaves closely and sharply serrate, rarely entire, teeth
not incurved. 12.

Veins often depressed above, dirty white tomentose and

very prominent on the under surface; northern only.
S. bebbiana.
Veins not depressed above, not very prominent beneath
nor especially hairy, frequently ferruginous.
S. dtscolor.

12. Pubescent on both surfaces. 13.

12. Pubescent only below. 14.

Leaves ovate, subcoriaceous, very glandular, rarely entire.

tomentose with matted wool, green on both sides; shrub,

northern. S. adenophylla
Leaves lanceolate, thin, not especially glandular, pubescent
with parallel hairs, glaucous beneath; tree. S. alba.

14. Pubescence evenly distributed. 15.
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14. ubescense mostly on the veins, wooly. 16.
Venation regular, neither arching nor looping; arborescent.
S. alba.
Venation irregular, arching or looping; sprawling shrub.
S. sericea.

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27.

OHIO STATE ACADEMY OF SCIENCE.

16. Primaries ascending a long way near the margin, usu-
ally not arching; leaves narrow, often auriculate
in rank growth. S. longipes.

16. Primaries more distant, forming a series of arches or -
loops near the margin; not auriculate. 17.

Leaves ovate; northern. S. adenophylla.
Leaves lanceolate. S. cordata.

18. Leaves green on both sides, only a little paler if at all

beneatinnl9:

18. Leaves glaucous beneath. 27.

Leaves relatively thin, dull. 20.

Leaves subcoriaceous, glossy above, strongly glandular es-
pecially on the stipules, ovate or broadly lanceolate,
often, especially in rank growth, very long attenuate;
buds and twigs bright shining brown, buds large; a
beautiful shining shrub; northern. S. lucida.

20. Marginal vein prominent, extending nearly to the base
of the leaf, primaries distant. 21.

20. Marginal not present, primaries closer. 23.

Leaves very finely serrate, reticulate venation very fine,
long acuminate, frequently falcate; growing into a scrag-
ly tree, along streams. S. nigra.

Leaves spinulose serrate to entire, teeth rather distant,
meshes coarse, acute, not falcate; shrubs tending to
form close thickets. 22.

22. Leaves, except those at the bases of the season’s twigs,

less than 1 cm. wide. S. interior.

22. Leaves more than 1 cm. wide. S. interior wheelert

Leaves dark purple green, entire or obscurely serrate. 24.

Leaves bright green, mostly distinctly serrate. 25.

24. Leaves elliptical; dwarf shrub in sphagnum bogs only.

S. pedicellarts.

24. Leaves mostly oblanceolate; shrub with many ascend-

ing branches, often planted. S. purpurea.

Venation irregular, primaries mostly arching or looping;
shrubs. 26.

Venation regular, primaries ascending to near the margin;

arborescent; water-shoots of S. fragilis.

26. Leaves sharply serrate; shrub with branches lopping
over onto the ground. S. cordata.

26. Leaves distantly serrate with incurved teeth; shrub
with stems ascending. S. discolor.

Leaves spatulate, elliptical, or oblong-oblanceolate, not
sharply serrate. 28.
Leaves lanceolate or broader than above, serrate. 31.

30.

33.

(SY)
Or

37.
37.

THE WILLOWS OF OHIO. PTE:

28. Leaves distantly serrate with incurved teeth, not

revolute. S. discolor.

28. Leaves entire or very obscurely serrate, frequently
revolute: “29.

Margin undulate-revolute, veins very prominent below,
bright green, only rarely completely glabrous. S. humilis.
Margin not undulate, veins not raised below, purplish
green. 30.
30.- Leaves elliptical, widest near the middle; dwarf shrub
in bogs. S. pedicellarts.

30. Leaves oblanceolate, widest above the middle; good-
sized shrub; often planted. S. purpurea.

Leaves with a strong marginal vein, smaller veins forming
a very fine meshwork, leaves small, narrow, often fal-
cate; tree with pendent branches. S. babylonica.
Marginal if present not extending much below the middle

Glave eae a2:

32. Leaves bluntly and coarsely or irregularly serrate with
incurved teeth or entire; shrubs. 33.

32. Leaves sharply serrate with projecting teeth. 36.
Leaves extremely glaucous below, glossy above, broadly
lanceolate or ovate; northern. S. glaucophylla.
Leaves with a thin bloom beneath, not glossy. 34.
34. Leaves lanceolate, long acute, venation regular; bark
of branches green, smooth; arborescent. S. jragilis.
34. Leaves mostly broadest above the middle, short acute;
bark of branches brown, soon roughened. 35.
( S. glaucophylla.

Water shoots, blackening in drying 4 and
l S. discolor.
Well developed leaves, remaining green. S. discolor.

36. Leaves held up by the stiff midrib so as to display their
white under surfaces, broadly lanceolate or ovate,
mostly glossy above and very heavily glaucous be-

low; northern. S. glaucophylla.
06. Leaves not held up against the twigs, mostly lanceo-
late (3,

Primaries mostly forking and arching; sprawling shrubs. 38.

Primaries mostly ascending without forking, arching or

looping; arborescent, not sprawling except S. lon-
gipes. 39.

38. Leaves dull above, not very glaucous, almost always

with a trace of hair on the veins above or below,

quite sharply serrate, petiole stoutish. S. cordata.

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OHIO STATE ACADEMY OF SCIENCE.

38. Leaves subglossy, frequently very glaucous, when
young sometimes pubescent but with evenly dis-
tributed sericeate hairs, inclined to be distantly ser-
rate with prominent glands, petiole slender, leaves
hanging down gracefully from the twigs.

S. petiolaris.

39. Venation mostly regular, reticulations not very fine, no
marginal; trees with smooth bark on the branches. 40.

39. Venation mostly irregular, reticulations very fine, with a
strong tendency toward a marginal; bark of branches

ridgey, brown. 41.
40. Bark of branches yellow to olive; serrations very fine
and sharp, not especially glandular. S. alba.

40. Bark of branches green; serrations rather distant,
glands prominent, glauscescence absent from the
veins beneath leaving them contrasted with the
rest of the leaf. S. fragilis,

41. Northern and western parts of state only, shapely tree,
especially in swamps; leaves ovate-lanceolate, twigs
smooth. S. amygdaloides.

41, Extreme south only, straggling tree or sprawling shrub in
river beds; leaves oblong-lanceolate, twigs usually
pubescent. S. longipes.

AMYGDALENAE, BLACK WILLOWS.

Mostly trees with rough brown bark, leaves long attenuate,
often falcate, with a marginal vein at the tips at least, reticula-
tions of the secondaries and tertiaries very fine. Catkins un-
folding with the leaves, stamens 5 or more, filaments pubescent,
scales one colored, deciduous, capsules glabrous, green, style
short or none. Conspicuous inhabitants of the temperate re-
gions of America and extending clear through the tropics to Chili
and Peru.

For a key to this section of the genus I feel that I cannot do
better than copy the following table in which the characters of
the three species are contrasted, from an article by Dr. Glatfelter
in Science Nov. 1, 1895, which I have found quite useful. It is
constructed for the forms about St. Louis and is particularly
servicable on that account; for Missouri is the only region where
the ranges of all three species overlap and all can be studied
together.

S. migra

Range extended
north and south

large tree, branches
crooked, ascend-
ing

stems in clumps

old bark flaky

twigs very brittle at
base

hardy

shoots pubescent

leaves oblong or lin-
ear lanceolate

bases acute to trun-
cate

under surface green

venation very min-
ute, strong mar-
ginal

petiole short

stipules pointed,
persistent, non-
glandular

blossoms about Ap-
ril 25

stamens mostly less ©

than 6
scales short, obtuse
capsules ovoid coni-
cal
pedicel short, slender
notched stigmas and
style

THE WILLOWS OF OHIO.

S. longipes
south of 39° lat.

small spreading top

single
deeply laticed ridg’y

tenacious

buds winter-killed
hoary pubescent
same or broader

acute to auriculate
whitish glaucous

without marginal

short
obtuse, persistent,
non glandular

May 5
4—7 mostly 5-6

ovate
globose conical

long, stout
both poorly devel-
oped.

273

S. amygdaloides
north and west

large, branches
straight

single

smooth or roughish
somewhat brittle

hardy
glabrous
ovate-lanceolate

acute to cordate
bluish glaucous

coarser more regular

very long

obtuse, caducous, al-
ways glandular
April 15

6-9

ovate-oblong, acute
ovoid conical

long, stoutish
as in migra.

SALIX NIGRA Marsh. BLiack WILLOW.

A tree occasionally reaching a height of 40 meters and a

trunk diameter of 1 meter but generally dying when about 15
meters tall. It most often grows in characteristic straggling
clumps with four or five crooked, leaning stems. In the fall it
drops off most of the season’s twigs leaving the old branches bare,
a habit which hinders their elongation and in a few years makes
them thick and stubby. This excess of self-pruning together
with the straggling clumps in which it grows give it a habit which
n typical specimens clearly distinguishes it from any other

oe

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Plate I, SaLix NIGRA.

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THE WILLOWS OF OHIO.

species, especially in winter when it is leafless. In summer the
long slender twigs sometimes give it almost the appearance of the
Weeping Willow from which, however, it can be easily distin-
guished by its leaves green, not glaucous. The winter buds are
very small, less than 3 mm. long, broadly ovate, acute, commonly
but not always without the mark of the leaf base across the back.
The leaves commonly are about 10 cm. long by 1 wide, narrowly
lanceolate with a very long attenuate-falcate tip, mostly very
finely serrate, green and glabrous on both sides. The character-
istic venation of the Amygdalenae is at its fullest development
in Salix nigra. The marginal vein often runs almost to the very
base and the secondaries and tertiaries blend into a system of
meshes finer than in any other of our species.

Sometimes broad, blunt leaves are found at the bases of
lateral twigs but they are still easily recognized by their fine re-
ticulation. The aments appear with the leaves, capsules glab-
rous, short conic, short pedicelled, forming thin close cylindric
catkins which, supported as they are by the characteristic leaves,
resemble those of no other species. The staminate resemble
very closely those of S. amygdalotdes (which see), stamens 5 or
more, filaments pubescent.

Everywhere throughout the state, Salix nigra is our com-
monest willow. But it assumes a much more important role in
plant society along the southern border than further north.
There it attains its greatest size and at the same time becomes
much more abundant than elsewhere. For long distances along
the Ohio River it is almost the only native willow met with and
occupies all the territory which in the north is divided up between
several species.

So far as Ohio is concerned it is perhaps the most constant
and easily recognized of our willows but in the south it is almost
identical with S. humboldtiana and is rather hard to separate
from S. longipes. In the west it is represented by several
variable varieties which seem to connect it with related forms
and render it a very difficult subject indeed.

Its typical habitat is along streams but it may be found in
wet places generally though it seems to prefer moving to stagnant
water and is much more infrequent in swamps.

The variety “‘jalcata”’ is a form with narrower more falcate
leaves. In my opinion it is scarcely worthy of consideration
since it is not genetically different but is merely an accidental
leaf variation without correlated variation in other characters

Plate I. Salix mera.

Leaves of ordinary growth and of ranker growth with stipules; flowers:
and fruit typical; natural size; drawings of the flowers and capsule made
with camera lucida and photographed, enlarged seven times.

276 OHIO STATE ACADEMY OF SCIENCE.

of the plant. All the leaves are more or less falcate; their falca-
tion varies greatly on the same plant.

A hybrid between S. migra and S. alba has been reported
from New York.* It is said to have the catkins of S. alba and
the leaves of S. nigra x amygdaloides. Such a cross is most
surprising in view of the distant relationship of the parents. It
has not been reported from Ohio.

SALIX LONGIPES Shuttlew. Warpb’s WILLOW.

This species has not as yet been found in the state; its near-
est reported station is at the falls of the Ohio at Louisville.
Though this is some distance south of our territory it is possible
that it may be found along the Ohio River. I include it here be-
cause of that possibility and in order to make the paper more
useful outside the state. Its range extends from Missouri to
Washington, D. C., and southward to the Gulf. Dr. Glatfelter
says that it is not found like S. mgra sometimes away from the
banks of the streams but is strictly confined to them.

sometimes it grows into a tree like S. nzgra but much more
bushy. Around Washington it is a low shrub resembling S.
cordata surprisingly, considering the remotene +s cf their relation-
ships. The leaves have short stout petioles which with the mid-
ribs and larger veins are usually hairy. The blades are extreme-
ly variable. Frequently in rank growth they are auriculate at
the base; this character when present segregates them at once
from any other of our species. Sometimes they are very long,
oblong-lanceolate with straight edges narrowing gradually to the
tip; and this again is like no other of our species. More often
they are lanceolate with upper surfaces varying from shiny and
glabrous to dull and hairy. Sometimes they resemble those of
S. mgra closely except for the glaucous under surface. Or they
may be so similar to those of S. cordata as to deceive even the
expert; often this resemblance is especially well borne out by the
under surface which is at times gray glaucous and hairy exactly
like that species when grown in a dry place. The two can best
be distinguished by the venation which 1s simular to that of the
rest of the Amygdaleneae except that the marginal is hardly per-
ceptible and in its place the primaries ascend a long-way near the
margin.

From printed descriptions Salix amygdaloides might be con-
fused with the present species but they are nct very similar.
Salix amygdaloides is a much cleaner more shapely tree, never
shrubby; its leaves are long petioled, decidedly broader, much

* Bebb, Abrorescent Willows of North America 3. Gard. & For. 8:423
1895. Fig. 58.

THE WILLOWS OF OHIO. DahG:

more sharply pointed and never pubescent. While both have a
bloom on the under surfaces, S. longipes is gray glaucous and
S. amygdaloides bluish glaucous. Besides all this their ranges
do not overlap in Ohio.

In fruit it is easily distinguishable from either of the other
Amygdaleneae. The capsules are similar to those of S. mgra
but larger and long pedicelled like those of S. amygdaloides.

SALIX AMYGDALOIDES Anders. PEACH-LEAVED WILLOW.

Salix amygdaloides grows to medium sized or rarely to a
large tree. Its bark and general appearance suggest at once its
affinity for the black willow of which it was once considered a
variety. Its habit, however, differs very considerably from
that of Salix nigra. It is generally single-stemmed and very
shapely, with clean branches and darker brown, smoother bark.
The winter buds are nearly twice as large as those of Salix nigra,
dark brown above with a much lighter base where they were pro-
tected by the petiole of the old leaf in the fall. Salix amygda-
loides is well-named for when in leaf the tree, at a little distance,
often bears a striking resemblance to a peach tree; the twigs and
petioles are often reddened and the leaf arrangement is similar to
that of a peach tree. Both twigs and leaves are entirely devoid
of hairs while young shoots, at least, of the other black willows
are pubescent.

The leaves are much broader than in the other members of
the amygdalenae, often being almost ovate, distinctly broadest
below the middle with a rounded base and an attenuate falcate
tip, bright green above, glaucous beneath. The venation while
of the black willow type begins to approach the regular type, ex-
emplified by S. alba; the primaries are close, the marginal vein
short, hardly extending as far as the middle of the leaf, and
the meshwork, though very fine is much coarser than in Salix
nigra, while the secondaries are often more or less regular.

The catkins are so similar to those of the black willow that
it is difficult to distinguish them in dried specimens. With the
carpellate aments the difficulty disappears in fruit but with the
staminate it continues in old flowers. The brightening of the
bark at flowering time, which is noticeable in all willows renders
the twigs in Salix nigra very similar to those of the present species
and its smaller buds are swollen to about the size o those of the
Peach-leaved willow so that they can be used as a diagnostic
character no longer. The leaves supporting the catkins are too
young to have assumed their characteristic texture and the
aments themselves are almost the same. But Salix amygdalot-
des flowers nearly two weeks earlier than Salix nigra, which to-
gether with the habits should distinguish them in most cases.

SALIX AMYGDALOIDES.

Plate II.

THE WILLOWS OF OHIO. 279

The ripe capsules of Salix amygdaloides are narrowly long-conic
on long pedicels, contrasting with the short pedicelled capsules
of Salix nigra and giving the aments a very different appearance.

S. amygdaloides is characteristically a swamp plant though
it is not absent from river banks. In Ohio its range is over the
northern and western parts of the state. Columbus is near its
southern limit in central Ohio; further east it does not extend
so far south while further west I suspect 1t may even reach the
river. It is a north-western species of which Ohio is near the
eastern limit.

While remaining for the most part recognizable in the west,
Salix amygdaloides loses the glaucescence of the under surface
of the leaf while the upper surface brightens till it is almost like °
Salix lucida. The shape of the leaves also changes and becomes
shorter and broader. Southwestward it grades into the varieties
of S. migra which occupy the region. About St. Louis there is
a great complex of the amygdalenae in which pure forms of the
three constituent species are uncommon and there are all sorts
of interconnecting variations. In Ohio we have only two species
together and intermediates are rare though several have been
collected.

LUCIDAE.

Trees or shrubs, branches shining; leaves often very long
acuminate, broadly lanceolate to ovate, thick, glossy, strongly
glandular; catkins leafy peduncled, thick and dense, scales cadu-
cous, often dentate, filaments pubescent, capsules glabrous, style
short, stigmas thick.

SALIX LUCIDA Muhl. SuHininc WILLOW.

A bush or very rarely a tree 8 m. tall; bark smooth or near-
ly so; twigs shining orange brown, glabrous; buds rather narrow-
ly ovate, large, (5-10 mm. long) bright reddish brown in spring,
duller earlier in the season. Leaves reaching an extreme length
of 18 cm. and a breadth of 8 cm., varying from ovate to lanceolate,
rounded or narrowed at the base, prcminently glandular-serrate,
especially on the deciduous stipules, often covered when young
with long tawny scattered hairs, becoming glabrous, coriaceous,
very glossy above so as to give the plant a very beautiful appear-
ance different from any other willow. The thickness of the leaf
makes the rather regular veining difficult to make out. Stami-

Plate Il. Salix amygdaloides.

All parts typical, natural size except the capsule which is enlarged
three times.

LUCIDA

5 AILS

late ITI

ie

THE WILLOWS OF OHIO. 281

nate catkins borne with the leaves; easily recognizable by their
large diameter and fine appearance; scales large, conspicuous, cre-
nate or dentate, stamens mostly 3-5, filaments pubescent at the
base. Carpellate catkins also large, dense, 5-8 cm. long in fruit,
long persistent, scales obovate, slightly pubescent, pedicel near-
ly half as long as the capsule, stigmas large thick, deeply notched,
style short, capsule narrowly cylindric, about 6 mm. long.

Salix lucida is a northern plant occuring in the northern
third of the state but not extending to Columbus.

Salix lucida hybridises with Salix alba and S. fragilis and
when all three come together in one plant it creates very great
confusion in a group already very difficult to handle. Though
hybrids are mostly individual cases and irreducible to any general
type, it may be said that hybrids between S. lucida and one of the
fragiles are likely to have some of the following characteristics.
Nearly always the leaves are dull instead of glossy, sometimes
they are pubescent like S. alba. They are likely to retain some-
what of the prominently glandular character of S. lucida. Though
they may be almost typical of one of the other species in other
respects, they are almost certain to have the large reddish brown
winter buds of S. lucida. The stamens are oftenest two but the
catkins are likely to be short and thick asin S. lucida and dentate
scales are frequent.

Salix pentandra L. furopean Shining Willow.

Salix pentandra, the European species corresponding to our
Salix lucida, has been detected as an eccape in two places in the
state.* - It is not infrequently cultivated as a basket willow and
may be met with anywhere in cultivation.

it is most difficult to distinguich frem S. lucida and in scme
forms of the native species they canact be told apart with cer-
tainty. Salix pentandra, however, never has the long attenuate
leaf tips common in S. lucida, not even on watershoots. Its
leaves are rather thinner and not quite so glossy as those of S.
lucida. It dces not grow so rank and does not succeed well in
this climate. The ocier growers around Columbus complain that
it winter-kills.

I should not advice a beginner in Salicology to try to dis-
tinguish the two forms in the range cf Salix lucida but any form
outside the range, with the less acuminate leaves of S. pentandra,
may be suspected.

* Ses O. Nat. 4:12

Plate III. Salix lucida.

Large leaf from rank growth; smaller pair from ordinary twigs; the
short, broad, blunt one from the base of a branch; fiowers and fruit typical;
natural size; capsule enlarged three times.

Nov. 1903.

SALIX BRAGG

Pinter Ve

THE WILLOWS OF OHIO. 28%

FRAGILES, CRACK WILLOWS.

Trees, originally introduced from Europe and though now
freely escaped, still largely planted and often found growing in
rows (except the Weeping Willow). The bark of the medium-
sized branches is smooth, and yellow or greenish as distinguished
from the rough brown bark of the Amygdalenae. The leaves of
all the species are glaucous beneath and without stipules unless
very young. The catkins are borne with the leaves on lateral
branches; the stamens are normally only two; the capsules
glabrous and green, in flower at least, like the Amygdalenae but
unlike the other diandrous willows.

The Fragiles are clearly intermediate between the polyan-
drous tree-willows and the diandrous shrubs. But among them-
selves their relationships are not so clear. By hybridisation and
the importation of various horticultural varieties the group is
very much confused and consists of many very closely similar
forms. It has been the despair of many Botanists and one finds
more mistakes in the determination of this group than in any
other.

Key.
From habit.
Growing in clumps.
Bark of branches green. S. fragilis.
Bark of branches yellowish green or yellow S. alba.
Large trees, not in clumps.
Branches long, drooping. S. babylonica.

Branches not pendulous.

Large branches and trunk disfigured by many ad-
ventitious twigs, a tall tree with a central
shaft, branches yellowish. S. alba.

Without many adventitious twigs, a low broad
topped tree without a central shaft, branches
green. S. fragilis.

From leaves.

Leaves with a marginal vein, reticulations very fine, vena-
tion irregular, leaf narrow, acuminate, often strikingly
falcate, sharply serrate. S. babylonica.

Without marginal, reticulations not so fine, venation regu-

ular, leaf broader, not acuminate nor falcate.

Plate IV. Salix fragilis.

Leafy twig tpyical of our American form; the single broader leaf re-
sembles more closely the European form; natural size; capsute enlarged
three times.

284 OHIO STATE ACADEMY OF SCIENCE.

Leaves sharply serrate, pubescent at least when young,

glaucescence bluish. S. alba.
Leaves distantly blunt-serrate, glabrous, glaucescence
greenish. S. fragtlts.

From capsules.
Capsules long conic, short pedicelled, green or black in dry-

ing. S. fragilis.
Capsules short, ovoid conic, pedicel very short or none.

Capsules yellowing, catkins often long. S. alba.

Capsules green, catkins short. S. babylonica.

SALIX ERAGILIS. le CRACK Wolrow:

A tree reaching a height of 25 m. and a trunk diameter of
2.1m. When in clumps it forms a tall slender tree, but a single
individual growing alone branches out so that the head is as thick
as high. The bark of the trunk is roughish, gray, that of the
smaller branches green and of the twigs sometimes red, winter
buds large (7 mm.) seldom well filled out. Leaves reaching a
length of 17.5 cm. and a breadth of 4 cm.; acute, narrowed to
the base, coarsely serrate with prominent glands, glabrous,
greenish glaucous beneath, petiole short, stipules early fugacious,
venation regular, showing through the glaucescence as a dark
net work, primaries ascending, straight, close together, not arch-
ing. The aments being accompanied by leaves are easily identi-
fied by the leaf characters; stamens 2, with pubescent filaments,
ripe capsules long conic, green, with a short but distinct pedicel.

Salix fragilis is a European species planted extensively along
streams to hold the earth or to act as a wind-break. It is also
pollarded for its twigs which are valuable in basket making. It
is one of the most abundant species in our area. It is found both
planted and escaped everywhere. Our American plant is differ-
ent from the typical European form in having narrower leaves
but it is in most cases sufficiently distinct from S. alba.

Because of its quick rank growth, shapely habit, and beau-
tiful gray green foliage which turns over very prettily with every
breeze Salix fragilis is the best of the willows to plant for a shade
tree. It is difficult to understand why it has not supplanted
Salix alba and its varieties long ago. But the nurseries nearly
always carry a larger stock of the latter sorts which are much in-
ferior because of their habit of sending out suckers all over their
trunks.

Yt

THE WILLOWS ‘OF OHIO. 23%

SALIX ALBA L. WHITE WILLOW.

A tree attaining a maximum height of 30 m. with a trunk
diameter of 2.5m. Like Salix fragilis often growing in clumps
but when single-stemmed it is taller and more slender and has
the trunk continued as a central shaft to near the top. It is not
clean like that species but is covered with a brush of suckers.
Bark of the twigs and branches yellowish green varying to yellow;
winter buds smaller (4 mm. long) than those of the crack willow,
oblong and well filled out. The leaves reach a length of 13 cm.
and a breadth of 3 cm., lanceolate, acute, narrowed to the base,
closely and finely serrate, sometimes almost entire, grayish or
bluish glaucous, pubescent on both surfaces at least till mature,
(hair mostly persistent below) with close, fine, appressed, parallel,
gray hairs, stipules deciduous; primary veins close (closer than in
S. fragilis), straight, ascending, regular, extending to the margin
without branching, secondaries conspicuously regular but often
forking like the letter Y. Catkins on lateral branches or some-
times supported only by bracts, scales hirsute, deciduous, cap-
sules ovate-conic, not more than 4 mm. long, greenish yellow in
fruit, obtuse, glabrous, pedicel very short, style short, stigmas
thick.

Salix alba is a European species planted in this country for
the same purposes as S. fragilis. In most parts of the state it
does not seem to escape so readily as that species and hence is not
quite as common but may be found planted almost anywhere.

As stated above, most observers have considerable difficulty
in separating this species from the preceding. The difficulty is
often assigned to their hybridising propensities. But in Ohio at
least hybrids are rather rare. I have found that the two species
are distinct and separable in nearly all cases though it was only
after long study that the ability to distinguish them was acquired.
The manuals state that the species in the typical form is rare in
this country, the majority of the American forms being the golden
osier (var. vitellina). It is certainly true that few of our plants
are the typical hairy plant of Linneus but study of the European
material at Washington leads me to the conclusion that in Europe
the typical form is about as scarce as here. Further the extreme-
ly bright yellow twigs and glabrous, half shiny leaves of the typi-
cal varietal form are scarcer in this country than the pure alba
forms. It seems to me good practice in a series of intergrading
forms to draw the line between species and variety close to the
variety and to call all but nearly typical varietal forms the species
simply, if for no other reason to avoid the use of a trinomial.

The variety vitellina then, of Salzx alba includes those plants
with bright golden yellow twigs and branches, and leaves soon
glabrous and bright green.

SALIX -ALBAL

Plate V.

THE WILLOWS OF OHIO. 287

The blue willow, Salix alba coerulea, has not yet been recog-
nized in Ohio. Indeed there are some willow students of high
authority who do not distinguish it at all in this country whatever
may be its status in the old world.

As hinted above Salix alba hybridises with Salix fragilis
though not so frequently as might be supposed. It also crosses
with S. lucida as described under that species.

SALIX BABYLONICA L. WEEPING WILLOW.

The Weeping Willow grows into a large graceful tree 20—25
m. tall, easily recognized by its very long drooping twigs, which
have a habit peculiar to themselves of sending out numerous
short branches at a very acute angle with the main stem. Leaves
commonly rather small, 7-10 cm. long, about 1 cm. broad, nar-
rowly lanceolate, long acuminate and often falcate at the tip,
narrowed to the base, sharply serrate, glabrous unless very
young, greenish glaucous or at least paler beneath, petiole short,
stipules apparently absent; primary veins forming regular acute
loops which run together into a more or less straight marginal;
this together with the very fine reticulations caused by the rela-
tive prominence of the tertiaries often gives the leaf a more or
less close resemblance to that of S. nzgra, which, however, is
never glaucous as in the present species. Aments on rather short
few leaved peduncles, rather dense, not more than 3 cm, long.
capsule short conic, glabrous, green, all but sessile, style short.

It is aremarkable fact that the staminate plant of this species
is unknown in America. It is sometimes stated that it does not
occur at all and it is sufficiently rare to warrant such an assertion
but yet in the national herbarium is what I believe to be a gen-
uine specimen of the staminate flowers. It was collected by
Coville at_Ithaca, N. Y., in 1885 (?). The leaves are similar to
those commonly appearing with the carpellate catkins. The
aments are short, less than 25 mm. long, densely flowered with a
thachis densely covered with short hairs. The stamens are sub-
tended by a very short ovate scale which is much shorter than in
any other of the Fragiles.

Doubtless it is with the Weeping Willow as with the Purple
‘Willow that the absence of one kind of flowers prevents the nat-
ural spread of the species and is responsible for the fact that so few
plants have escaped when conditions for their growth seem so
favorable. But whatever the reason it is certain that the species
though commonly planted, esceaps very rarely. During the last

Plate V. Salix alba.

Leaves and flowers typical; natural size; capsule enlarged three times.
photographed and brightened with pencil.

SALIX LONGIPES AND SALIX BABYLONICA.

Plate VI.

THE WILLOWS OF OHIO. 289

seven years I have been continually on the lookout for it but
have seen less than half a dozen individuals which were not
clearly planted. One of these was on the Hocking River near
Sugar Grove; the others were along the lake shore in Ashtabula
county.

In Europe Salix babylonica hybridises freely with S. fragilis.
But in this country the manuals have not included such a cross.
A single plant was discovered at Sandusky during the season of
1903. * The leaves and habit were so exactly intermediate be-
ween the two that there could be no doubt of its identity.

At Ashtabula was found a plant which from leaf and habit
I take to be a hybrid between the present species and Salix alba,
which is not reported in the manuals.

LONGIFOLIAE, THE LONG-LEAVED WILLOWS.

The longifoliaé comprise a very distinct and compact group
of American willows. They have no close affinities with any
other group and do not intermingle with any. They have two
stamens (a specimen in the Ohio herbarium has three) and light
one-colored deciduous scales which show their relationship with
both the polyandrous and diandrous willows. Within the group
the many described species are difficult to recognize; Bebb, him-
self, said after he had described two or three of them that he did
not know but what they were all one species after all. The group
is very easily recognized by the venation of the leaves which is
different from any other willow and much resembles that of many
Onagaraceae, for example the fire weed. There is typically a
prominent marginal vein running clear round the leaf, connected
with the midrib by a series of distant nearly straight primaries be-
tween which there are practically no secondaries and no mesh-
work, only a few costals running parallel to the primaries. But
in young leaves the veins ascend at a much sharper angle and
the marginal vein is not so prominent while the secondaries and
tertiaries have not yet faded from view so that the above descrip-
tion will not hold. The leaves in some of the species are very
long and sometimes so narrow that it is difficult to find any veins
at all.

* Ohio Nat. 4:13 Nov. 1903.

Plate VI. Salix longipes (left) and Salix babylonica (right).
Typical specimens natural size; capsules enlarged three times.

SALIX INTERIOR AND SALIX INTERIOR WHEELERI.

Plate VII.

THE WILLOWS OF OHIO. 291

Key.

Leaves narrow, not over 1 cm., sometimes very long, second-
ary, (auxilliary) aments very much younger than the primary;
forming dense clumps of slender stemmed shrubs. S. interior.

Leaves often 1.5 cm. broad or more, not long in proportion,
secondary aments of about the same age as the primary, forming
a cymose cluster; low, bushy, not in close clumps; northern.

S. interior wheelert.

SALIX INTERIOR ROWLEE. LONG-LEAVED WILLOW, SANDBAR
WILLow.

(S. longifolia and §. fluviatilis of the manuals in part.)

The characteristic habit of this plant is to grow in clumps
about 4 meters high with a great many slender stems coming up
very close together from a common root system. These stems
are more slender than those of any other of our willows and
when the species assumes this habit it may be recognized
from a considerable distance. Unfortunately, however, it does
not always do so, but sometimes grows by itself as a bush or small
tree and then it can best be distinguished by its leaves. The
twigs are very slender, thickly branched and straight ascending
so that the stems have a peculiar brush-like appearance which
gives a pleasing softness to the landscape. The winter buds are
small and the twigs resemble those of the black willow more or
less closely as the bark is of about the same color. Many of the
buds are defective and drop off early inthe autumn. The places
‘of such are taken by small lateral buds which develop one on each
side of the old scar. This habit when present is characteristic.
It is largely responsible for the large number of small branches
which come out for there thus arise two twigs instead of one at
each node. The leaves are sometimes very long (to 15 cm.) and
not more than 1 cm. wide, of almost the same breadth throughout,
straight linear-oblong, serrated, but with shallow distant spinu-
lose teeth. When young they are spatulate. The venation is
very characteristic except in young leaves as described above.
Sometimes they are very hairy even being covered with matted
wool persistent until very old, but often they are glabrous and
green as soon as they unfold.

Plate VII. Salix interior and Salix interior wheeleri (marked W).

Leafy branch of the narrow leaved form of S. interior; flowers typical;
two fruiting aments shown, one nearly glabrous, the other with densely
tomentose capsules; natural size; capsule enlarged three times. Variety
uheelert typical cluster of staminate aments; the densely tomentose leaves
in the upper corner are from the form which connect with the species;
the larger broad glabrous leaves shown below are the extreme form of
the variety farthest from the species, natural size.

292 OHIO STATE ACADEMY OF SCIENCE.

The flowering season of Salix intertor is much longer than
that of any other of our willows; it begins just after the pussy
willows have gone by and continues late into the summer. I
have even seen blossoms in October and on Cedar Point they are
not uncommon in July and August. The first catkins come out
on short peduncles with a few small bracts. Later when the
season’s twigs have developed, they also bear aments at their
tips. Just below these terminal catkins develop other lateral a-
ments which blossom later and so prolong the season. We have
no other willow which does this and the presence of these small
undeveloped aments is very characteristic. The carpellate
aments are generally but not always quite lax as they grow older.
The flowers have a tendency, sometimes very marked, to appear
in fascicles of from three to five on the rachis with a distinct in-
terval between them. This is another characteristic feature
present in no other species. The scales are yellow, deciduous,
the filaments frequently pubescent. The ovularies at anthesis
are scarcely longer than the scales with sessile stigmas on their
summits. They vary much in shape being sometimes, especially
when very hairy, thick and short with a squarely cut off tip,
sometimes nearly rostrate especially when glabrous. The ma-
ture capsules are narrowly conic, blunt pointed so as to be almost
cylindric if not well fertilised. When well developed they are
quite large (1 cm.) sometimes glabrous sometimes tomentose.
This variation makes them a puzzling problem and one would sup-
pose there were several species instead of one but there seem to
be no lines of cleavage between the different forms.

Salix interior is with the exception of its own variety wheelert
and the Texan S. thurberi the only representative of the longifo-
liae east of the Rockies. It extends all oves the Mississippi valley
and is occasionally met with east of the Alleghanies. In Ohio
it 1s common everywhere.

Salix interior var. wheeleri Rowlee. Wheeler's Long-leaved
Willow.

This variety as I have seen it in Ohio sometimes acquires a
slender tree form, but more generally is a low much branched
dwarf bush, spreading in the sand by the sprouting of buried
stems. These do not as in the species produce a dense clump of
stems close together but come up only at distances of a meter or
so and the result is a loose clump the members of which appear
like independent plants. In extreme forms, the leaves especially
the older ones from the axils of which branches come out, are
very much broader than in the species (7-10 cm. x 2 cm.), dark
green and glabrous with the typical venation of the longifoliae,
except that the primaries are rather closer and more ascending.
These extreme forms as they intergrade into the narrow glabres-
cent leaves of the species pass through a series of forms which

THE WILLOWS OF OHIO. 293

are somewhat narrower but so extremely wooly that neither sur-
face of the leaf can be seen. It was from these that Prof. Rowlee
took the type of the variety. These intermediate leaves shade
into narrow tomentose forms which connect with the narrow
glabrous leaves of the typical species. One might doubt, if he
had not studied the plant in the field, that the broad glabrous
leaves were the extreme form, were they not accompanied by an
extreme in floral development which is more significant than the
leaf characters on which Prof. Rowlee separated the variety.
This is in the development of the secondary aments at the base
of the terminal. In the variety the aments often form clusters
of half a dozen, all opening at nearly the same time—a thing
which is rarely seen in S. znterior itself. The flowering period is
also distinctly later in the variety than in the species. On Cedar
Point it seems to be at its height the first of July and continues
into August. Fully 99 % of the plants on Cedar Point are stam-
inate. To what the dearth of carpellate plants is due it is not
possible to say at present.

As yet the variety wheeler: has been found in Ohio only at
Painesville (H. C. Beardslee no. 67, fide Rowlee) and on Cedar
Point where it is abundant. It possibly occurs all along the lake
shore and possibly for some distance back into the country. But
I was unable to find it in Ashtabula county though the conditions
seem favorable. Its author limits its range to the basin of the
Great Lakes.

CAPREAE.

Low trees or shrubs with leaves ordinarily broad in propor-
tion to their length, generally glabrescent above, mostly tomen-
tose beneath, catkins appearing very early, oftenest in pussies,
capsules, in our species, villous.

Key.
From leaves.

Leaves ordinarily very tomentose below especially on the

rugose veins, venation strongly sunken above, northern.
S. bebbiana.

Leaves often glabrous, veins not strongly raised on the
under surface, nor depressed above, when hairy often
with red-brown hairs. - S discolor.

From flowers.

Catkins appearing with the leaves or only a little while be-
fore them, small at anthesis; scales yellow or darkened
only at the tip, capsules narrowly cylindric, filaments
not coarse nor long. S. bebbiana.

Catkins appearing much before the leaves in large pussies,
scales dark brown, capsules elongate-conic, filaments
coarse, long. . S. discolor.

P’ate VIII.

SALIX BEBBIANA.

2 it

THE WILLOWS OF OHIO. 295

SALIX BEBBIANA Sarg. BeEBB’s WILLOW.

A shrub or small tree occasionally reaching a height of 8
meters, with a habit almost exactly like that of S. discolor and
conspicuously different from that of all of the other shrub wil-
lows in that there is scarcely any tendency to sprawl, but the
stems all ascend from the root. Leaves generally elliptical,
varying from sharply serrate through undulate-serrate to entire
or often slightly revolute, generally glabrescent above, wooly be-
low at least on the veins, primaries and secondaries prominently
raised on the under surface making them very conspicuous, pri-
maries rather distant, inclined to be crooked and often forking.
The whole system of veins strongly sunken from above. Catkins
appearing just before or with the leaves, with leafy bracts or, in
fruit, on leafy branches; staminate 3.5 cm. long or less; carpel-
late sometimes 6 cm. in fruit; scales yellow or slightly darkened
at the tip, pubescent, persistent in fruit; capsule long pedicelled,
villous with white silky hair, cylindric, obtuse, sometimes 11 mm.
long in fruit.

Salix bebbiana is found across the northern third of the
state. :

This species in its normal forms is very distinct from Salix
discolor and can be separated from it withont the least difficulty
but the western forms though most keep their flowers like the
type, have leaves resembling those of S. discolor more or less
closely; sometimes even so closely as to be indistinguishable
from it. One of these plants from the middle west almost half
way between the eastern and western forms of the species Dr.
Rydberg has named S. perrostrata. Unfortunately, however, the
difficulty in separating the two species, though worst in the west,
is not confined to that region. Some specimens from Ohio are
so nearly intermediate that they can scarcely be determined, but
these are rare. There is no danger of any specimen with mature
leaves or in fruit being confused with any other species than S.
discolor for it resembles none, but both kinds of flowers, while
the bracts are yet small, resemble those of S. petiolaris and the
staminate are similar to those of S. candida.

Plate VIII. Salix bebbiana.

Typical specimens; two fruiting aments one with and the other with-
out leafy bracts, natural size; capsule enlarged three times.

» SALES DISCOLOR:

Plate 1X

THE WILLOWS OF OHIO. 297

SALIX DISCOLOR Muhl. Pussy WILLow.

The pussy willow is typically a swamp shrub growing in
clumps differing from those of S. sericea or S. cordata in that each
plant is usually a close clump, separated from its neighbors by a
distinct interval, while those species run over a considerable area
in a loose clump. The stems are not ordinarily recumbent but
strictly upright and straight. Twigs of swamp plants rank,
sometimes almost 10 mm. in thickness varying from glossy to
densely tomentose, with very large well filled purple-brown buds.
In less luxuriant growth the twigs may be smaller, sometimes
wooly, with smaller buds. Leaves varying from ovate to spatu-
late, coarse serrate with blunt incurved teeth to entire or even
slightly revolute, glabrescent above, beneath glabrous and paler
to glaucous or sometimes tomentose or pilose. Hair soft and
wooly as in S. bebbiana or short, straight and ferruginous.

Pussies before anthesis larger than in any other species and
consequently this is the favorite species with the children in
search of pussies in the spring. At anthesis the staminate with
their long coarse filaments are larger than any other of our wil-
lows except S. lucida; carpellate also very large, sometimes 13
em. in fruit, scales dark brown, capsules long (8 mm.), rostrate, °
gray pubescent to glabrate in age, pedicel sometimes nearly as
long as the capsule but usually shorter. The flowering time is
earlier than any other of our willows and it lasts such a short time
that it frequently happens that Salix discolor blossoms and goes
by before one gets out after it, a difficulty not met with in any
other of our willows. When the other pussy willows are found
in flower it is generally in fruit so that there is little danger of
confusing it with them.

As descrlbed above Salix discolor includes forms differing
from each other very strikingly. But the longer I study them
the surer I am that, diverse as they are, all are one species. The
great differences are all in characters like the shape, surface and
pubescence of the leaves, which are subject to considerable vari-
ation and are to a great extent the outcome of various environ-
mental conditions. The catkins also vary somewhat but in
studying specimens from marked trees taken in flower and leaf I
have been able to find no correlation between the separate varia-
tions in flower and leaf.

Plate IX. Salix discolor.

Leafy twig of the most common form; other leaves shown below from
left to right as follows: broad, blunt, tomentose form resembling S.
nigricans of Europe connecting with S. bebbtana (rare); narrow ferrugi-
nous form often found on plants which would pass as S. ertocephala;
long, narrow, nearly entire form (var. prinoides); twig with winter bud
and young pussies; staminate flowers typical; carpellate ament a little
narrower than usual; fruiting ament typical except that it is unusually

long; natural size; capsule enlarged three times.

SALIX HUMILIS AND VAR. TRISTIS.

Pileiye

THE WILLOWS OF OHIO. 299

Salix eriocephala Michx. of Britton’s manual includes those
forms with ferruginous hair on the leaves and tomentose twigs.

Salix prinoides Pursh, 1s a narrow leaved form of the type.

It is common in swamps all over the state.

This species is most difficult to separate from S. cordata in
leaf but its upright habit and the coarse serration as contrasted
with the sprawling habit and sharp-toothed leaves of S. cordata
are sufficient to distinguish them. As described under Salix
bebbiana it sometimes connects with that species. Narrow revo-
lute-leaved forms are sometimes found which connect this species
with S. humilis, probably some of them are hybrids.

SALIX HUMILIS Marsh. PRAIRIE WILLOW.

A shrub not more than 3 meters tall with spreading often re-
cumbent branches. Leaves oblong or spatulate, gradually nar-
rowed to the base, abruptly acute and sometimes mucronate at
the tip, mostly revolute, entire to undulate-dentate, puberulent
or glabrous above, tomentose, especially on the prominently rais-
ed veins beneath. or glabrous and glaucous, primary veins rather
distant, inclined to be horizontal, looping or oftener branching
and arching, with several costals between them, secondaries
quite irregular, catkins born very much as those of S. discolor,
long before the leaves but smaller than in that species, from
short stubby pussies, staminate 2 cm. long or shorter, carpellate
occasionally 4.5 cm. in fruit, bracts small or none, scales dark-
ened above, long pilose on the back, glabrous in front, capsules
elongated, often rostrate-conic in fruit, hirsute at least when
young, pedicelled, sometimes almost 1 cm. long when ripe, style
distinct, red.

Salix humilis though common nowhere is generally distri-
buted over the state. It will probably be found growing on dry
hillsides in nearly every county.

In most forms Salix humilis is easily recognizable in leaf
because of the long narrow revolute leaves. In flower it is char-
acterized by the short stubby pussies from which the flowers
come. The leaves of the ranker shoots take on an appearance
very similar to those of Salix discolor. It is from such branches
that many of the so-called hybrids of our herbaria came but real
hybrids undoubtedly do occur. Sometimes also the present
species is very similar to S. candida, but ordinarily it is a gray
plant while S. candida has snowy white wool on the under side

Plate X. Salix humilis and variety trtstzs.

Series of leaves from S. humilis (lower row) connecting with the va-
riety tristis (upper row); flowering aments from var. tristis, fruiting from
S. humilis; natural size; capsule from S. humilis drawn in with camera
lucida, enlarged five times.

X SERICEA.

SAL

Plate XI

THE WILLOWS OF OHIO. 301

of its leaves contrasting strongly with the rich green of the upper
surface. In some hybrids, however, most noteworthily Salix
candida x. S. petiolarts the resemblance is so close as to make it
all but impossible to separate the two. But such hybrids usually
have a distant scolloped serration derived from S. petiolaris
which is different from any form of S. humilis. At flowering
time there will be no difficulty in separating them.

Salix humilis var. tristis (Ait.) Dwarf Gray Willow.

A depauperate form of S. humilis with which it is connected
by many intermediates. It may be described as smaller and
hairier throughout. It is quite low (to 6 dm.) with smaller
leaves (to 5 cm. long) more strongly gray tomentose, and catkins
sometimes scarcely 5mm. long. This can hardly be regarded as
a distinct species. There is not a single constant character by
which the two differ and what differences there are, are such as
would be likely to be caused by differences in environment.
Such forms should be considered as varieties rather than as
species.

SERICEAE.

Swamp shrubs, leaves narrowly elliptic-lanceolate, com-
monly tapering to both ends, serrate with blunt cartilaginous
teeth to entire, glabrous to glaucous, sericeate below when young,
generally blackening in drying. Aments born before the leaves,
sessile or short peduncled, scales darkened at the tip, pilose,
capsules pedicelled, silvery sericeate at least when young.

Key.
From leaves.

Leaves dull above, silvery silky beneath at least until very
old, scarcely glaucous. S. sericea.

Leaves shining above, glabrous on both sides except when
young, pubescence on young leaves mostly ferruginous,

quite glaucous below. S. petiolaris.
From flowers.

Fruiting catkins dense, sessile, capsule ovoid-conic, obtuse,
short pedicelled, sericeate even when ripe, hardly +4 mm.
long. S. sericea’

Fruiting caktins, looser, short peduncled, capsule becoming
cylindric conic, acutish, long pedicelled, nearly glabrous
when mature, more than 4 mm. long. S. petiolarts.

Plate XI. Salix sericea.
Typical leaves flowers and fruit; natural size; ripe capsule drawn in
with camera lucida, enlarged five times.

302 OHIO STATE ACADEMY OF SCIENCE.

SALIX SERICEA Marsh. SitKy WILLow.

A shrub seldom 5 m. tall, with straggling branches generally
lopping over onto the ground, forming clumps very similar in
appearance to those of Salix cordata, buds similar to those of
that species but without the loose inner membrane, twigs also
similar but very brittle at the base. Leaves narrowly lanceolate,
cuneate or rounded at the base, sharply serrate to almost entire,
rather dark green and glabrous above, typically dull, below sub-
glaucous, silvery sericeate (rarely somewhat ferruginous or glab-
rate when old) with lustrous hair which reflects the hight irregu-
larly at different angles almost like changeable silk; venation
very similar to that of Salix cordata but the primaries ascend less
rapidly toward the margin than in narrow leaves of that species,
the looping is more prominent and blunter or sometimes, even
the primaries are quite irregular. Catkins opening before the
leaves, supported by a few small green bracts or unbracted,
10-30 mm. long, very dense, scales dark and pilose at the tip,
light below, anthers generally red, capsule short conic or ovoid,
obtuse, silvery sericeate even when ripe, not more than 4 mm.
long, style short but distinct, pedicel about half as long as the
ripe capsule.

Salix sericea prefers boggy land and is so common that it
may be found in almost any swamp in the state but it seldom
grows along river banks with Salix cordata.

The greatest difficulty in defining Salix sericea is to separate
it from S. petiolaris, see below, and from S. cordata. In both
cases the carpellate aments are entirely diagnostic and it is with
the leafy specimens that the difficulty occurs. The leaves of
Salix cordata are ordinarily wider than those of S. sericea and
broadest below the middle while in this species the greatest
width is near the middle; the serrations are sharper and finer in
S. cordata and the leaves of S. cordata do not blacken in drying.
But most important of all are the silvery hairs below which dif-
ferentiate it from everything else. In old specimens, however,
they are sometimes almost absent and in such cases it frequently
becomes almost impossible to separate them.

Salix sericea subsericea, (Anders.) Rydb. is the name that
has been given to the form of the present species which connects
it with S. petiolaris. It is characterized by leaves much less pu-
bescent than the typical form and less dull or even subglossy on
the upper surface so that from leaf characters alone it cannot be
distinguished from that species. It is very common in our area,
see under S. petiolaris.

wo

THE WILLOWS OF OHIO. 30

' SALIX PETIOLARIS Smith. SLENDER WILLOW.

A shrub very similar to Salix sericea in habit. But typical
leaves are quite different from those of that species in general
appearance. Unfortunately, however, in this region connecting
forms sometimes render it impossible to distinguish the two from
the leaves alone. They are rather narrower, margin (sometimes
entire) serrate with more prominent, blunt cartilaginous teeth,
subglossy above, quite glaucous beneath, or sericeate when young
oftenest with ferruginous hair, venation so similar to that of S.
sericea that it cannot be distinguished unless by the more ascend-
ing primaries and the finer reticulations. Carpellate aments
short peduncled, becoming somewhat lax in fruit, scales yellow,
mostly darkened above, pilose, capsules long pedicelled, acutish,
in fruit cylindric conic, more than 4 mm. long.

The character of the leaves in the extreme form is very simi-
lar to that of the Broad-leaved Willow with which they might
be confused were it not for their narrowness. The difficulty in
separating them from S. sericea comes especially late in the sea-
son when that species begins to lose its pubescence and to become
more glossy on the upper surface. The catkins sometimes re-
semble those of Salix bebbiana but can be distinguished by the
darkened rather than yellow scales. With these exceptions the
present species is not likely to be confused with any other in our
area. .

Salix petiolaris is the western form of Salix sericea or rather
since that species was later named, it is the eastern form of S.
petiolaris of which it was made a variety by Andersson. It almost
seems as though that were the proper treatment and that we
should be doing well to return to Andersson’s view but the carpel-
late aments seem fairly distinct.
= Weare near the eastern border of the range of Salix petio-
larts. It occurs rather rarely in Ohio and has been found only in
the north-western portion of the state, though it may extend well
eastward along the lake shore.

Salix pettolaris gracilis Anders. Frequently one meets with
forms of the Slender Willow with even more slender and graceful
twigs than the usual form. This variety has been given the name
gracilis. by Andersson. It is characterized by narrower, more
sharply serrate leaves and longer pedicels than the typical form.
It may perhaps be taken as the extreme development of the
type farthest from S. sericea. It is to be expected wherever S.
petiolaris is found.

PETIOLARIS.

D SALTEEX

FAN

- SALIX CANDIDA

Il

xX

Plate

THE WILLOWS OF OHIO. 305

SALIX CANDIDA Fluegge. SaceE WILLow, Hoary WILLow.

This little shrub seldom grows more than a meter tall. It
may be recognized anywhere by its leaves which are narrowly
oblong and revolute, veins deeply depressed on the upper sur-
face and prominent below; under side covered with a thick white
tomentum contrasting strongly with the rich dark green of the
upper surface, petiole short and stipules lacking. Flowers ap-
pearing before the leaves; ovularies densely covered with silvery
white wool, nearly sessile, with a very long conspicuous deep
rose red style like no other of our species; staminate catkins with
few bracts below them and hardly presenting diagnostic charac-
ters; small and delicate resembling those of S. bebbiana some-
what but easily distinguished from them by the dark colored
scales.

Salix candida is quite rare in Ohio. It was first reported by
Mr. Moseley from Castalia prairie but has since been found in
Wyandot county also. It ranges over the eastern and northern
portions of North America. But in the west and north the
leaves apparently become broader almost elliptical and not mark-
edly revolute.

Salix candida hybridises with S. cordata, with S. sericea and
with S. petiolaris. At Castalia a fine series of hybrids with S.
cordata and S. petiolaris may be found.

CORDATAE.

The cordatae are a group of shrub willows with very variable
leaves, characterized by glabrous capsules borne from wooly
pussies. A peculiarity of the opening buds is that the inner bud
scale grows out beyond the outer, enveloping the base of the
ament and looking like the wing of a beetle imperfectly folded
under the elytron. This so far as I know occurs in no other
group and is therefore an important diagnostic character at a
time when the species are particularly hard to separate. In this
state the species though variable are fairly well marked but in
the west the group is represented by a number of forms whose
relationships have not been satisfactorily worked out as yet.

Plate XII. Salex candida (left )and Salix petiolaris (right.)

S. candida—typical but rather short leaves, other parts typical;
natural size; capsule and carpellate flower enlarged three times, photo-
graphed and brightened.

S. petiolaris—leafy twig of the extreme type farthest from S. sericea,
with shiny, glaucous, glabrous leaves; two small leaves belonging to the
variety gracilis in the lower corner; flowers and fruit typical, natural size;
capsule enlarged three times.

SALIX CORDATA.

Plate XIII.

THE WILLOWS OF OHIO. 307

Key.
From leaves.

Leaves glossy above, heavily glaucous below, mostly broad,
ascending so as to show their white under sides, glab-
rous, northern. S. glaucophylla.

Leaves neither glossy nor very glaucous.

Leaves thick, ovate, with emarginate bases, tomentose

on both sides, northern. S. adenophylla.
Leaves thin, seldom pubescent above, lanceolate, gen-
erally distributed. S. cordata.

From flowers.

Bracts strongly glandular, before anthesis very villous,
aments dense, capsules green, short pedicelled, medium
sized, northern. S adenophylla.

Bracts not glandular, tomentum quickly evanescent, pedicel
longer.

Bracts green, obscurely serrate or entire, capsules green,
small, medium pedicelled. S. cordata.
Bracts blackening, glaucescent, capsules large, rostrate,
becoming brown, long pedicelled, aments lax;
northern. S. glaucophylla.

SALIX CORDATA Muhl. HErart—LEAVED WILLOW.

Salix cordata is the botanist’s bugbear. It is one of the
most common and most variable species with which we have to
deal. Its leaves seem to be under no restraint of heredity and
may assume almost any form and any character. I have seen
them all the way from linear-lanceolate to orbicular, from sub-
glossy to heavily tomentose, from green to glaucous. But var-
ious as are the leaf forms they are all united by the flowers and
one cannot doubt that they all belong to a single species.

The separation of the bud scales alluded to above is particu-
larly well marked in Salix cordata and in the southern portion of
the state or anywhere out of the range of the other species at
once differentiates this species from S. sericea or S. discolor
which it resembles much in early spring.

Leaves very variable but generally of the lanceolate type,
broader or narrower, but widest below the middle, serrations
always sharp and usually fine. In leaf it is most likely to be con-
fused with S. discolor and S. sercicea. From the former it can

Plate XIII. Salix cordata.

Leaf showing venation and the two others in the lower portion of
the plate typical; large leaf from very vigorous water shoot, perhaps indi-
cating some affinity with S. adenophylla; upper narrow leaf from variety
angustata; flowers and fruit typical; natural size; capsule enlarged three
times.

X ADENOPHYLLA

SALI

Plate XIV.

THE WILLOWS OF OHIO. 309

be distinguished by its sharp fine serration contrasted with the
distant blunt and often coarse teeth of the pussy willow. The
habits of the two are sufficientiy different to put aside all confu-
sion when the plants are seen together. From S. sericea it can
generally be distinguished by the absence of the silvery white
pubescence on the under surface. The shape of the leaves is also
different, in most cases in that species the leaf is widest near the
middle; in this it is widest below the middle. Salix cordata also
lacks the peculiar leaf habit of Salix sericea. The flowers come
very early from small pussies. As they mature the carpellate
aments come to be supported by large leaves and much of the
wool of the pussy drops off from the fruiting rachis. The anthers
just before the elongation of the filaments are almost as red as
those of Salix sericea. The capsules are green and glabrous,
the stigmas frequently red.

Salix cordata angustata Anders. includes the narrow leaved
forms of the species. In Ohio most plants have leaves wider
than those of the typical angustata but decidedly narrower than
the typical specific form. It is therefore difficult to distinguish
two forms in our area and since the leaf variation may be con-
sidered as accidental and without significance it 1s perhaps hardly
advisable to separate them.

Salix cordata is abundant all over the state. Its usual
habitat is along streams while the other species with a similar
habit and leaf are typically swamp plants. This is not to say
that the present species never grows in swamps nor that S. sericea
and S. discolor never grow along river banks—for they do—but
that they attain their best development in the habitats given
and are usually found there.

To increase the difficulty of dealing with Salix cordata it
hybridises very freely. It forms with S. candida a fine series of
connecting forms. With S. sericea hybrids occur though not so
frequently as has been supposed. It is also said to mix with S.
discolor but I have seen no unquestioned specimens from Ohio.

SALIX ADENOPHYLLA Hooker. Furry WILLOW.

A straggling shrub of about the same size as S. cordata
which it resembles most closely. It looks like a xerophytic
adaptation of that species. The leaves are thicker, shorter and
broader, ovate, more or less tomentose on both sides, with an
emarginate base, very sharply serrate or entire; ordinarily in
rank growth with the leaves closely crowded on the twigs and

Plate XIV. Salix adenophylla.

__ Typical leaves, flowers and fruit, showing fruiting aments with and
without bracts; a bract from a fruiting ament in the upper corner; natural
size; capsule enlarged three times.

eae ee eee eee

LLA.

SALIX GLAUCOPHY

Plate XV.

THE WILLOWS OF OHIO. Bil

generally accompanied by large stipules. The flowers differ
from those of our other species of cordatae in having broader
bracts, much woolier before anthesis; in fruit by the denser cat-
kins with shorter pedicelled capsules which are smaller than
those of S. glaucopiylla and rather larger than the average of
S. cordata.

In typical forms the wooly broad leaves will distinguish it
from everything else in our area. It is sometimes difficult to
separate, however, from S. cordata in its more tomentose forms,
and intermediates seem to occur. In shape the leaves are not so
different from S. bebbiana but they will be quickly distinguished,
among other things, by the sharply serrate margins of the present
species.

Salix adenophylia is our rarest willow. It is a plant of the
Western Great Lake region and reaches its best development in
Michigan, being rare east of that state. Until recently it was
not supposed to extend into Ohio but I have seen several speci-
mens collected at Erie, Pennsylvania. It should therefore ex-
tend the whole length of the Ohio Lake Shore. But it is very
scarce indeed. Though I have hunted for it on Cedar Point and
in Ashtabula County I have seen from Ohio but a single undoubt-
ed specimen which was taken by A. D. Selby on Cedar Point.

SALIX GLAUCOPHYLLA Bebb. BROAD—LEAVED WILLOW.

A shrub sometimes 5 m. tall, growing in clumps like S. cor-
data; leaves mostly broad, ovate or ovate-lanceolate, shiny above
except in rank growth, with very fine irregular venation and
stiff midribs which hold them up at an acute angle with the stem
so as to display the white under surfaces. In character the leaves
resemble much the narrower leaves of S. petiolaris and like them,
when succulent have a strong tendency to blacken in drying.
Flowers appearing before the leaves, bracts mostly narrow, glau-
cescent; staminate aments generally larger than those of S. cor-
data with smaller bracts; carpellate long, lax in fruit, capsules
very long (1 cm. or more) glabrous, sometimes rostrate, long ped-
icelled, turning brown.

Bebb labelled some of the first material of this species he
sent out, Salix cordata x S. lucida. This determination though
very far from correct as he himself soon recognized, is descriptive
of the species. Its affinity to Salix cordata is evident at once
from inspectoin of either leaves or flowers but yet in both there
is a strong resmeblance to Salix lucida. The large thick catkins
with the long capsules and the beautiful glossy leaves cannot but
suggest that species.

Plate XV. Salx glaucophylla.
Typical leaves and aments, natural size; capsule enlarged three times.

Plate XVI. Satix PURPUREA AND SALIX PEDICELLARIS.

THE WILLOWS OF OHIO. ole

The Broad—leaved Willow is a species of the Great Lake ba-
sin and in Ohio is confined to the northern part of the state. It
is common along the western portion of the Lake Shore but pro-
bably does not extend beyond Cleveland.

Salix glaucophylla at times seems to grade into Salix cordata
by what are probably a series of hybrids. In other cases it is
very difficult to separate from S. discolor with which it also pro-
bably hybridises.

MYRTILLOIDES. BOG WILLOWS.

Low shrubs with mostly elliptical, glaucous, glabrous leaves
strongly reticulate veined and purplish green; aments and bracts
usually reddened, small, few flowered; filaments often partially
united showing their affinity with the following group; a group of
three or four species all very similar to the European Salix
myrtilloides.

SALIX PEDICELLARIS (Anders). AMERICAN BocG WILLOow.

A low shrub seldom reaching a height of 1 meter, aereal
shoots erect, slender, supported in the sphagnum by long creep-
ing stems which run far down into the bog putting out numerous
rootlets along their length. Leaves when fully grown sometimes
9 cm. long and 20-25 mm. broad, but ordinarily smaller, elliptical,
oblong, spatulate or rarely obovate, pointed at both ends, entire,
slightly revolute, dark purplish-green above, slightly glaucous
below, not hairy unless when very young, nearly sessile, venation
conspicuously reticulate with meshes large and coarse consider-
ing the size of the leaf. Catkins appearing with the leaves,
loosely flowered, not more than 25 mm. long in flower, but occa-
sionally 5 cm. in fruit, scale short, often no longer than the nec-
tary, as broad as long, filaments often more or less united; cap-
sules long pedicelled, nearly always glabrous, short conic to cylin-
dric-conic, obtuse, sometimes 8 mm. long in fruit, with a decided
tendency to turn red or purple.

Salix pedicellaris is a northern species growing in cold peat
bogs where it may be easily recognized by its small size and pe-

Plate XVI. Salix purpurea (left) and Salix pedicellaris (right.)

S. purpurea. Typical branch with opposite and scattered leaves:
flowers and fruit typical, the latter from a European specimen; natural
size; staminate flower and carsule enlarged three times, photographed
and brightened.

S. pedicellaris. Typical summer branch with small ovate leaf from
the base of a twig to its left; staminate and carpellate flowers and fruit
typical, the latter not ripe; natural size; staminate flowers and capsules
enlarged three times, one of them, marked M, between the leaves of S.
purpurea. 4

314 OHIO STATE ACADEMY OF SCIENCE.

culiar leaves. It it necessarily quite local in its distribution but
probably grows in most of the sphagnum bogs in the state.
There are specimens in the state herbarium from Williams and
Portage counties and from the Licking Reservoir.

I am informed by Mr. P. A. Rydberg that the typical S.
myrtillordes of Linneus was a European plant and that it differs
slightly but constantly from the American species, hitherto con-
sidered as the variety pedicellaris, which should consequently
be considered distinct.

As stated above Salix pedicellaris is easily distinguished from
all the other species with which we have to deal. But more than
once I have strongly suspected it of hybridising with S. discolor
which occupies the same territory. But so far I have not yet
been able to satisfy myself of the hybridity and so leave the
question in abeyance.

PURPUREAE.

Shrubs, leaves mostly oblong, oblanceolate or linear-oblan-
ceolate, nearly entire, glabrescent, stamens more or less perfectly
united into one with 4 anthers, capsules globose-conic, nearly
sessile, silky.

SALIx PuRPUREA L. PuRPLE WILLOW.

A shrub reaching a height of about 3 m. putting forth a

dense growth of slender wands from the larger branches. Leaves
scattered or opposite, sometimes 10 cm. long and 2 cm. broad,
oblong-oblanceolate, oblong or rarely elliptical, mostly broadest
above the middle, abruptly acute, generally gradually narrowed
to the round base, entire or obscurely serrate, glabrous, dark pur-
plish-green, paler beneath, petioles short, stipules apparently
absent. Catkins expanding from pussies, sessile with a few small
green bracts, scales more or less pilose on the back, oblong, blunt
with conspicuously purpled tips, very concave above and strongly
reflexed in the staminate, less so in the carpellate, capsule broadly
ovoid, silky, sessile, style very short or none.
Salix purpurea is an Old World species planted in America largely
for its twigs which are much used in basket work. It has been
long reported in the manuals as escaped in this country but
though it is fairly common over the state it is rarely that one sees
a clump growing in a place where it would not likely have been
planted. It certainly has not esceped to any such degree as have
S. alba and S. fragilis. In America the carpellate plant is very
rare and the species is mostly propogated by cuttings. This may
account for its inability to spread as in the case of Salix babylon-
aca.

ae