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TABLE OF CONTENTS

* I ^"^^No. I. Symbiosis and Saprophytism, by Dr. D. T. MacDougal.
iS'^'^No. 2. New Species from the western United States, by Dr. Per Axel
Rydberg.
/^'l ^ No. 3. The dichotomous Panicums : some new Species, by Geo. V. Nash.
l'?'^^ No. 4. Delphiniuvi Carolinianum and related Species, by Dr. Per Axel

Rydberg.
f[(>0 No. 5. Studies on the Rocky Mountain Flora— I. by Dr. Per Axel

Rydberg.
{^OO No. 6. Notes and Descriptions of North American Plants 1 and II, by

Dr. J. K. Small.
C^^DNo. 7. Vegetative Reproduction and Multiplication in Ery t hroniuin, hy

Frederick H. Blodgett.
|9 J No. 8. Two new Species of Grimmia from Montana, by R. S. Williams.
|9(;, uNo. 9. Studies on the Rocky Mountain Flora — II, by Dr. Per Axel
Rydberg.
\°(0(:jNo. 10. Studies on the Rocky Mountain Flora— III, by Dr. Per Axel

Rydberg.
\^q\ No. II. Life history oi Schizaa piisilla, by Elizabeth G. Britton and Alex-
andrina Taylor. ^

■ q^l No. 12. Studies on the Rocky Mountain Flora— IV, by Dr. Per Axel

Rydberg.
\'^0\ No. 13. Further Studies on the Potentilleae, by Dr. Per Axel Rydberg.
icj^, I No. 14. Studies on the Rocky Mountain Flora— V, by Dr. Per Axel

Rydberg.
\'\6\ No. 15. Studies on the Rocky Mountain Flora — VI, by Dr. Per Axel
^i Rydberg.

\^0\ ^ No. 16. Vanishing Wild Flowers, by Elizabeth G. Britton.
I^oli^-- No. 17. The Tylostomaceae of North America, by V. S. White.
|C\Ol "^ No. 18. Shrubs and Trees of the Southern States, by Dr. J. K. Small.
l^Ol'^ No. 19. The American Species of Lvnnorchis and Piperia, North of
■^ Mexico, by Dr. Per Axel Rydberg.

1 '^ 0 7-No. 20. The Morphology and Physiology of the Seedlings of Arisaema

ttiphyllum and A. Dracontiuni, by R. J. Rennert.
I '^O LNo. 21. Two new Western Mosses, by R. S. Williams.
iGj2.,No. 22. Studies on the Rocky Mountain Flora — VII, by Dr. Per Axel

Rydberg.
/7o3^No. 23. Studies on the Rocky Mountain Flora — VIII, by Dr. Per Axel

Rydberg.
/ ^o/UNo. 24. The Nidulariaceae of North America, by V. S. White.
/ ^t^A-^^- -5- Notes on American Hepaticae, by Marshall A. Howe.

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SYMBIOSIS AND SAPROPHYTISM

BY DANIEL TREMBLY MacDOUGAL

NEW YORK
1899

[Reprinted from the Bulletin of the Torrey Botanical Club. 26 : 511-530. 16 Oct., 1899 ]

Symbiosis and Saprophytism

By Daniel Trembly MacDougal
[Plates 367-369]

General Discussion

It is customary to desii^nate all chlorophylless seed-forming
species which have no nutritive connection with other vascular
plants as saprophytes, or more exactly, holosaprophytes (allotropic
or heterotropic forms according to Pfeffer's classification), and
others of similar physiological tendencies as hemisaprophytes
(mixotropic forms), without regard to the nutritive unions formed
by the roots or absorbing organs in mycorhizas, tubercles and
other associations. It is obvious that the term saprophyte, or
holosaprophyte should be applied only to those species which de-
rive their supply of food from organic products directly without the
intervention of the activity of chlorophyll, and unaided by other
organisms. In this sense, and it seems to the author to be the
only meaning admissible, the holosaprophytes include numerous
bacteria and fungi, but so far as present investigations show, only
one seed-forming species, Wiillsddaegdia aphylla : Cephalaiitliera
Orcgana was erroneously grouped in this class in a previous
publication.*

* An abstract of this discussion of terms was read before the Society for Plant Phys-
iology and Morphology, at New York, Dec. 28, 1898, and was published in American
Naturalist, for March, 1899 (lo). See also note on same in Science for Feb. 3, 1899,
and Botanical Gazette for Feb. and Sept., 1899.

[Issued October 16.] ( 511 )

512 MacDougal : Symbiosis and Sai'Ropiivtism

As a consequence of the above limitations all species furnished
with mycorhizas, tubercles, or which enter into direct mechanical
or nutritive associations must be classed as symbionts, or if it is de-
sirable to maintain connection with existing literature dealing with
these forms it would be permissible to refer to them as symbiotic
saprophytes, although such designation must be regarded as ten-
tative, and justified by expediency only (i i).

It is a matter of common knowledge that seedlings are holo-
saprophytic in the stage in which they are wholly dependent upon
the reserve material of the seed or fruit, and during the whole
period previous to the formation of chlorophyll in general. This
period is practically obliterated in those species in which chloro-
phyll is formed in the seed. On the other hand, the development
of this tendency has been twofold. The increase of the capacity
for the absorption of organic products has played an important
part in the reduction of certain seeds to their present minute form,
and again the retention, or extension, of this capacity throughout
a greater or less portion of the life of the sporophyte has resulted
in varying stages of true saprophytism, complete in one species of
the higher plants only. The duration of the holosaprophytic stage
shows very wide variations in different species. In certain arums
it extends over two years under natural conditions, and may be
extended by cultural methods so that the seedling ma)' not form
chlorophyll until the third or perhaps even the fourth year of ex-
istence, according to experiments now in progress. Only those
species which show a marked capacity for the absorption and use
of organic products during the greater part of the life of the sporo-
phyte should be classed as hemisaprophytes. The hemisapro-
phytes would consist chiefly of the carnivorous plants. The
greater majority of the species now included in this category are
in fact more or less symbiotic by means of mycorhizas, tubercles
or other nutritive associations.

This paper describes an extension of the investigations detailed
in a previous paper (i i), and an effort has been made to determine
whether such associations constitute a single physiological type, or
whether several types of nutritive adjustment arc presented.

MacDougal : Symbiosis and Saprophytism 513

Cephalanthera Oregana Reichenb. (13)

A number of living specimens of this plant were examined in
the field in Washington and Idaho in 1892, and alcoholic material
from this region was obtained in 1899. Besides the notes and
material thus obtained, the author has had the opportunity of in-
specting herbarium material representing the entire known range
of the species, from middle California northward into British
Columbia, west of the main continental divide.

The plant consists of an upright subterranean rhizome 5 to 40
cm. in length, from the internodes of which arise the adventitious
roots. The internodes are provided with short sheathing scales.
The aerial stem is slender, waxy white, 20 to 50 cm. long, and
bears short sheathing leaves which are wholly devoid of chloro-
phyll. The flowers form a dense terminal raceme and perhaps
agree with those of other members of the genus in being self-fer-
tilizing (Plate 367, Fig. i).

The seedling has not been observed. The rhizome is peren-
nial, and the stumps of two or three old aerial stems may be seen
adhering to the most recently formed internodes. Although re-
ported from open meadows by some collectors, the author has met
it only in the deep humus of coniferous forests. Its deep penetra-
tion of the loose substratum, which is generally woven together by
the interlacing roots of neighboring trees, makes the collection of
the entire plant veiy difficult and as a consequence the greater
number of herbarium specimens are lacking the root system, and
on none of these has the writer found the mycorhizal rootlets
described below.

The Roots

The roots are wavy cylindrical organs 1.5 to 3 mm. in diam-
eter, and from 5 to 12 cm. in length. One to four arise at each
internode of the premorse rhizome, and penetrate the substratum
at an angle of forty-five degrees with the horizontal. Mature
organs have a yellowish-brown appearance due to the decay of the
outer epidermal wall. A many-layered root-cap sheathes the tip
for a distance of a millimeter, and shows a fairly normal structure.
The epidermal cells are rectangular in surface section, with the
radial walls separated to allow a slight outward convexity of the

514 MacDougal: Symbiosis and Saproph\tism

outer surface, which may in some instances assume a papillose
form, or may be extended to form typical root-hairs. The latter,
and all of the epidermal elements, are rich in protoplasm. The
outer walls of the epidermis are covered more or less thickly with
crystals of calcium. The sub-epidermal layer is hardly differen-
tiated, and consists of muriform elements rich in protoplasm, and
devoid of reserve material. The cortex is composed of ovoid,
cylindrical or globose cells, the outer and inner layers of which
are composed of elements slightly smaller than those of the middle
region. The middle and inner regions are heavily loaded with
starch, especially in the older or basal portions, and the cells of
the middle region show a tendency to elongation in the radius of
the root. Raphide cells are indifferently distributed throughout
the cortical tissues, but perhaps more abundantly in the outer
layers. Two types of roots which do not differ greatly in outward
aspect may be distinguished ; a fibrous form, and a fleshy form
about twice the diameter of the first. External to the endodermis
the two are similar except in the amount of development of the
cortex. In the development of the thicker storage organs, the
central parenchyma becomes slightly lignified and sclerotized, the
xylem bundles increase from six to seven or eight, the pericycle
shows two or three layers underneath the endodermis, and this
sheath is composed of slightly thickened and pitted elements with
no special passage cells. The xylem and the endodermis become
lignified (Plate 369, Fig. 2). In the development of the fibrous
roots the xylem undergoes such centripetal development, and
lateral fusion that it changes from a hexarch to a tetrach. The
heavily lignified xylem and the weakly developed central paren-
chyma form a four-rayed star, with the phloem lodged in the
sinuses (Plate 369, Fig. 2). The endodermis is heavily thickened
in the regions external to the phloem regions and consists of thin-
walled passage cells opposite the xylem. The xylem shows great
degeneration in both types of roots, and consists of scalariform
vessels and elongated tracheids with transverse pits. The phloem
exhibits no ereat deviation from the normal, and consists of narrow
sieve tubes, companion cells and some parenchyma. A layer of
elongated elements containing protoplasm and apparently adapted
to the conduction of proteids lies immediately outside the phloem

MacDougal : Symbiosis and Saprophytism 515

proper. The two types of roots show a remarkable resemblance
in stelar alterations to those of the two types in Wnllschlacgelia
apJiylla as described by Johow (7, p. 427), though not so distinctly
separated by external characters.

The older roots often assume a dark-brown color over certain
regions 6-10 cm. in length, due to the presence of an ectotropic
fungus, which forms a permanent inycelium in three or four of the
outer layers of the cortex (see page 523). The hyphae are heavy
walled and septate. The inner branches form large vesicles which
occupy the greater part of the cells of the medio-cortex. The outer
branches of the mycelium pass through the short root-hairs into
the soil. The region inhabited by the fungus gives rise to a few
rootlets which assume the form of lozenge shaped branches with
a length not greater than 3 cm. These branches arise endogen-
ously, are brownish to within a few millimeters of the tips and are
furnished with a well developed root cap. The mycorhizal fun-
gus adv^ances toward the tips as in the branches of the Coral-
lorhisa, and the entire structure shows an arrangement generally
similar to the coralloid branches of that plant.

In the half dozen perfect specimens in the hands of the writer
the branches arise only from regions inhabited by the fungus and
it is conjectured that the presence of this organism may act as a
stimulant in setting up such action. The fungus is found in por-

FlG. I. Mycorhizal portions of roots of Cepkalanthera.

tions of the root-system 50 to 60 cm. below the surface of the
soil, and as these organs are very brittle, especially the mycorhizal
portions they are generally broken off in collection. The absence
of chlorophyll and mycorhizas from numerous specimens sent in
by the most careful collectors led the writer to announce that the

516 MacDouoal : Symbiosis and Saprophvtism

species was a true saprophyte (ii). Recently, however, Mr. O.
D. Allen, of Ashford, Washington, by special request, dug up
some entire specimens with the result that the mycorhizas were
preserved, and are shown in Figure i.

This species is noted as a parasite in systematic texts, but the
author has failed to find any direct evidence of its nutritive connec-
tion with any other seed-forming species, either by his observa-
tions or from the reports of collectors.

The structural features of the roots agree with those of other
mycorhizal forms in the reduction of the absorbing surfaces, the
lack of spiral and annular vessels, the formation of tracheids with
narrow transverse pits and the radial elongation of the mediocor-
tical region. This last named feature has been supposed to be due
to the influence of the fungus in mycorhizal roots, but Groom has
shown that it may occur in portions of the root unoccupied by the
symbiont. Its presence here, in portions of the root not occupied
by the fungus, indicates that it is due to the necessities of absorp-
tion and use of organic food perhaps rather than to the stimulation
of the presence of a symbiotic organism.

The Rhizome

The rhizome is an upright stem, slightly compressed, with in-
ternodes i to 2 cm. long, from the terminal internodes of which
annually arise one or two flower scapes.

The epidermis is composed of elongated muriform elements
with the outer walls slightly convex. No transpiratory openings
are to be found. The cortex is ten to fifteen layers in thickness,
the cells are ovoid, cylindrical, pitted and separated by spare inter-
cellular spaces. The starch stored here during the resting season
is usually exhausted in the formation of the inflorescence. The
pericycle is two or three layers in thickness, heavily sclerotized
and lignified and is interrupted in places by thin-walled elements.
The crowded xylem ring contains a large number of scalariform
ducts, and some imperfectly differentiated spiral and annular ves-
sels, all deeply lignified. The phloem consists of cambiform ele-
ments with spare protoplasmic content. The interfascicular paren-
chyma shows collenchymatous thickenings in places, and numbers
of elongated conducting cells with unlignificd walls are placed
near the vessels (Plate 369, Fig. 4).

MacDougal: Symbiosis and Saprophytism 517

The rhizome exhibits no marked or general degeneration, unless
the condition of the phloem could be interpreted in this manner.
The xylem is amply developed for the conduction of the supply
of water necessary for the aerial shoot, and the elongated elements
near the vessels appear to be suitable for the transmission for pro-
teinaceous compounds, a function served by certain cells immedi-
ately internal to the endodermis in the root. The rhizome of
Ccplialanthcra differs from similar members in symbiotic sapro-
phytes in not being provided with organs for the excretion of water.

The Inflorescence

The flowering stem is composed of flattened internodes 2 to 5
cm. long, bearing short sheathing leaves, and a terminal raceme
of flowers. The epidermis is composed of flattened cylindrical
elements with oblique ends, resembling tracheids in general form,
and with the inner and radial walls dotted with numerous perfora-
tions. Irregular masses, globules and networks of a yellowish
brown substance are abundant in the epidermal and cortical cells
of material preserved in alcohol. Ordinary chemical tests are
without definite reaction, though Raspail's reagent gave slight in-
dication of proteids in these masses. The epidermis is totally
devoid of transpiratory openings.

The cortex is eight to twelve layers in thickness, and consists
of elongated cylindrical elements with no intercellular spaces. It
appears to serve as a tissue for the storage of water. Internal to
the cortex is a sclerenchyma sheath, in the inner margin of which
lies a circle of 32 to 40 bundles, and centrally placed is a second
ring of six to eight bundles, which are fairly identical with those
of the rhizome. Each bundle is enclosed in its own schleren-
chyma sheath. The sheath and the xylem are heavily lignified.
No marked degeneration is to be seen outside of the lack of chlo-
rophyll, the transpiratory organs, and the intercellular spaces of
the cortex. The heavy sclerenchyma sheath is a feature of the
aerial stems of the symbiotic saprophytes. The xylem shows a
very typical development.

The Leaves
The leaves are reduced to sheathing bracts, and are destitute of
chlorophyll, as shown by an examination of the alcoholic extract

518 MacDougal: Symbiosis and Saprophytism

with the micro-spectroscope. The free portion representing the
lamina is i to 2 cm. in length, and the outer dorsal surface is pro-
vided with stomata, the guard cells of which are motile and con-
tain starch (Plate 369, Fig. 5). The possibility that the guard
cells may contain a small amount of chlorophyll is not excluded,
though it could not be detected by ordinary tests in the specimens
examined. The epidermis is composed of muriform cells with the
outer walls convex and not cuticularized. The mesophyll consists
of two layers of irregularly globoid elements rich in protoplasm,
and separated by large air-spaces. A third layer of thin -walled
elements, cylindrical in form are to be found lateral to the simple
fibrovascular bundles. This tissue is devoid of protoplasm and
may serve for the storage of water. The sclerenchyma sheath is
incomplete at the point of contact with this supposed storage tis-
sue, thus permitting the ready passage of water between the xylem
and the thin-walled cells.

The degeneration of the leaf is seen to consist in the loss of
chlorophyll, the lack of differentiation of the mesophyll, and the
reduction of the surface. This degeneration has been accompanied
by the development of a tissue for the storage of water, and by the
retention of the stomata. Ccplialanthcra is to be added to the list
of chlorophylless species furnished with motile stomata.

General Considerations
It is evident that the mycorhiza of Ccplialanthcra is to be con-
sidered as adventitious or accidental in its occurrence. In a large
number of ectotropic forms the fungus gains entrance to the un-
derground organ very early in its development, and then keeps
pace with its growth. In this instance, however, it is found only
in certain regions, and might be mistaken for a parasite were it not
for its characteristic vesicles or organs of interchange. The com-
paratively small area of the mycorhizal structures suggests that
Ceplialanthera is capable of absorbing largely from the humous
products independently.

Calypso bulhosa (L.) Oakcs
The author has called attention to the occurrence of adventi-
tious mycorhiza in Calypso {\ i), and cited Lundstrom's description

MacDougal : Symbiosis and Saprophytism 519

of these formations, which have been found by him and others in
Europe and America. Since that paper was sent to the press a
shipment of plants has been received, which had been collected
in northwestern United States, and a number of the specimens
showed the coralloid mycorhiza. A careful reexamination has
been made of these structures in the light of the generalizations
drawn from previous material, and the results are presented below.
The subterranean stem of Calypso consists of an ovoid taper-
ing corm 1.5 to 2 cm. in length, comprising two or three inter-"
nodes. The single ovate, or ovate-cordate leaf is terminal, while
the inflorescence arises from the first node below. The plant is
reproduced vegetatively by a short offset of such reduced length
that the new corm formed from its apical internodes stands upright
in contact with the old corm (Plate 367, Fig. 6). The iow short
roots arising from the base of the corm are mycorhizal as de-
scribed in the previous paper.

Stem-mvcorhiza

The old or spent corms of the preceding season's activity may
give rise to offsets from the basal internodes, and these may de-
velop into coralloid structures by the repeated branching due to
the development of all the buds, as in Aplcctnun. The general
anatomy of the coralloid formations is too nearly like that of the
stem-mycorhizas of Aplcctrum to warrant description here.

The fungus is seen to be a loose skein of hyphae in the three
or four outer layers of the cortex passing outwardly through the
thin-walled epidermal cells into the substratum, and do not,' so far
as observations go, traverse through the nodal trichomes. In this
respect Calypso differs from other coralloid plants. Occasionally
small globular or ovoid structures resembling sporangioles are to
be found terminating the branches of the hyphae in the outer cor-
tex. The three or four layers of the medio-cortex are filled with
dense masses of interwoven hyphae. The hyphae form irregu-
larly swollen branches upon entering the cells of this region, and
one or two of these branches near the nucleus of the cortical cell
expands into a vesicle, which in turn gives off a large number of
branches nearly filling the cell. The hyphae are unseptate and
have definite heavy walls. The form, irregular outlines and in-

520 MacDougal : Symbiosis and Sapkofiivtism

definite membranes of the hyphac in the medio-cortex led Lund-
strom to believe that the fungus was plasmodial in its earlier
stages (9). This appearance is heightened by the adhesion of the
cytoplasm of the cortical cell to the hyphae. The inner cortex is
free from hyphae, and contains starch in abundance, especially in
the apical region. Starch is also present in the outer cortex, but
quickly disappears from the cells invaded by the hyphae. The
nuclei of the outer cortical cells are nearly normal, but those of
the medio-cortex, occupied by the fungus, are hyperchromatic,
distorted, and in some instances undergo fragmentation, as in
Pcramiinii ( 1 1 ). The stele is not differentiated into xylem and
phloem, and consists chiefly of cylindrical elements rich in pro-
toplasm, not differing greatly from plerome. The endodermis
cannot be made out. The fungus of the coralloid structure and
that of the roots are quite similar, but their identity is not es-
tablished.

A comparison of the specimens which have come under inspec-
tion makes it apparent that the tendency to form coralloid myco-
rhizas may play as important a part in Calypso as in Aplectriini.
The coralloid stems were small in some instances, and attached to
the base of corms two years old, while in others their bulk was
greater than that of the corm of the previous season to which they
were attached. The extreme development was found in one
specimen in which the coralloid structure was very large, and the
old corm to which it was attached was shrunken to half its orig-
inal size, but was still sound and normal (Plate 367, Fig. 8). A
second offset had sprung from the node nearest the apex of the
corm. This offset was about a centimeter in length and bore two
roots at the first node while the three upper internodes had begun
to swell in the formation of a new corm. The terminal portion
bore a rounded cordate leaf and a flower bud. The members of
this specimen named in order from the basal end were : stem-
mycorhiza, spent corm, offset, roots, developing corm, flower bud
and leaf. A comparison of the specimens at hand showed a fairly
well established correspondence between variation in the outline of
the leaf and the development of the stem-mycorhiza. Such varia-
tion might be due to the increased capacity of the plants furnished
with coralloid structures for the absorption of humus products.

MacDougal : Symbiosis and Saprophytism 521

The v'ariation in the leaf is also accompanied by increase in the
length of the offsets, the decrease of the capacity of the storage
organs, and the diminution in the number of the roots. The coral-
loid offsets of Calypso have not been seen to give rise to new plants
as in Aplcctniiii.

The very great divergence of individuals from the type in leaf
and flower characters in this species has been a subject of remark
among systematists for many years, and Mr. Heller has recently
raised one of the most pronounced forms to specific rank (4).
Whether such variations of the shoot are directly correlated witii
the development of the mycorhizal structures or not can not be
definitely stated. The matter may be determined only by the
careful examination of specimens in the field throughout the habitat
of the species.

Corallokhiza Arizonica Wats.

(No. 94 of collection of 1898 by MacDougal.)

The general morphology, and occurrence of the fungus of the
coralloid branches of Corallorhi::a have been somewhat fully de-
scribed by Schacht, Irmisch, Reinke and others, chief attention
having been paid to Corallorldza Corallorliiza{\^^ Karst (C innatd).
The results at hand, however, were obtained quite early in the
history of investigation upon this subject, and certain details now
known to be of great interest were not touched upon in these
earlier researches. It was deemed advisable, therefore, to make a
reexamination of the mycorhizal structures of another representa-
tive of the genus, with especial attention to the physiological rela-
tions of the members of the symbiotic union. Jennings and
Hanna (6) have recently published a short paper on C. innata in
which it is stated that the symbiotic fungus is a " hymenomycete
and commonly an argaric." Clitocybc iiifundibiiliformis Sch., was
found attached to the coralloid formations in one instance, and
'' Hystcraiigiuin stoloiiifenivi of Tu/asne" in another in a manner
indicative of the identity of these species with the symbiotic fungus.

CorallorJiiza Arizonica is a native of the upper part of the tran-
sition zone and the Canadian zone in the Rocky Mountains. The
subterranean portion of the plant consists of a dense mass of coral-
loid stems which lie as deep as 20 cm. below the surface, and from

522 MacDougal: Symbiosis and Saprophytism

which extends upwardly a premorse rhizome lo to 15 cm. in
length. The thick aerial stem readies a height of i 5 to 25 cm.,
is sheathed by membranous leaves, and terminated by a strongly
developed racemose inflorescence. The leaves and stem are irreg-
ularly colored with blotches of reddish, purplish and brownish
tints, and appear to be wholly free from chlorophyll. Specimens
with rudimentary aerial stems and others in bloom were collected
by the writer on Mormon mountain, and on the San Francisco
mountain in Arizona in 1891 and 1898, and preserved in alcohol.
The results described below are based upon this material.

The coralloid mvcorhiza

The mycorhiza of this plant is a dense mass of club-shaped
branches arising from the upright underground rhizome, on which
the true roots are to be seen as minute papillae. The germination
of the seed has not been observ^ed, and it is impossible to say
whether the primary roots are developed or not, or at what stage
the symbiotic fungus invades the offsets which constitute the coral-
loid mass. Reinke figures a young plant, probably of the second
year's growth, in which the underground member consists of a
coralloid stem only (14). The external anatomy and method of
branching need no further description in this species. The rhizome
attains a thickness of 4 to 6 mm., with internodes 2 to 4 mm. in
length.

The phloem consists of two or three layers of prosenchymatous
cells with yellowish thickened walls and slimy contents, most
nearly like companion cells. The phloem forms two crescents with
the tips nearly touching with the 3 to 5 xylem bundles lying inter-
nally, or the phloem may form a complete ring enclosing the
xylem. The xylem consists almost entirely of scalariform vessels
and one or two tracheids in which the perforations are oval and
elongated obliquely. The central parenchyma is made up of short
cylindrical cells often richly loaded with starch. The pericycle is
present as one or two layers of cambiform cells, and the endo-
derniis forms a sheath of flattened cylindrical elements. External
to the stele is a cylinder of cortical tissue 10 to 15 layers in thick-
ness composed of ovoid or globoid elements, with large intercel-
lular spaces, and containing starch during the resting period. The

MacDougal : Symbiosis and Saprophytism 523

medio-cortex is i 5 to 20 layers in thickness and is composed of
cells with the radial diameter twice the axial. The outer cortex
consists of 3 to 5 layers of very thin-walled elements with the
tangential diameter greater than the radial. Both the medio-
cortex and the outer cortex are provided with intercellular spaces.
The epidermis is composed of flattened cells, irregular in outline,
with the lateral and inner walls pitted and the outer wall slightly
thickened and brownish in color. It is furnished with a large
number of stomata, with the motile guard cells of cfescentic form
(Plate 368, Figs. 5 and 6) containing starch. These and the large in-
tercellular spaces constitute a very efficient aerating sy.stem, and
makes the coralloid structure independent of the aerial shoot in
transpiration, and at the same time allows free access of atmos-
pheric oxygen. The apices of the rudimentary sheathing leaves
are soon converted into a number of blackish shreds and their
bases persist as wedge-shaped rudiments with no distinct function.
Clumps of large trichomes resembling root-hairs arise from papillae
which are infra-axillarv to the leaves.

The fungus obtains access to the coralloid offset at quite an
early stage of its existence and constantly grows toward the elon-
gating .apex forming convolutions of pale gray shining hyphae
with numerous septae in the outer cortex. The growth of the
hyphae keeps pace with the offset in its slow growth, and they
extend forward to the shoulder of the blunt tip of each branch,
curving inward at this place toward the embryonic tissue. The
hyphae in the outer cortex remain active even in old mycorhiza
and may be designated as forming the permanent mycelium. The
permanent mycelium is thus in the shape of a sub-epidermal cylin-
der, and when new branches are formed the sheathing cylinder ot
the fungus is continued out in it. Branches from the apical por-
tion of the permanent mycelium penetrate the medio-cortex while
it is still in an undifferentiated condition, and these branches grow
and ramify with the development of the cortical cells until the
latter are almost filled with their dense convolutions. For some
unknown reason the development of the hyphal branches is great-
est immediately internal to the nodal trichomes in the medio-
cortex. The hyphal branches are generally cylindrical but occa-
sionally portions become swollen to twice the normal diameter,

524 MacDougal : SvMinosis and Saprophvtism

but no enlargements constituting sporangioles, vesicles or other
organs of interchange are to be seen. The cortex of the younger
portions of the coralloid structures is filled with starch granules
which are slowly corroded by the action of the developing hyphae.
Cells in which the hyphae have made many convolutions still
contain some starch, but it finally disappears. The hyphae in the
medio-cortex a distance from the tip are yellowish and collapsed,
but no solid bodies are to be seen as a result of the liberation of
their disintegration products in the cortical cells. The proto-
plasm of the latter is w^ell spent but normal, and the nuclei are
normal and active. The permanent mycelium sends out external
branches through the trichomes into the soil. The permanent
mycelium is, therefore, in the form of a sheathing cylinder with
numbers of branches opposite each other extending out into the
substratum* and into the cortex. It is to be pointed out in this con-
nection that the numerous statements to the effect that the fungus
gains access to the interior of the coralloid structure through
the trichomes are obviously incorrect. Entrance to the offset in
the initial stage of the formation of the coralloid branches is per-
haps made in this way, but once inside the branch the permanent
mycelium is found which keeps pace with growth of the cortex and
sends branches outwardly through the trichomes. The continued
and repeated entrance of the fungus through the trichomes is an
assumption only, and is based on the necessities of the theory of
mycorhizas as fungus traps rather than on the actual facts.

The chemotropic reactions of the fungus as shown by its method
of extension are of great interest. The permanent mycelium tra-
verses the coralloid branches in the layers of cortical tissue first
differentiated. The tips of the hyphal branches are attracted out
through the trichomes, presumably by atmospheric oxygen, or by
the humus products, which would increase in concentration from
the base of the epidermal cells to the apices of the trichomes. The
attraction of the branches into themedio-cortical cells nnist be due
to a carbohydrate, rather than a nuclear product, since it is quite
noticeable that all convolutions of the hyphae are made in regions
of the cell some distance from tlic nucleus. The tip of a hypha
may pass within its own diameter of the nucleus of the cortical
cell with mutual indifference, and only in a small number of in-

MacDougal : Symbiosis and Saprophytism 525

stances does the presence of the fungus affect the nucleus. Ex-
cretions from the hyphac cause some distortion of a few nuclei,
which are also hyperchromatic. The hyphae may be traced around
the cell in several circuits. Penetration of the wall and entrance
into a neighboring cell is not made until the supply of starch is
nearly exhausted, and the solution in the contiguous cell would
form a stronger chemotropic stimulus. The portion of the hypha
in the wall appears but half the normal diameter of the typical fila-
ment, and is nearly colorless even in old formations.

The ascending rhizome

The ascending rhizome consists of four or five napiform seg-
ments, each consisting of two or three internodes and representing
one season's growth. As each segment is formed it gives rise to
an inflorescence from an apical node. Later in the season it forms
offsets which reproduce the premorse rhizomes with their coral-
loid branches. As a consequence of this mode of growth, a dozen
plants may be found adherent in a colony, with the coralloid ni}'-
corhizas closely crowded in a huge clump.

The epidermis of the rhizome consists of very irregular ele-
ments, some of which contain starch at all times. Numerous
hyathodes with a central oval or oblong central cell filled with a
dense mass of yellowish brown secretion, surrounded by six or
seven radially arranged elements are to be seen (Plate 368, Fig.
4). The cortex shows a very copious development, consisting of
ovoid or globoid elements with ample air-spaces, and embracing a
large number of mucilage cells. This member is therefore fur-
nished with a very efficient mechanism for the extrusion of water :
an important provision in an organ devoted to the condensation
of carbohydrates of soluble carbohydrates to starch, and this pro-
cess may continue irrespective of the presence of the aerial shoot.
The peri cycle forms a dense heavy ring of 7 to 10 layers rich in
proteids. The stele in general exhibits a degree of degeneration
corresponding with that of the coralloid branches. Its parenchy-
matous elements as well as the cortex are loaded with starch.

The Inflorescence axis
The epidermis consists of flattened cylindrical cells with oblique
ends, are rich in protoplasm, and entirely devoid of stomatal open-

526 MacDougal : Symbiosis and Saprophytism

ings. The lo to 15 layers of cortical tissues are furnished with
very large intercellular spaces, which may denote an epidermal
transpiration of some importance, though no special adaptation for
this purpose could be detected. Crystal cells are scattered through-
out the cortex and appear even in the epidermis. The pericycle
is composed of several layers of elongated cells with thickened
walls and yellowish brown contents. The bundles are scattered
in the stele with the xylem and phloem radially arranged. Then'
degeneration is fairly uniform with that of the rhizome and its
branches. The leaves are sheathing, destitute of stomata, and
show no differentiation of tissue for food-formation, or transpira-
tion, except that the globular parenchyma shows great intercellu-
lar spaces. As noted above they are free from chlorophyll
C. Arizonica differs from C. Corallorhiza (C. innnta) in the formation
of a bulky premorse rhizome furnished with stomata, the total lack
of chlorophyll in the shoot, with absence of stomata, and in the
greater degeneration of the stele. The differentiation of the fungal
symbiont into a permanent mycelium with short-lived and external
and internal branches seems to be a mycorhizal character described
here for the first time, though it is present in C. Corallorliiza and
other species. The hyphae in the coralloid formation of C. Co-
rallorldza, C. uiultiflora and others are applied to the nuclei in the
cortical cell while in C. Arizonica this is not the case, indicating a
difference of chemotropic reaction of the fungus in the two in-
stances.

Relations of the mycorhizal components

The fungus in the coralloid formations of CorallorJiiza draws its
nourishment from two sources ; .from the humus products of the
soil and from the carbohydrates in the cortex. The material thus
obtained is used in the construction of extensions of the mycelium
and its branches. With the growth and progression of the my-
corhiza, the older internal branches of the mycelium which have
formed dense masses in the medio-cortex undergo disintegration
and the products thus liberated may be drawn in two directions :
toward the apex of the mycorhiza and toward the premorse rhi-
zome. On arrival at both places the surplus matei'ial is converted
into starch. The starch of the rhizome is used in the construe-

MacDougal : Symbiosis and Saprophytism 527

tion of the reproductive and other branches. The starch in the
apex of the coralloid mycorhiza is used in the construction of em-
bryonic tissue and a portion of it remains in the medio-cortex and
becomes available to the fungus as a highly advantageous food.
Janse and others have upheld the theory that endotropic my-
corhizas are similar in physiological value to leguminous tuber-
cles (ii), but the only actual proof adduced in favor of this view
is the evidence obtained by Nobbe and Hiltner from experiments
with Podocarpiis ( 1 2). That endotropic fungi may cooperate in
the fixation of free nitrogen in the roots of plants in which they
occupy only a portion of the absorbing system is readily ad-
missible and may be considered as proven. Such an explanation
is wholly inadequate to account for the arrangement of the my-
corhizal components and transpiratory structures in Corallorhiza^
however, on purely anatomical grounds. The underground mem-
bers of this genus are furnished with a complete sub-epidermal
sheath of mycelium, which fills every cell of the outer cortex in
two or three layers, except a minute area at the tip of the coral-
loid branch, and usually the 10 to 15 layers of the medio-cortex.
It is obviously impossible for the CorallorJiiza to absorb substances
from the soil except through and by the agency of the fungus.
The fungus may be capable of accomplishing the fixation of free
nitrogen, but that it is not its sole, or its major function in the
symbiosis, since all of the food-material of the association must
pass through its hyphae ; a statement equally true of such forms
of ectotropic mycorhiza as those of Ptcrospora, Monotropa, etc.
The higher plant affords a lodgment for the fungus, from which it
sends out absorbent and reproductive branches. Food-material
taken in by the fungus is yielded to the higher plant and consti-
tutes its sole supply. To this extent the higher plant is parasitic
upon the fungus. But the higher plant accomplishes transforma-
tions of chemical energy in the food thus obtained of which the
fungus is incapable and yields the elaborated product in an ad-
vantageous form in the apex of the mycorhiza, where it serves as
a food for the advancing mycelium. The higher plant is, there-
fore, not a fungus-trap pure and simple, as the association is of
^reat mutual advantage.

The principal conclusions which may be drawn from the facts

528 MacDougal : Symbiosis and Saprophytism

adduced in the foregoing paper may be briefly stated as follows :

I. The term saprophyte should be appHed to those species
only which derive their food-supply from organic products, un-
aided by chlorophyll, and without the intervention of other organ-
isms. The true saprophytes therefore include numerous bacteria,
fungi and but one seed-forming species — Wullsclilaegelia apJiylla.
The saprophytic capacity of the seedling has been extended to
cover periods of varying length in the life of the hemi-saprophytes
and with symbiosis has reacted to diminish the tendency to store
reserve material in seeds.

II. The degenerations of the true saprophytes are generally
parallel to those of mycorhizal forms.

III. Ccplialantlicra Oregana and Corallorliiza Arizonica are to
be added to the list of chlorophylless plants furnished with stomata.

IV. The offsets of Calypso are occa.sionally converted into
coralloid mycorhizas as in Aplectruiii. The stele of such structures
is not differentiated into xylem and phloem. The occurrence of
the coralloid mycorhiza is accompanied by variations in the form
of the leaves, and"o£ the decrease in the capacity of the storage
organs in the specimens examined.

V. Corallorliiza Arizonica exhibits greater development of the
symbiotic adaptation than C. Corallorliiza. The stele is quite
primitive throughout, chlorophyll islacking, and stomata are pres-
ent on the coralloid branches only. The epidermis of the pre-
morse rhizome is furnished wath hyathodes. The aerial shoot
is furnished with large intercellular spaces, but may carry
on epidermal transpiration only. The fungus in the coral-
loid structures consists of a permanent mycelium, with external
and internal branches ; the former are organs of absorption and
reproduction, the latter are organs of interchange between the
members of the symbiosis. The higher plant affords lodgment
for the fungus and carries on chemical transformations the prod-
ucts of which are available to the fungus. The latter absorbs
and yields to the higher plant in a more or less complex form the
products of the humous soil.

VI. All endotropic mycorhizas do not conform to a single phys-
iological type. The theory of Janse that endotropic fungi are
negatively chemotropic to oxygen, and bear the same relation to

MacDougal : Symbiosis and Saprophytism 529

the higher plant as the organism of the leguminous tubercle, is not
capable of general application. Such relation has been proven be-
tween Podocarpus and the peronosporous fungus of its mycorhiza
only.

VII. Two types of endotropic mycorhizas may be distin-
guished ; one adapted for nitrogen fixation, and a second for the
absorption and modification — perhaps oxidation — of the soil prod-
ucts before liberation in the tissues of the higher plant. The ex-
tension of information will doubtless result in the further division
of the second type.

Literature to which Referenxe is made.

1. Bonnier and Mangin : Recherches sur la respiration des tissues
sans chlorophyll. Ann. Sc. Nat. VI. i8 : 203. 1884.

2. Engler u. Prantl : Die naturl. Pflanzenf., 2 : 11. .

3. Pf effer : Pflanzenphysiol. i : 349. 1897.

4. Heller : New plants from western North America. Bull. Torr.
Bot. Club, 25 : No. 4. April, 1898.

5. Janse : Les endophytes radicants de quelques plantes javanaises.
Ann. d. Jard. d. Buitenzorg. 14: 53. 1896.

6. Jennings and Hanna : Corallorhiza innata R. Br. and its my-
corhiza. Sc. Proc. Roy. Soc. Dublin, N. S. 9: [pages?]. 1899.

7. Johow : Die chlorophyllfreiHe Humusbewohner West Indiens,
biologisch-morphologisch dargestellt. Pringsh. Jahrb. f. wiss. Bot. 16 :
445. 1885.

8. Lory : Sur la respiration et la structure des Orobanches, et
autres plantes vasculaires deporvues de partes vertes. Ann. Sc. Nat.
III. 8: 158. 1847.

9. Lundstrom : Einige Beobachtung ueber Calypso borealis. Bot.
Centralb. 38: 697. 1889.

10. MacDougal: Symbiosis and Saprophytism. Am. Nat. 33:
210. 1899.

11. MacDougal: Symbiotic saprophytism. Ann. Bot. I : i. 1899.

1 2 . Nobbe u. Hiltner : Die endotropische Mycorhiza von Podo-
carpus, und ihre physiologische Bedeutung. Landw. Versuchssta. 51 :
241. 1898.

13. Reichenbach : Orchidiographische Beitrage. Linnaea, 41 : 53.
1877.

0/

530 MacDougal : Symbiosis and Sapkophytism

14. Reinke : Zur Kenntniss des Rhizomes von Corallorhiza und
Epipogon. Bot. Zeit. 56: 145. 1873.

15. Thomas: The genus C^ra//W«'2<2. Bot. Gaz. 18 : 166. 1893.

Hxplanatiou of Plates

Plate 367. (l) Entire specimen of Cephalanthera Oi-egana Reichenb : A, base
of aerial stem. (2) Premorse x\^\zo\nQ o'i Corallorhiza Arizouica : A, X^xxmx^zS. bud;
B, B, B, B, scars of preceding buds ; C and D, offsets from which arise coralloid
branches. (3, 4 and 5) Coralloid branches of Corallorhiza Arizonica. (6) Typical
specimen of Calypso borealis : A, young corm ; B, corm of previous season's forma-
tion. (7) Aberrant specimen of Calypso : A, o\A. corm with coralloid branch. (8)
Widely aberrant form of Calypso : A, young corm ; B, old corm with large coralloid
branch, C.

Plate 368. Corallorhiza Arizonica Wats, (i) Longitudinal section of tip of old
coralloid mycorhiza : a, a, epidermis ; 6, />, medio-cortex, containing disintegrating
branches of fungus ; <r, stele ; m, m, permanent mycelium ; d, d, branches ; e, leaf.
(2) Portion of transverse section of mycorhiza : ^, epidermis ; r, permanent mycelium;
w, internal branches in medio-cortex ; /, inner cortex; b, phloem; 0, xylem. (3)
Cells from cortex: a, a, hyphae of permanent mycelium ; c, masses of hyphae in medio-
cortical cells ; n, n, n, nuclei of cortical cells. (4) Hyathode from premorse rhizome.
(5) Stomata from coralloid branch. (6) Section of epidermis of coralloid branch
showing structure of stoma : e, ^.epidermal cells; a, guard cells; «, nucleus. (7)
Transverse section of aerial stem : e, epidermis ; c, cortex ; (/, sclerenchyma sheath ;
X, X, fibrovascular bundles.

Plate 369. Cephalanthera Origana Reichenb. ( i ) Transverse section of stele
of storage root : /, medulla ; x, x, xylem ; c, endodermis. (2) Transverse section of
stele of fibrous root : v, passage cells of endodermis ; e, thickened cells of endodermis
outside of phloem; s, phloem; z, xylem; c, medulla. (3) Longitudinal section in
stele ; 0, s, vessels ; vi, medulla ; n, endodermis. (4) Portion of transverse section of
rhizome : e, epidermis ; h, cortex ; /, sclerenchyma sheath ; g, fibrovascular bundles
with heavy sheath ; r, parenchyma. (5) Stoma from leaf.

Bull. Torr. Bot. Club.

Pl. 367.

F. K. Deniston, del.

MACDOUGAL ON SYMBIOSIS.

THE HELIOTYPE PRINTING CO., BOSTON.

Bull. Torr. Bot. Club.

Pl. 368.

F. K. Deniston and H. B. Humphrey, del.

MACDOUGAL ON SYMBIOSIS.

THE HELIOTYPS PRINTING CO.. BOSTON.

Bull. Torr. Bot. Club.

Pl. 369

H. B. Humphrey and F. K. Deniston, del.

MACDOUGAL ON SYMBIOSIS.

THE HEU0TYP6 PRINTING CO.. BOSTON.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 2

NEW SPECIES FROM THE WESTERN
UNITED STATES

BY P. A. RYDBERG

NEW YORK
1899

[Reprinted from the Bulletin of the Torrey Botanical Club. 26 : 5U-5u6. 16 Oct., 1899.]

New Species from the Western United States

By 1*. A. Ryduerg

Juncus Suksdorfii

Stem about 3 dm. high, strict, light green, 2-3 mm. in diame-
ter ; leaves terete or slightly flattened, distinctly septate ; the basal
ones short ; stem leaves, except the upper ones, often 3 dm. long
all with a conspicuous, scarious sheath ; heads in a contracted pan-
icle, brown and shining, 5-8-flowered ; bracts ovate, cuspidate-
acuminate ; perianth segments subequal, about 4 mm. long, nar-
rowly lanceolate, acute or acuminate ; stamens 6 ; anthers longer
than the filaments ; style long-exserted ; capsule dark brown and
shining, oblong, acuminate, 3-angled.

Dr. Watson has labeled this /uticus Nevadensis var., to which it
may be nearest related if the structure of the flower is taken in
consideration. It is different in habit, however, being much stouter,
having more numerous and larger heads, and longer leaves.

Washington: Falcon Valley, iSS^,, Suksdorf, 21"/; 1885, (56*0;
Spangle, Spokane Co., 1884, ^6^ (all in Gray Herbarium).

Allium Neo-Mexicanum

Bulb oblong, membranaceous, crowning a more or less persis-
tent rhizome ; coat membranaceous ; scape slender, terete ; leaves
narrow, 1-3 mm. wide, apparently almost flat, slightly keeled ;
umbel 8— 20-flowered, nodding ; involucre very small ; perianth-
segments oblong-ovate, acute, nearly white, without a distinct mid-
vein ; stamens and style exserted ; capsule 6-crested.

This resembles most A. ccrnnuin, but differs in the fewer flow-
ered umbel, the narrower perianth-segments, and in the thinner
and narrower leaves, which are only slightly keeled.

New Mexico: Organ Mountains, 1894,^. 0. Wooton; 185 1-2,
C. WrigJit, 191 3.

South Colorado : 1 86 1 , C (T. Parry, j^o.

Arizona: Tanners Canon, 1892, Dr. T. E. Wilcox.

Astragalus Cusickii

Perennial from a creeping rootstock : stem about 5 dm. high,
strigose, somewhat branched ; leaves pinnate of 6-9 pairs of linear

(541)

642 Rydberg : Species from Western United States

leaflets which are 2-3 cm. long and about 2 mm. wide, glabrous
above and slightly strigose beneath ; raceme with a 1-2 dm. long
peduncle, rather lax and few-flowered ; flowers almost sessile, about
12 mm. long ; calyx about 7 mm. long, strigose with dark hairs ;
lobes short, lanceolate and unequal ; corolla yellow ; pod with a
stipe which is about i cm. long and curved upwards, upright, ob-
long, obcordate in cross-section, with the dorsal suture strongly
inflexed to about half-way to the ventral one, subcoriaceous, the
body being about 2 cm. long.

The specimens were named A. arrcctus Gray?, to which species
it has a superficial resemblance, differing in the pod, the struc-
ture of which places it near A. Dritmmondii and A. scopulorum.
From these it differs, however, in the short erect pod. It grows
on dry hillsides.

Oregon: Malheur, 1885, W. C. Ciisick, 1238 (Gray Herba-
rium).

Potentilla rosulata

Glandular and viscid pubescent throughout ; caudex thick and
lignose, topped with dense rosettes of leaves and short stems ; the
laUer, at least in the type specimens, less than i dm. high ; basal
leaves 4-5 cm. long, long-petioled, pinnately 5-foIiolate ; stem-
leaves ternate, short-petioled, or the upper subsessile ; lower
stipules lanceolate and thin ; the upper ovate and rather thick ;
leaflets thick, densely viscid and glandular pubescent, broadly
obovate, or the terminal orbicular, deeply crenate, or somewhat
cleft, 7-10 mm. long; pedicels 5-15 mm. long; hypanthium
about 5 mm. in diameter, densely viscid pubescent ; bractlets
ovate, about half as long as the broadly triangular ovate acute or
acuminate sepals ; petals small, oblong, whitish or light yellow,
about as long as the bractlets; stamens between 30 and 40;
anthers decidedly didymous ; pistils 20-40; style filiform, at-
tached near the apex of the ovary.

This is nearest related to Potentilla saxosa Greene,* but differs
in the less numerous leaflets of the basal leaves, the much thicker
and less incised leaflets, the shorter and stouter stems, the smaller

* In my monograph I transferred this species to Horkelia, on account of its close
resemblance to Horkelia Bailcyi, but a study of better material in Mr. T. S. Bran-
degee's herbarium has persuaded me that I made a mistake. The species is a true
Potentilla. There are three species, all belonging to the Potetitilleae, that are almost
identical in the vegetative parts, but still must be placed in three different genera.
These are: Potentilla saxosa Greene, Horkelia Bailey i Wats,, and PurJ>usia saxosa
Brandegee.

Rydberg : Species from Western United States 543

petals and the shorter hairs of the receptacle. It resembles also
P. rivalis somewhat in habit and leaves, but it has a thick peren-
nial caudex, much more numerous stamens and filiform style.

California: 29 Palms, Colorado Desert, 1898, A. H. Alvcr-
son (type in the herbarium of T. S. Brandcgee).

Horkelia chaetophora

Caudex stout, covered with the remains of leaf-stalks and
stipules from former years ; stems several, 1-1.5 dm. high, almost
scapose, finely puberulent ; basal leaves numerous, about i dm.
long, with 1 5-20 pairs of leaflets ; their stipules broad, brown,
obtuse, bristly ciliate ; leaflets 3-5 mm. long, divided to near the
base into linear-oblong segments, densely puberulent and tipped
with bristles ; cyme rather many-flowered and open ; hypanthium
5-7 mm. in diameter, puberulent and hirsute ; bractlets linear-ob-
long, one third shorter than the broadly lanceolate acute sepals ;
petals yellow, oblong, about equaling the sepals; stamens 10;
filaments filiform ; pistils about 20.

This is intermediate between H. Utahensis and H. pygmaca. It
resembles the former most in habit and flowers, but has the bristles
and obtuse stipules characteristic to H. pygmaca. From the latter
it differs in the larger size of the plant and flower and the many-
flowered and open cyme. It grows in rocky places in the moun-
tains at an altitude of 3000-3400 m.

California : Farewell Gap and Little Kern River, Tulare Co.,
1896, C. A. Pnrpus, 140^; Keweah Peak, 1895 (both in the her-
barium of T. S. Brandegee).

Horkelia Congdonis

Perennial with a woody caudex; stems erect, 3-4 dm. high,
few-leaved, somewhat branched above, glandular puberulent ; basal
leaves 1-1.5 dm. long, with 30-40 pairs of leaflets ; stem-leaves
similar but smaller ; upper stipules deeply cleft ; leaflets 3-5 mm.
long, cleft to the base into 4-5 oblong divisions, obtuse, glandular
puberulent ; cyme with a few ascending branches and short-pedi-
celed flowers ; hypanthium campanulate, 7-8 mm. in diameter,
glandular puberulent ; bractlets lanceolate, one half or two thirds
the length of the lanceolate acuminate sepals ; sepals almost equal-
ing the sepals, oblong, obtuse ; stamens 20 ; filaments slightly
dilated, subulate ; pistils numerous.

This is nearest related to Horkelia piirpiirasc ens, but differs in the

644 Rydberg : Species from Western United States

taller habit, the more branched cyme, the more acuminate sepals,
which in the type specimens are not reflexed, and the petals, which
are not retuse or emarginate as in that species.

California: Casa Diabolo, 1895, /. JT. Congdon (type in the
herbarium of T. S. Brandegee).

Mertensia tubiflora

Perennial ; stem 2—3 dm. high, glabrous striate, and somewhat
angled, branched above ; basal leaves oblanceolate, short petioled;
stem-leaves sessile, lanceolate to ovate, about 4 cm. long and 1-2
cm. wide, glabrate, except the hispid ciliolate margins, muricate
above, obtuse ; panicle contracted ; pedicels very slender and
drooping, about i cm. long, strigulose ; calyx slightly strigose,
about 4 mm. long, cleft half-way into oblong-lanceolate acutish
lobes ; corolla 13-15 mm. long ; tube about i o mm. long and 3 mm.
in diameter, more than twice as long as the limb ; the latter cam-
panulate with very short lobes ; nutlets very strongly muricate.

This species combines the general habit of M. lanceolata with a
corolla which is most like that of JZ oblongifolia.

Wyoming : Headwaters of the Tongue River, Big Horn
Mountains, July, 1898, F. Tivccdy, iig.

Symphoricarpos Utahensis

Syniplioricarpos nioiitaiins W^ats. King's Exp. 5: 132 partly;
not H.B. K.

Shrub a meter or more high, with brownish bark ; leaves
broadly ovate, more or less rounded at both ends, obtuse or often
mucronate, often coarsely sinuately toothed, pubescent when
young, glabrate in age, 3-4 cm. long and 2-3 cm. wide ; flowers
in terminal one-sided, drooping short spikes, or with smaller
clusters in the upper axils ; corolla somewhat funnelform, about
8 mm. long.

This resembles most .S". racemos^is in inflorescence and leaves,
but differs in the form of the corolla. The inflorescence, the
larger and less pubescent leaves and the size of the bush separate
it from S. rotimdifolius.

Utah: Logan, August, 1895, P. ^. Rydberg {l^y'^€)\ Wah-
satch Mountain, 1869, .S". Watson, .//j, in part.*

* Watson includes under this number not less than three distinct forms. Of these
one belongs to this species, one from Virginia Mountain, Nevada, to S. oi-eophilus, and
the third from the Uintahs to the next or an undescribed species.

Rydberg : Species from Western United States 545

Symphoricarpus Parishii

Apparently rather tall shrub for the group ; bark of the old
stems gray, of the young twigs brown ; leaves of older stems
small, about 1.5 cm. long, narrowly oval, acutish at both ends,
densely pubescent, more or less bluish green, rather thick ; those
of the young shoots larger, about 3 cm. long, deeply 3-lobed and
coarsely toothed ; corolla elongated campanulate, 6-7 mm. long.

This resembles mostly 5. rottuidifolius in pubescence and flow-
ers, but is evidently a larger plant and the leaves are bluish green
and acutish at both ends. It seems to be confined to Southern
California.

California: San Bernardino Mountains, 1892, 5. B. Parish,
2514; 1894, J o_v.

Erigeron flabellifolius

Perennial with a long slender creeping rootstock ; stem 1-2
dm. high, few-leaved, glandular puberulent above ; basal leaves
petioled, about 3 cm. long, slightly glandular puberulent, cuneate-
flabelliform in outline, deeply 3-5 -cleft into cuneate 3-lobed divis-
ions or the lower simply 5-9-lobed at the apex ; stem-leaves
cuneate or obovate, smaller, subsessile and less divided ; heads
about 10 mm. high and 10-15 "i"^- ""^ diameter; bracts linear,
acuminate, with more or less spreading tips, dark brown or purplish
black, glandular puberulent; rays 7-8 mm. long and 1.5-2 mm.
wide, light pink or white.

This is a member of thciS". coinposititni group, easily distinguished
from its relatives by the form of the leaves, which are never com-
pound, but simply cleft two thirds their length or less. It is also
characterized by the lack of hirsute pubescence generally found in
that group. It grows in rocky slides at an altitude of 3600 m.

Wyoming : Yount's Peak, Teton Forest Reserve, August,
1897, Tweedy, 536.

Erigeron spathulifolius

Perennial from an ascending rootstock ; stems 5-8 cm. high,
generally ascending, glabrous or slightly puberulent above, 3-5-
leaved ; basal leaves about 2 cm. long, perfectly glabrous, some-
what fleshy, broadly spatulate, tapering into a short petiole, entire-
margined, obtuse or acutish; stem leaves 1-1.5 cm. long, linear-
oblong or oblanceolate, sessile, obtuse ; head solitary, 7-8 mm,
high and 10-15 mm. in diameter, excluding the rays; bracts

54G Rvdberg: Species from Western United States

linear-lanceolate, acute, black, slightly puberulent ; rays light
blue, in age white, about 8 mm. long and 2-3 mm. wide.

In leaves and heads, this resembles most E. siinplex Greene, but
has a different root-system, is a much more glabrous plant and
lack altogether the long villous hairs on the involucre character-
istic of that species. On account of its root-system, it may be
associated with E. icrsiims and E. r'adicatus, but lacks the hirsute
pubescence of those species and has broader rays. The same char-
acters, together with the single head and broad leaves, separate it
from E. Eatonii, which also has somewhat the same habit. It is
an alpine species growing at an altitude of 3000 m. or more.

Wyoming : Black Rock Creek, Teton Forest Reserve, August^
1897, Tzueedv, 54.3.

Antennaria angustifolia

Surculose-proliferous ; leaves of the stolons linear or linear-
oblanceolate, about 1.5 cm. long, finely tomentose on both sides;
stem-leaves narrowly linear, erect, the uppermost subulate ; heads
few in a subcapitate cluster, 4-5 mm. high ; involucre campanu-
late, tomentose at the base ; bracts of the fertile head linear-
oblong, acute, yellowish or brownish white.

This is nearest related to A. pmvifolia and A. microphylla, from
which it differs in the subcapitate heads and the very narrow
leaves.

California: Yosemite Valley, 1865,/. Torrcy (labeled A.
stenophylla ? ) ; Hat Creek, J. S. Newberry (labeled A. luzidoides ;
both in the Torrey Herbarium).

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 3.

THE DICHOTOMOUS PANICUMS;
SOME NEW SPECIES-I.

BY GEO. V. NASH

NEW YOEK
1899

[Reprinted from the Bttllbtin of the Tobbey Botanical Club, 26: 568-581. 15 Nov., 1899.

The dichotomnus Panicums; Some new Soecies.— I

By Geo. V. Nash

Panicum Bushii

A tufted perennial, glabrous, with the exceptions noted below.
Culms about 3 dm. tall, the nodes sparingly barbed, finally much
branched : leaves about 3 ; sheaths about as long as the internodes,
those on the branches short and overlapping ; ligule a dense ring
of short hairs about 0.2 mm. long ; blades erect, linear, acuminate,
serrulate and very rough on the margins, ciliate at the base with a
few very long hairs arising from papillae, the larger primary blades
8-10 cm. long, 3-4 mm. wide, those on the branches usually 6 cm.
or less long : panicle much exserted, 6-7 cm. long, its branches
erect-ascending, the larger 3-3.5 cm. long, the secondary panicles
much smaller: spikelets 2.5 mm. long and about 1.2 mm. broad,
obovate, the scales glabrous, the first scale orbicular or very broadly
ovate, I -nerved, rounded or obtuse at the apex, about one-third as
long as the spik^let, the second and third scales 7-nerved, the
second a little the shorter, the fourth scale yellowish white, about
2 mm. long and 1.2 mm. wide, about as long as the second.

Collected by B. F. Bush, in dry ground, in McDonald Co.,
Missouri, July 24, 1893 no. 413. Related to P. augustifolium,
but the spikelets are glabrous and of a different shape -and the
sheaths and blades glabrous.

Panicum ciliosum

A tufted perennial. Culms 3-5 dm. tall, rather stout, ascend-
ing, papillose-hirsute with spreading hairs, finally much branched :
leaves 4 or 5 ; sheaths shorter than the internodes, densely hirsute
with spreading hairs, ciliate on the margins ; ligule a dense ring of
hairs about 0.5 mm. long; blades erect or ascending, narrowed
toward the base, glabrous above, ciliate on the margins, the stiff
hairs arising from papillae, the lower surface densely pubescent
between the nerves with short spreading hairs : panicle broadly
ovate, about 8 cm. long, included at the base, its axis pubescent
with short hairs, the branches spreading, the larger about 3 cm.
long : spikelets a trifle less than 2 mm. long and about 0.9 mm.
wide, elliptic, the outer 3 scales strongly pubescent with rather long
spreading hairs, the first scale broader than long, about one quar-
ter as long as the spikelet, i -nerved, rounded at the apex, the

(568)

569 Nash : The uichotomous Panicums

second and third scales 7-nerved, about equal in length, the fourth
scale white, a little longer than the third, about 1.6 mm. long and
0.8 mm. wide.

Type collected by S. M. Tracy, at Biloxi, Mississippi, Septem-
ber I, 1898, no. 4580. In habit and general appearance much
resembling P. piibescens, but the ciliate blades with the upper sur-
face glabrous at once separate it. The specimen above described
is the late state and has the panicle included ; the early form of the
plant will probably be found to have an exserted primary panicle.

Panicum Glutei

A tufted perennial, glabrous, with the exceptions noted below.
Culms rather stout, 6-8 dm. tall, at length branched : leaves 4 or
5 ; sheaths rather loose, minutely pubescent at the apex and
usually more or less ciliate on the exterior margin, the exterior
basal ones pubescent; ligule a dense ring of hairs about 0.3 mm.
long ; blades firm, ascending, often appearing as if erect in press-
ing, lanceolate, smooth on both surfaces, rough on the margins,
the lower and larger 7-14 cm. long, 7-12 mm. wide, the basal
blades ovate-lanceolate, 3-4 cm. long and 10-13 mm. wide, long-
ciliate on the margins : panicle considerably exserted, broadly
ovate, 6-10 cm. long, its ascending branches smooth, the larger
ones 4-6 cm. long: spikelets oval, acutish, about 2.3 mm. long
and about 1.3 mm. wide, the first scale broadly ovate, obtuse or
somewhat acute, i -nerved, glabrous or nearly so, the second and
third scales densely pubescent with very short ascending hairs,
9-nerved, the second one shorter than the third and usually a little
shorter than the fourth, the fourth scale yellowish, oval, about 2
mm. long and about 1.2 mm. wide.

Pine-barrens of southern New Jersey. Collected by Mr. W.
N. Clute, after whom I take pleasure in naming it, on a trip from
Tuckerton to Atsion, July 3-6, 1899. A most distinct species.

Panicum curtifolium

A tufted perennial, glabrous, with the exceptions noted below.
Culms slender, weak, 2-3 dm. tall, finally much branched : leaves
3 or 4 ; sheaths less than one half as long as the internodes, usually
about one third as long, sparsely pubescent with long weak spread-
ing hairs ; ligule a dense ring of hairs about 0.3 mm. long ; blades
widely spreading, lanceolate, minutely serrulate and rough on the
margins, a few long hairs on the upper surface just back of the
ligule, the culm blades 1.5-3 cm. long, 3-4.5 mm. wide, the basal
leaves 4-5 cm. long : panicle considerably exserted, broadly ovate,

Nash : The dichotomous Panicums 570

2.5-3.5 cm. long, its slightly hispid branches widely spreading, the
larger 1.5-2 cm. long: spikelcts about 1.5 mm. long and about
0.75 mm. wide, elliptic, the scales glabrous, the first scale about
one third as long as the spikelet, broadly ovate, obtuse, i -nerved,
the second and third scales 7-nerved, the second shorter than the
third, the fourth scale about equalling the third, about 1.25 mm.
long and about 0.6 mm. broad, yellowish-white, ob.scurely apiculate.
Collected by S. M. Tracy at Ocean Springs, Mississippi, May
2, 1898, no. 4598. Related to P. lucidicin As\\e,hn{. distinguished
by its smaller spikelets and sparsely pubescent sheaths.

Panicum decoloratum

A more or less purplish tufted perennial, glabrous, with the ex-
ceptions noted below. Culms 4-6 dm. tall, stout, finally branched,
the nodes barbed : leaves 5 or 6, extending to the base of the
panicle, the upper one including its base ; sheaths loosely embrac-
ing the culm, usually overlapping and hence concealing the culm,
the lower and basal ones papillose-hirsute between the nerves, the
hairs of the former early deciduous, the upper sheaths ciliate on
the external margin and with a minutely pubescent ring at the
apex ; blades erect or ascending, variously colored with black-
purple, broadly lanceolate, a little rough on both surfaces, especially
above, cordate-clasping at the base, 7-12 cm. long, 1.2-2 cm. wide,
minutely serrulate and very rough on the margins : panicle in-
cluded at the base, its branches hispidulous : spikelets on hispid-
ulous pedicels, 2.7 mm. long and 1.3 mm. wide, elliptic, the first
scale from nearly orbicular to broadly ovate, about one third as
long as the spikelet, i -nerved, obtuse or rounded at the apex,
glabrous, the second and third scales rather sparingly pubescent
with ascending hairs, 9-nerved, the second scale a little shorter than
the third, the fourth scale slightly yellowish-white, about 2.4 mm.
long and i mm. wide, obtusely and rather obscurely apiculate,
minutely pubescent at the apex.

Collected by Mr. E. P. Bicknell on a sandy railroad bank at
TuUytown, Pennsylvania, May 30, 1899. The panicle in the ma-
terial at hand may not be fully developed, and so a later stage
may show it exserted instead of included at the base. In habit
much resembling P. clandestimim, but that species is larger in
every way, the pubescence is harsh and hispid and occurs on all
the sheaths, the blades are much longer and with the margins near
the base conspicuously ciliate with long stiff hairs, and the spike-
lets are larger (exceeding 3 mm. in length).

571 Nash : The dichotomous Panicums

Panicum Earlei

A densely tufted perennial. Culms slender, 1-1.5 dm. tall,
usually with a few long weak scattered hairs below, finally branched,
the nodes rather sparingly barbed : leaves about 3 ; sheaths rather
sparingly hirsute with long hairs ; ligule a dense ring of hairs
about 0.3 mm. long ; blades ascending, lanceolate, rather spar-
ingly hirsute on both surfaces with long spreading hairs, 1—3
cm, long, 2-6 mm. wide, minutely serrulate and roughened on the
margins : panicle broadly ovate, 2-3 cm. long, its smooth branches
spreading, the larger ones 1-1.5 cm. long: spikelets about 1.3
mm. long and 0.7 mm. wide, elliptic, obtuse, glabrous, the first
scale orbicular or broadly oval, i -nerved, obtuse, the second and
third scales 7-nerved, the second scale shorter than the third and
fourth, the fourth scale white, about i mm. long and 0.6 mm.
wide, oval.

Type collected at Auburn, Lee Co., Alabama, on May 7,
1898, by Messrs. F. S. Earle and C. F. Baker, no. 1532; no.
1535, of the same place and date, also belongs here.

Panicum epilifolium

A tufted perennial, glabrous, with the exceptions noted below.
Culms 2-3.5 dm. tal : leaves 2 or 3 ; sheaths shorter than the
internodes, minutely pubescent at the apex, ciliate on both mar-
gins with long slender hairs ; ligule a scarious ciliolate ring about
0.2 mm. wide ; blades widely spreading, linear-lanceolate, 4-7
cm. long, 5-7 mm. wide, minutely pubescent on the upper surface
between the nerves, serrulate and very rough on the margins :
panicle exserted, ovate, 5-7 cm. long, its branches spreading or
ascending, the larger ones 2-2.5 cm. long: spikelets 3 mm. long
and about 1.5 mm. broad, oval, obtusely apiculate, the first scael
glabrous or with a few scattered hairs, i -nerved, nearly orbicular,
acute, a little less than one half as long as thespikelet, the second
and third scales densely pubescent with short spreading hairs, 9-
nerved, the second usually a little shorter than the third and fourth,
the latter scale yellowish, 2.5 mm. long and a little over i mm.
wide, elliptic, obtusely apiculate.

Type collected by the writer in a scrub hammock at Eustis,
Lake Co., Florida, March 12-31, 1894, no. 45. Also secured at
the same place by Professor L. M. Underwood, on March 22,
1891, no. 2250. It bears some resemblance in habit to P. cilii-
fcriun, but is readily distinguished from that species by its glabrous
sheaths, naked blade-margins and more acute spikelets.

Nash : The dichotomous Panicums 672

Panicum flavovirens

A densely tufted liijht green perennial, glabrous, with the ex-
ceptions noted below. Culms 2-3 dm. tall, slender, finally much
branched : leaves 2 or 3 ; sheaths very short, those of the primary
leaves about one third as long as the internodes, one margin
usually extending above the other, making the summit of the
sheath more or less oblique, the lower sheaths ciliate on the
margin, the exterior basal ones pubescent all over ; ligule a dense
ring of short hairs about 0.2 mm. long; blades thin, erect, with-
out a white margin or nearly so, entire or very minutely serrulate,
hence smooth or nearly so on the margin, linear-lanceolate, those
on the main culm 2.5-4 cm. long, 2.5-4 mm. wide, commonly
minutely pubescent on the lower surface between the nerves, those
on the branches much shorter, the basal blades longer, 4-6 cm.
long : panicle much exserted, 3-4 cm. long, broadly ovate, its
branches spreading, the larger 1-2 cm. long, the secondary
panicles much smaller, barely exserted and with spreading
branches : spikelets 1.5 mm. long and 0.7 mm. broad, elliptic, the
outer 3 scales densely pubescent with spreading hairs, the first
scale I -nerved, broadly ovate, obtuse, about one half as long as
the spikelet, the second and third scales 7-nerved, about equal in
length, the fourth scale yellowish-white, 1.3 mm. long and about
0.6 mm. wide.

Type collected by the writer in Lake Co., Florida, June 16-30,
1895, no. 2061 ; growing in swampy woods along the edge of
road leading to the ford near the J. T. & K. W. R. R. bridge
across the Wekiva river. No. 2487a, collected in a similar habitat
at Lake City, Columbia Co., in the same state, on Aug. 30, 1895,
is also referred here.

Differs from P. albomarginatum in the thin linear-lanceolate
blades which are entirely or almost devoid of the white margin,
and barely if at all rough on the margins. In P. albomarginatum
the blades are very thick, much broader, and with a wide strongly
serrulate white margin which is much thickened.

Panicum Helleri

A tufted perennial, glabrous, with the exceptions noted below.
Culms 2-4 dm. tall, appressed-pubescent below with long hairs,
the nodes sparingly barbed, finally branched : leaves 5 ; sheaths
shorter than the internodes, the middle ones only about one half
as long, ciliate on the exterior margin, bearing between the promi-
nent nerves scattered papillae, from which sometimes arise stiff

573 Nash : The dichotomous Panicums

hairs, the internerves of all but the upper sheaths minutely pu-
bescent : ligule a dense ring of hairs about 0.6 mm. long : blades
broadly lanceolate, thin, a little narrowed toward the sparsely
ciliate rounded base, the margins minutely serrulate, rough, 6-8
cm. long, 6-12 mm. wide: panicle included at the base, 6-8 cm.
long, its branches ascending, the larger ones 3-4 cm. long, the
secondary panicles smaller: spikelets 3.25-3.5 mm. long and
about 1.5 mm. wide, the first scale broadly triangular-ovate, .1-
nerved, the second and third scales pubescent toward the base
with a few scattered hairs, the second scale 11 -nerved, the third
scale 9-nerved, the fourth scale yellowish-white, 2.5 mm. long and
about 1.3 mm. wide, oval, obtusely and obscurely apiculate.

Collected at Kerrville, Kerr Co., Texas, by A. A. Heller, May
14-21, 1894, no. 1759. Differs from P. pernervoswn in the pu-
bescent culm and sheaths, the broader blades of a different shape
and the narrow sp kelets which are usually sparsely pubescent.

Panicum paucipilum

A tufted perennial, glabrous, with the exceptions noted below.
Culms 6-10 dm. tall, finally sparingly branched : leaves 5-8 ;
sheathes one third to one half as long as the internodes, the ex-
ternal margin ciliate toward the summit ; ligule a dense ring of
hairs about 2 mm. long : blades erect or ascending, thickish, rather
firm, sometimes minutely puberulent on the lower surface, usually
with a few hair-bearing papillae at the base, the lower and larger
6-9 cm. long, 5-7 mm. wide : panicle finally considerably exserted,
rather dense, oblong, usually 5-10 cm. long, its branches erect-
ascending or erect, the larger ones 2.5-4 cm. long: spikelets
numerous, about 1.4 mm. long and 0.8 mm. wide, oval, the first
scale about one third as long as the spikelet, i -nerved, orbicular,
the second and third scales pubescent with spreading hairs, the
former plainly, the latter obscurely, 9-nerved, the fourth scale yel-
lowish, about 1.3 mm. long and 0.8 mm. wide, obscurely apiculate.

In wet soil, southern New Jersey to Mississippi. Type col-
lected by Mr. E. P. Bicknell, at Wildwood, New Jersey, May 30
and 31, 1897. The following specimens from Mississippi are also
referred here :

Ocean Springs, July 19, 1889, F. S. Earle ; Biloxi, Sept. i,
1892, S. M. Tracy; Beauvoir, May 13, 1898, S. M. Tracy, no.
4594 (distributed as P. octonodinn S. & S.).

Intermediate between P. octotioduju and P. Eatoni. Differs
from the former, to which it is closely related, in the ciliate margin

Nash : The dichotomous Panicums 574

of the sheaths, the few basal hairs of the blades, and particularly
in the strongly pubescent spikelets. From P. Eatoni it may be
distinguished by its much smaller spikelets with the first scale
glabrous and orbicular.

Panicura longiligulatum

A tufted perennial, glabrous, with the exceptions noted below.
Culms 4—5 dm. tall, slender, finally branching, the branches fasci-
culately much divided and forming dense masses at their ends ;
leaves 4 or 5 ; sheaths usually from one third to one half as long
as the internodes, minutely pubescent between the prominent
nerves ; ligule a ring of long erect silky hairs about 3 mm. in
length ; blades ascending, lanceolate, obtusely and minutely pubes-
cent on the lower surface, the margins serrulate and very rough,
the primary culm blades 2.5-3 cm. long, about 3 mm. wide, those
on the branches much smaller, the basal blades thick, broadly
lanceolate, 4—5 cm. long : panicle oval, 5—6 cm. long, considerably
exserted, its branches spreading, the larger ones about 3 cm. long:
spikelets about 1.3 mm. long and 0.8 mm. wide, oval, the outer 3
scales densely pubescent with spreading hairs, the first scale ovate,
I -nerved, about one third as long as the spikelet, the second and
third scales 7-nerved, about equal in length, the fourth scale yel-
lowish white, about 1.2 mm. long and 0.7 mm. wide.

Collected by Dr. Geo. Vasey, at Apalachicola, Florida, in
1892. Its relations-hip is with P. parvispiaduni, but its more
slender culms, smaller blades and spikelets and the glabrous mar-
gins of the sheaths at once distinguish it.

Panicum patentifolium

A tufted purplish perennial, glabrous, with the exceptions noted
below. Culms erect or nearly so, 2—4 dm. tall, puberulent,
slender, finally much branched : leaves 2-4, rather distant ; sheaths
less than one half as long as the internodes, minutely pubescent,
especially at the apex on the margins, rather loosely embracing
the culm ; ligule a dense ring of hairs about 0.25 mm. long ; blades
widely spreading, firm, lanceolate, 2. 5—4 cm. long, 2—4 mm. wide,
puberulent at the very base on the upper surface, smooth on both
sides, rough on the margins, the basal ones similar in shape and
texture but larger : panicle at length considerably exserted, broadly
ovate, 4-6 cm. long, its axis and spreading branches puberulent,
the larger branches 2—3 cm. long: spikelets about 2.5 mm. long
and 1.3 mm. wide, the first scale orbicular, clasping, purple, at
least at the base, i -nerved, rounded at the apex, one half as long

575 Nash : The dichotomous Panicums

as the spikelet, the second and third scales pubescent with spread-
ing hairs, 7 -nerved, the fourth scale white, a little exceeding 2 mm.
in length, oval, minutely pubescent at the apex.

Type collected by the writer at Eustis, Lake Co., Florida,
March 12-31, 1894, no. 72, in dry sand in a scrub hammock. No.
52 of the same collection also belongs here.

Related to P. Wcbberianiun, but the more slender culms and
the smaller and widely spreading blades readily separate it.

Panicum perlongum

A tufted pubescent perennial. Culms 2-4 dm. tall, simple,
glabrous or sparingly pubescent, the nodes barbed, later with short
basal culms : leaves i or 2 ; sheaths hirsute with long ascending
hairs; ligule a dense ring of hairs about 0.7 mm. long ; blades
elongated, linear, erect, papillose-hispid beneath, glabrous, rough
above, 2-3 mm. wide, the upper blade commonly 8-14 cm. long,
occasionally shorter : panicle much exserted, generally extending
beyond the apex of the upper leaf, 4-6 cm. long, its branches erect
or erect-ascending, the larger ones usually 2-3 cm. long : spike-
lets, on hispidulous pedicels, obovate, about 3.25 mm. long and
i.c^-x.j^ mm. wide, the outer 3 scales with a few scattered long
hairs, especially near the base, the first scale one quarter to one
third as long as the spikelet, orbicular-ovate, i -nerved, the second
and third scales 9-nerved, about equal in length, the fourth scale
oval, 2.5 mm. long and about 1.5 mm. wide, yellowish white, ob-
tusely apiculate, its summit reaching the apex of the third scale.

On prairies and dry soil, Illinois to North Dakota, south to
Indian Territory. Type collected in Indian Territory at Creek
Nation, by M. A. Carlton. April 25, 1891, no. 98, and distributed
as P. depanperatttm Muhl. It differs from that species in the
smaller pubescent obtuse spikelets with the second and third scales
not exceeding the fourth scale. From P. liiicarifoliuni, to which it
is more nearly related, it is separated by its larger sparsely pubes-
cent spikelets.

The following specimens, distributed as P. depmiperatuvi Muhl.,
are also referred here :

Illinois: June 7, 1848, 5. B. Mead.

South Dakota: Custer, July 18, 1892, P. A. Rydberg, no.

7/00.

Iowa: Ames, June 22, 1896, C. R. Ball, no. /yj.

Kansas: Prairie, Riley Co., 1896, A. S. Hitchcock, no. 881.

Nash : The dichotomous Panicums 576

Panicum pernervosum

A glabrous perennial. Culms 3-5 dm. tall, finally branching :
leaves 3 or 4 ; sheaths ciliate on the exterior margin, the lower
longer, the upper shorter than the internodes ; ligule a dense ring
of hairs about 0.5 mm. long ; blades erect or ascending, narrowed
toward the base, serrulate and very rough on the margins, the
intermediate and upper blades 5- 10 cm. long, 5—9 mm. wide,
ciliate toward the base with a few long hairs, the lower ones usu-
ally pubescent on the lower surface, shorter and broader, ciliate for
two thirds their length : panicle considerably exserted, 7—12 cm.
long, its branches ascending, the larger ones 4—6 cm. long : spike-
Jets 3 mm. long and 1.8 mm. wide, broadly oval, turgid, rounded
at the apex, the scales glabrous, the first scale broader than long,
I -nerved, about one third as long as the spikelet, the second and
third scales coarsely 9-nerved, the second a little shorter than the
third, the fourth scale yellowish white, 2.5 mm. long and about
1.6 mm. broad, obtusely and rather obscurely apiculate.

Type collected by Elihu Hall in woods, at Houston, Texas,
A.pril 16, 1872, no. 830. Mr. G. C. Nealley also secured it in
the same state in 1886.

Panicum psammophilum

A tufted perennial. Culms 2-4 dm. tall, appressed-hirsute
below, puberulent above, finally much branched : leaves on the
main culm about 4, occasionally 3 or 5 ; sheaths shorter than the
internodes, appressed-pubescent, the basal ones with long hairs,
the upper and those on the branches with very short hairs ; ligule
a dense ring of hairs about i mm. long ; blades erect or nearly so,
thick, firm, serrulate on the margins, puberulent beneath, the
ovate-lanceolate basal ones, and occasionally also those on the
culm, with a few very long scattered erect hairs on the upper sur-
face, the primary blades lanceolate, 2-5 cm. long, 2-5 mm. wide,
those on the branches 2-3 cm. long, 2-3 mm. wide ; primary
panicle broadly ovate, usually 2-3 cm. long, rarely larger, its axis
and usually ascending branches puberulent : spikelets broadly ob-
ovate, 1. 3- 1. 5 mm. long, rarely a little larger, and 0.8-1 mm.
wide, obtuse, the outer 3 scales densely pubescent with spreading
hairs, the first scale usually about one third as long as the spikelet,
sometimes a little larger, i -nerved, orbicular or broadly ovate,
acute or obtuse, the second and third scales 9-nerved, the sec-
ond usually a little shorter than the third, the fourth scale yel-
owish, 1. 2- 1. 5 mm. long, oval.

In sandy soil, on or near the coast, Massachusetts to New Jer-
sey. Specimens examined :

577 Nash : The dichotomous Panicums

Massachusetts : Martha's Vineyard, Miss Witinan, July, 1890 ;
Ipswich, Geo. V. iVcrs/i, Aug. 25, 1898, no. J2.

New York : Fisher's Island, C. B. Graves, Aug. 29, 1898,
no. 12.

New Jersey : Wildwood, May 30 and 31,1 897, E. P. Bick-
ne//; Tom's River, July 25-31, 1898, IV. N. Clute, no. 175 (type);
Tuckerton to Atsion, July 3-6, 1899, W. N. Clntc.

Related to P. Addisonii, but at once distinguished by its smaller
and relatively broader spikelets.

Panicum pseudopubescens

A densely tufted pubescent perennial. Culms usually rather
stout at the base, 2-4 dm. tall, hirsute with ascending hairs, finally
branched, the nodes barbed : leaves usually 3 ; sheaths less than
one half as long as the internodes, densely hirsute with spreading
hairs 1.5-2 mm. long ; ligule a dense ring of hairs about 0.5 mm.
long ; blades erect, rather firm, lanceolate, serrulate and very rough
on the margins, 4-10 cm. long, 3-1 1 mm. wide, densely hispid on
the lower surface with spreading hairs, the upper surface with a
ring of long stout erect hairs just back of the ligule and usually
also more or less hirsute with spreading hairs : panicle consider-
ably exserted, broadly ovate, 5-9 cm. long, its axis glabrous or
with a few scattered hairs below, the branches with the axils spar-
ingly hirsute, ascending or spreading, the larger ones 2.5-5 cm,
long: spikelets 2.25-2.5 mm. long and i — 1.2 mm. wide, obovate,
a little pointed, the first scale usually about one third as long as
tfie spikelet, i -nerved, glabrous or sparingly pubescent, broadly
ovate or nearly orbicular, acute or obtuse, the second and third
scales densely pubescent with spreading hairs about 0.25 mm. long,
commonly 9-nerved, the second a little shorter than the third, the
fourth scale nearly white, a little less than 2 mm. long and about
I mm. wide, oval.

Type collected at Auburn, Lee Co., Alabama, May 7, 1898, by
Messrs. F. S. Earle and C. F. Baker, no. 1537. Nos. 1522, 1524,
1526 and 1529, of this same collection, are also referred here. It
is distinguished from any form of P. piibeseens by its much larger
spikelets, and from P. Atlauticum by the shorter pubescence and
stouter culms

Panicum pubifolium

/'. latifolium var. niolle Vasey, Contr. U. S. Nat. Herb. 3: 33.
1892. Not Z'. ;;/^//^ Sw. 1788.

Nash : The dichotomous Panicums 578

P. Portcriannui Nash, Bull. Torr. Bot. Club, 22 : 420. In part.

1895.

A softly pubescent densely tufted perennial. Culms 3-7 dm.
tall, pubescent with soft weak spreading hairs, those at the base
the longer, finally branched, the nodes densely barbed with long
hairs : leaves 3-5 ; sheaths shorter than the internodes, often only
one half as long, ciliate on the margins, densely pubescent, at least
all but the uppermost, with spreading weak usually long hairs, also
a dense ring of short hairs at the apex ; blades spreading or ascend-
ing, minutely serrulate and very rough on the margins, ovate-
lanceolate to ovate, acuminate, gradually narrowed to the rounded
cordate-clasping base, often inequilateral, pubescent on both sur-
faces with short soft spreading hairs, the upper primary blades 7-
1 1 cm. long and 2-3 cm. broad, the lower primary blades, as well
as those on the branches, smaller : primary panicle usually but
little exserted, sometimes included at the base, 7-1 1 cm. long, its
axis, as well as the branches, densely pubescent with short soft
spreading hairs, the branches spreading or ascending, the larger
ones 3-4 cm. long, the secondary panicles much smaller, included
at the base: spikelets 4-5 mm. long and about 1.6 mm. broad,
narrowly obovate, the scales distantly inserted on the rachilla, the
outer 3 scales strongly pubescent with long spreading hairs, the
first scale 3 -nerved, from two fifths to one half as long as the
spikelet, obtuse or acute, the second scale 1 1 -nerved, the third
scale 9-nerved, the former a little shorter than the latter, the fourth
scale yellowish, 3.5 mm. long and 1.4-1.6 mm. broad, pubescent
at the obtusely apiculate apex.

Usually in rocky woods. New York to Missouri, south to
Florida and Mississippi.

Among a large number of specimens of this grass examined,
the following are referred to as well representing this species :

Pennsylvania: Chestnut Hill, Easton, July i, 1887, T. C.
Porter (distributed as P. Walteri viollc).

District of Columbia : Washington, June 9, 1894, Th. Holm,
no. 14 (distributed as P. latifoUuni).

Virginia : Between Fall Creek and Danville, June 3, 1891, /.
K. 5wrt//and A. A. Heller, no. 466 (distributed as P. latifoliuni).

Missouri : McDonald Co., July 24, 1893, B. F. Bush, no. 415
(distributed as P. latifoliuni) ; Montier, June 30, 1894, B. F. Bush,
no. 754 (distributed as P. Walteri).

Tennessee: Knox Co., July 9, 1893, T. H. Kearney, Jr. (dis-

579 Nash : The dichotomous Panicums

tributed as P. Waltcri mollc) ; Knoxville, July, 1898, A. Ruth, no.
y^ (distributed as P. Portcrianuni).

Georgia : Stone Mt., Aug. 1-6, 1895, /. K. .S";//rt// (distributed
as P. Portcriamivi).

Readily distinguished from P. Portcriaiium by the pubescent
sheaths and the lower surface of the blades and the hirsute panicle.

Panicum pyriforme

A densely tufted perennial. Culms 3—4 dm. tall, rather slender,
glabrous, rather weak, finally much branched ; leaves usually 2,
or sometimes 3 ; sheaths much shorter than the internodes, densely
papillose-hirsute with reflexed hairs ; ligule a ring of hairs about
0.3 mm, long; blades thin, lax, glabrous on both surfaces, serru-
late and very rough on the glabrous margins, long-acuminate,
narrowed to the barely rounded base, 1—2 dm. long, 8—12 mm.
wide, the basal ones often 2.5 dm. in length, the blades on the
branches much shorter : panicle much exserted, ample, loose and
open, 6—1 1 cm. long, its branches widely spreading, the larger ones
4-6 cm. long: spikelets rather few, about 2.5 mm. long and 1.5
mm. wide, broadly obovate, the first scale about one third as long
as the spikelet, broadly triangular-ovate, i -nerved, the second and
third scales densely pubescent with long hairs, 9-nerved, the fourth
scale yellowish white, about 2 mm. long and 1.5 mm. wide, oval,
strongly apiculate.

Type collected by the writer in clay soil, at Orange Bend,
Lake Co., Florida, March 12 31, 1894, no. 239. The larger
spikelets and glabrous blades at once distinguish this from P. loxi-
floniin, to which it is otherwise related. The following numbers of
my collection of 1895 are also referred here: 2034, 2156, 2531a.

Panicum strictifolium

A tufted perennial. Culms 2.5—5 dm. tall, strongly pubescent
toward the base with long stiff hairs, puberulent toward the summit,
finally much branched : leaves 3 or 4 ; sheaths much shorter than
the internodes, the lower ones densely pubescent with long stiff ap-
pressed hairs, the upper ones more sparingly so ; ligule a dense
ring of hairs about 1 mm. long ; blades erect or nearly so, rather
firm, narrowly lanceolate, appressed -pubescent beneath with stiff
hairs, the upper surface often with a few scattered long hairs, the
lower blades also with a ring of very long stiff hairs just back of
the ligule, serrulate and rough on the margins, the primar)- blades
4—7 cm. long, 3-5 mm. wide : panicle broadly ovate, 5-7 cm. long,
its axis and spreading branches minutel}' pubescent, the lower

Nash : The dichotomous Panicums 580

branches 2.5-3.5 cm. long: spikelets obovate, 3 mm. long and
about 1.5 mm. wide, the first scale a little more than one half as
long as the spikeiet, broadly ovate, clasping at the base, 3-nerved,
sparingly pubescent, the second and third scales densely pubescent
with rather long spreading hairs, 11 -nerved, the second distinctly
shorter than the third and the fourth, the fourth scale yellowish-
white, oval, 2.5 mm. long and about 1.3 mm. wide.

Collected by the writer in the high pine land at Eustis, Lake Co.,
Florida, May 3, 1894, no. 603. Most nearly related to P. viala-
con, but distinguished by the less copious pubescence which is
much finer and softer, and by the glabrous upper surface of the
blades. From P. ciliifcnim it is at once separated by its narrower
blades which are not ciliate on the margins.

Panicum trifolium

A much tufted perennial, glabrous, with the exceptions noted
below. Culms slender, 2-4 dm. tall, finally a little branched :
leaves usually 3, rarely 4, the uppermost one much above the
middle of the culm and generally but a little below the panicle ;
sheaths less than one half as long as the internodes, sometimes
but one quarter as long : ligule a dense ring of hairs about 0.4 mm.
long ; blades erect or nearly so, firm, lanceolate, often minutely
pubescent on the lower surface, the margins thickened and carti-
laginous, serrulate and very rough, 1.5-6 cm. long, 1.5-5 m"^-
wide, the basal ones numerous, 4-6 cm. long : panicle more or less
exserted, broadly ovate, 2.5-6 cm. long, its branches ascending,
the larger ones 1.5-3 cm. long : spikelets 1.5 mm. long and about
0.7 mm. wide, elliptic, the first scale nearly orbicular, glabrous,
I -nerved, one quarter to one third as long as the spikeiet, the
second and third scales densely pubescent with short spreading
hairs, 7-nerved, the fourth scale white, 1.3 mm. long and about
0.6 mm. wide, obtusely and obscurely apiculate.

In sandy soil. North Carolina to northern Florida, west to
Mississippi. Type collected by Dr. John K. Small, in the Ocmul-
gee River Swamp, below Macon, Georgia, May 1 8-24, 1 895. The
following specimens are also referred here :

North Carolina : Chapel Hill, W. W. Ashe (distributed as
P. ensifoliimi).

South Carolina: Aiken, 1867, H. W. Ravcnd.

Florida: Chapman, 1890, no. 3; Apalachicola, 1892, Dr.
Geo. Vasey.

581 Nash : The dichotomous Panicums

Alabama : Buckley ; Auburn, May 5, 1898, Earle and Baker,
nos. 1534 and 1547a.

Mississippi: 5. M. Tracy, Biloxi, Aug. i, 1894, no. 2865,
March 28, 1898, no. 4602, April 2, 1898, no. 4612 ; Avondale,
April 28, 1898, no. 4610; Horn Island, June i, 1898, no. 4613.

Related to P. alboinarguiatinn , but distinguished by its thinner
blades and more slender culms, which are leafy nearly to the
panicle.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 4.

DELPHINIUM CAROLINIANUM AND
RELATED SPECIES

BY P. A. RYDBERG

NEW YOKK
1899

[Reprinted from the Bulletin of the Toerby BoTiNiCAL Club, 26: 58Z-587. 15 Nov., 1899.

Delphinium Carolinianum and related Species

By p. a. Rydberg

None of our native larkspurs has been more misunderstood
than Ddphiiiiiun Carolinanuin Walt. , or D. azitreum Michx. This
species is found in the southern states only, its northwestern limit
being in Missouri. All specimens from the western states referred
to it belong to one of the other species described below. Any one
who has collected specimens of so-called D. aziireum in the prairie
states or in the Southwest has found trouble in trying to harmonize
the specimens collected with the descriptions in our manuals.
Some years ago I came to the opinion that the D. azureuin of Ne-
braska and neighboring states was quite different from the D.
azureiim of the South. It is only lately, however, that I have had
occasion to give closer attention to the matter, in connection
with a partial revision of the Ranunculaceae of Dr. Britton's Flora.
I have come to the conclusion that D. azureuin, as treated in Gray's
Synoptical Flora, contains about ten species. I regret that I have
not seen mature seeds of a few of the western species, as the seeds
afford excellent characters for determination.

All the species treated here have a leafy stem and seeds with
a loose cellular coat, that becomes transversely rugose squamel-
late. The most prominent characters by which they may be dis-
tinguished from each other are the following :

Bractlets some distance below the calyx and below the thickened portion of the pedicel.
Sepals deep blue ; principal segments of the leaves cuneate, cleft nearly or quite
to the middle. l- D- gerattiifolhtm.

Sepals white, tinged with blue ; segments cleft beyond the middle into narrow
oblong or linear lobes.
Spur about twice as long as the petals ; sepals obovate.
Spur straight or slightly curved.

Lobes of the lateral petals not diverging ; lower pedicels not much

elongated; spur mostly horizontal. 2. D. albescens.

Lobes of the lateral petals diverging ; lower pedicels elongated ; spur
mostly erect. 3- ^- camporum.

Spur strongly curved upward. 4. D. Penardi.

Spur thrice as long as the petals, slightly j-curved ; sepals oblong.

5. D. macroseratilu.

(582)

583 Rydberg : Delphinium Carolinianum

Bractlets close under the calyx on the thickened end of the pedicels.

Sepals greenish or yellowish white ; segments of the upper leaves oblong.

6. D. virescens.
Sepals blue or bluish ; segments of the upper leaves narrowly linear.

Plant tall, slender, green ; bractlets narrowly linear, almost subulate.

Seeds strongly wing-margined, only slightly rugose- squamellate ; raceme

simple and narrow. 7. D. Carolinianmn.

Seeds not wing-margined, strongly squamellate ; raceme often branched.

8. D. vimineum.
Plant low, stout, more or less canescent ; bractlets linear or lanceolate.
Sepals deep blue ; pedicels ascending. 9. D. Geyej-i.

Sepals light blue or white, tinged with blue or purple ; pedicels erect.

10. D. Wootoni.

I. Delphinium geraniifolium sp. nov.

Stem from a deep woody perennial root, stout, 3-4 dm. high,
finely grayish-strigose ; leaves numerous, especially at the base,
long-petioled, grayish-strigose, mostly 5-divided to the base ; di-
visions cuneate, twice 2-3 -cleft into broad oblong divisions ;
racemes many-flowered, somewhat branched ; pedicels ascending ;
bractlets linear, 2-4 mm. below the blue calyx ; spur stout, hori-
zontal, about one half longer than the petals, slightly curved ;
upper petals brownish, tipped with blue ; seeds unknown.

This is evidently nearest related to D. Gcycri, but differs by the
broad leaf segments and the form and position of the bractlets.
Dr. Gray referred it to D. viinincuin, which it resembles very little.

Arizona: Charles Valley, 1883, H. H. Rusby.

2. Delphinium albescens sp. nov.

Generally tall, 3-15 dm. high, from a woody branched root,
finely pubescent or glabrate below, somewhat viscid above ; leaves
rather variable, from 5 to i 5 cm. in diameter, repeatedly divided into
linear or the lower often into oblong divisions ; raceme long and
simple, sometimes 5— 6 dm. long; pedicels erect, 1—2 cm. long ;
bractlets narrowly linear, borne 2—4 (in fruit often 6-8) mm. be-
low the calyx ; sepals white with a blue spot and sometimes tinged
with blue ; spur stout, about twice as long as the petals, straight
or slightly curved, tinged with blue, generally horizontal or as-
cending ; upper petals very oblique at the summit, tinged with yel-
low ; the lateral ones bearded, 2-cleft, but the lobes not diverging;
follicles cylindric, pubescent; seeds 1.5—2 mm. long, brown,
sharply angled but not wing-margined, rather strongly squamellate.

This species has been included in D. Carolmiammt, but is easily
distinguished by the color of the flowers, the form and position of
the bractlets, the stouter habit, and especially by the seeds. Its

AND RELATED SpECIES 584

distinctness from D. caiuponim is not so clear. The principal char-
acter used by Prof. Greene in order to distinguish the latter from

D. Carolinianum, viz., the erect spur, does not hold, for D. Caro-
linianum sometimes has an erect spur ; a'lso D. albescens, as
shown in Dr. Houghton's specimens from Lake Winnipeg.
These were included by Greene in D. caiiiponim, but are
very unlike the plant from Texas, New Mexico and Arizona,
which must be taken for the type of D. cauiponim. Houghton's
plant differs in no respect, except the erect spur, from the common
Larkspur of the Prairie Region. The specimens from Arkansas
cited below are more slender and have narrower leaf-segments than
is usual. The following specimens of D. albescens are found in the
herbaria of the New York Botanical Garden and Columbia Uni-
versity.

Manitoba : Lake Winnipeg, Dr. Houghton.

Minnesota: Hennepin Co., \^go,J. H. Sandberg ; Fort Snell-
ing, 1889, E. A. Mearns.

South Dakota : Scalp Creek (collector not given).

Nebraska: Lincoln, 1887, H.J. Webber (type) ; Crete, 188 1,
G. D. Sivcezey.

Kansas: Ft. Riley, 1892, E. E. Gayle, 484; Atchison, 1892,

E. B. Knerr ; Riley Co., 1895, / B. Norton, 8 ; Topeka, 1891,
B. B. Smyth; Manhattan, 1889, W. A. Kellerman.

Colorado: Ft. Collins, 1896, C. F. Baker.
Arkansas : Dr. Pitcher.

Indian Territory: Sapulpa, 1895, B. F. Bush, logi.
Texas: San Antonio, 1894, A. A. Heller, ij8j.
Missouri : Eagle Rock, 1898, B. F. Bush, 228 ; Independence,
1894, 7.

Illinois : Augusta, 5. B. Mead.

3. Delphinium camporum Greene, Erythea, 2 : 183
This species is closely related to the preceding, but is generally
lower and stouter, with numerous basal leaves and {ew stem-leaves ;
the lower pedicels are much elongated, often 5 cm. long; the pedicels
as a rule are strongly curved at the end, bringing the spur into
an erect position. The flowers are similar to those of the preced-
ing, but generally more purely white, and the lobes of the some-

585 Rydberg : Delphinium Carolinianum

what longer lateral petals are more divergent. The upper petals
are less oblique at the top. To this species, I refer the following
specimens :

New Mexico: Jorunda del Murto, 185 1, Geo. TJiurber, 2gr ;
Mangus Spring, 1881, H. H. Rusby, ^ ; 1852, C. Wright, 8^0.

Texas: Rio Bravo del Norte, 1852, Schott.

4. Delphinium Penardi Huth, Helios, 10: 27. 1892

I have seen no specimens of this species and from the descrip-
tion one would come to the conclusion that it is simply a form of
D. albescens, the curved spur notwithstanding. We have seeds,
however, received from M. E. Autran of the Boissier Herbarium,
and these are very unlike those of D. albescens. They are large,
black, very irregular, and only slightly squamellate ; in fact, they
are almost identical with those of D. Geyeri. The upper petals are
also described and figured as being toothed at the apex, a condi-
tion I have never seen in D. albescens.

5. Delphinium macroseratilis sp. nov.

Stem slender, about 3 dm. high, finely grayish -pubescent ;
leaves divided to the base into 3—5 segments, these again 2—3-
cleft ; raceme simple, strict, many-flowered ; pedicels erect, about
I cm. long ; bractlets 2—4 mm. below the calyx, linear ; sepals
white, the lower ones with a bluish spot, oblong, more than twice
as long as the upper petals ; spur about three times as long as the
upper petals, tinged with bluish, slightly s-curved ; upper petals
very oblique and pointed at the summit ; lateral ones much longer,
2-cleft and bearded ; fruit and seeds unknown.

This is also closely related to D. albescens, differing in floral
characters and the fewer segments of the leaves. Further study
of the species is needed, especially as seeds are lacking.

Texas: Tom Greene Co., 1879, Frank Tiveedy.

6. Delphinium virescens Nutt. Gen. 2: 14. 18 18

There is only a fragmentary specimen in the Torrey Herbarium,
but this is enough to show that it is quite distinct from D. Caro-
linianum. Not only is the corolla of a different color, but it is
much larger and the spur is strongly hooked at the end. The
plant is much stouter than D. Carolinianum, stouter even than any

AND RELATED SpECIES 586

of the species given below. As the seeds are not known, the
species may not belong to the group with squamellate seeds.

7. Delphinium Carolinianum Walt. Car. 155. 1788

D. azureiun Michx. Fl. Bor. Am. i : 314. 1803.

D. Carolinianum is confined to the South, ranging from North
Carolina, or perhaps Virginia, to Missouri and south to Florida,
Lousiana and Arkansas. It is characterized by the slender habit,
the small deep blue flowers, the long straight narrow raceme, the
slender and slightly curved and usually horizontal spur.* The
seeds are quite different from those of the related species, being
smaller, only 1.5 mm. long, with broad wing-margins on the
angles and only slightly squamellate on the sides. The following
specimens are in the herbaria of the New York Botanical Garden
and Columbia University.

Georgia: Augusta, 1888, G. McCarthy; Flint River at Al-
bany, i%g^, John K. Small.

Florida: Jackson Co. (collector not given); A. W. Chapman
(locality not given) ; West Florida, Chapman.

Alabama: .Sz^^/^/^j (locality not given); Milstead, 1896,/.. M.
Underzvood ; Auburn, i^gy, Earle & Baker.

Mississippi: Agency, 1897, .S. M. Tracy.

Missouri: Swan, 1898, B. F. Bush, 188.

Arkansas : Nuttall.

8. Delphinium vimineum Don ; Sweet, Brit. Fl. Gard. ser. 2, //.

374

The flowers are similar to the preceding, but the plant is taller,
often over a meter high, with slender lax branches. The most
striking difference, however, is found in the seeds. These are
scarcely angled at all, but with exceedingly strong transverse
lamellae and over 2 mm. long. It is a species confined to the
Gulf Region.

Texas : Driimmond.

Louisiana : Alexandria, and Red River, Hale.

* The specimens from Missom-i cited here have almost erect spm-s.

587 Rydberg : Delphinium Carolinianum

9. Delphinium Geyeri Greene, Erythea, 2 : 189

This species is characterized by the strigose-canescent pubes-
cence, the ascending instead of erect pedicels, the large blue flow-
ers, the rather large bractlets, which often are half as long as the
sepals and borne close under them. The species would not be
placed in this group if it did not have squamellate seeds. These
are, however, only finely so, rather large, about 3 mm. long,
black, irregularly angled, but not wing-margined. The following
specimens are in our herbaria :

Wyoming: Cheyenne, 1872, E. L. Greene; Laramie River,
1894, Ave7i Nelson, /(.oo.

Colorado: Fort Collins, 1896, C. F. Baker.

10. Delphinium Wootoni sp. nov.

Perennial from a deep woody root, 1.5-2 dm. high, finely
grayish-strigose ; basal leaves rather numerous, grayish-strigulose,
rather firm, 3-5 -cleft to the base, the divisions cuneate in outline,
cleft beyond the middle into oblong or linear acute leaves ; stem-
leaves similar, very few ; raceme short, rather few-flowered ; ped-
icels erect, 1—2 cm. long ; bractlets linear, close under the calyx,
the latter light blue or white and tinged with blue or purple :
petals white, the upper ones tinged with yellow ; seeds unknown.

This is closely related to D. albescens, but the position of the
bractlets, the grayish pubescence, the low stout habit and the firm
leaves with broad segments indicate some relationship with D ,
Geyeri. In this, as in several other species, mature seeds are a de-
sideratum.

New Mexico: Organ Mountains, 1893, E. 0. Wooton (type).

Arizona : Ft. Huachuca, T. E. Wilcox.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 5

STUDIES ON THE ROOKY MOUNTAIN

FLORA— I

BY P. A. RYDBERG

NEW YORK
1900

[Reprinted from the Bulletin of the Toreey Botanical Club, 27 : 169-139. 21 Apr., 1900,1

Studies on the Rocky Mountain Flora.— I*

By p. a. Rydberg

(With Pi ates 5 and 6.)

SPECIES OF SENECIO OF THE LOBATUS, AUREUS, SUBNUDUS AND TOMENTOSUS GROUPS

It may seem strange that I present here a paper on Sexecio,
when it is well known to me that Mr. Greenman -is occupied in
preparing a monograph of the genus in all North America, from
the arctic regions to the Isthmus. My work was begun a year ago,
and before I knew of Mr. Greenman's work. I have not been able
to present my results in print before now, and I do it with the
good will of the gentleman mentioned, and with the understanding
that I confine myself to the Rocky Mountain region.

The four groups treated here are closely related and grade
into each other. They might have been treated as a single
group but even this would have been more or less artificial and
ill-defined, because there are several intergradations with related
groups. The more foliose species of the Aurei as S. platylobiis
and S. Idahoensis described below connect with the Eremophili ;
5. cyvibalarioidcs with the Alpicolae, and 5. canovirens with the
Cani.

The work presented here is based on my own studies in the
field and the specimens found in the herbaria of the New York
Botanical Garden, Columbia University and College of Pharmacy,
all in New York City.

LOBATI

Annuals or biennials or perennials with a taproot, perfectly
glabrous in age or slightly floccose at the bases of the leaves, more
or less leafy throughout, 3 dm. or more high : leaves, all except

*The author intends to publish under this title a series of papers on the Botany of the
Rocky Mountain Region. The intention is not however to limit these studies to the
botany of the mountains proper, but will include also the Great Plains to the eastward.
They will comprise the following states and territories : eastern British Columbia, Alberta.
Saskatchewan, Assiniboia, Montana, Idaho, Wyoming, Colorado, New Mexico, east-
ern Utah, and the extreme western portions of the Dakotas, of Nebraska, Kansas, Okla-
homa and Texas.

(169)

170 RvDBERG : Studies ox Rocky Mountain Flora

the very first, pinnately divided : heads rather small and minutely
calyculate. In habit resembling the Aurei and Tomentosi, to
which the last two form a transition, but have more divided leaves
and a root of less duration.

Leaves thin ; root annual or biennial.

Head about l cm. high ; tenninal segments of the basal leaves 3-5 cm. wide.

1. S. sangiiisorboides.
Head 5-8 mm. high ; terminal segments of the basal leaves less than 2 cm. wide.

Basal leaves with few segments ; segments crenate or dentate.

2. S. Greggii.
Basal leaves with numerous segments ; segments lobed or cleft.

3. ^". inillelobalw.
Leaves rather thick ; root perennial.

Bracts thick, much shorter than the disk. 4. S. niuUilobahts.

Bracts thiii, almost equaling the disk. 5. S. Nelsonii.

I. Senecio sanguisorboides sp. nov.

Tall and simple, perfectly glabrous, annual or maybe biennial :
stem terete, about 6 dm. high, leafy : basal leaves thin, 1-2 dm.
long, petioled, pinnately divided with 3-5 segments ; terminal seg-
ments reniform, 3—5 cm. wide, coarsely crenate ; lateral segments
almost orbicular, crenate : lower stem leaves similar ; the upper
with 7— II segments, short-petioled or the uppermost subsessile ;
the base of the petioles with large round, laciniate auricles ; ter-
minal segment ovate, incised-crenate ; the lateral ones obovate
or broadly cuneate : cyme rather contracted : heads about i cm.
high; bracts 12-16, linear-lanceolate, acute, about i mm. wide,
with membranous margins ; the calyculate ones very few and
minute, lanceolate : rays about lo, 8 mm. long and 3 mm. wide,
about 4-nerved : achenes glabrous and angled.

This has been referred to .S". Sangiiisorbae DC; but a compari-
son with De Candolle's description shows several discrepancies.
According to the description S. Sangiiisorbae should have 6nly
8-IO bracts and 5 rays, the leaves should be puberulent beneath
and the terminal 'segment orbicular, characters not found in the
present species. The latter grows at an altitude of 2500-3000

m. [Plate 5, f. 14.]

New Mexico: Santa Fe Caiion, 1897, A. A. & E. Gertrude
Heller, j8 20 (type in the herbarium of the N. Y. Botanical Garden) ;
White Mountains, 1897, E. O. IVoofoji, ^g^.

2. Senecio Greggii sp. nov.
Sefiecio Tanipicamis A. Gray, PI. Fendl. 109. 1849; not DC,
1837; 5. lobatiis A. Gray, PI. Wright. 2 : 99. 1852; not
Pers. 1807.

RvDBERG : Studies ox Rocky Mountain Flora 171

Biennial or annual, with several stems from the base, perfectly
glabrous in age : stems about 3 dm. high : leaves pinnately divided
or the very first ones simple : basal leaves with 3-5 segments, the
terminal one round and crenate, about i cm. in diameter ; the
lateral ones small, obovate ; stem leaves with 7-9 small segments
and more or less auricled at the base of the very short petiole :
cyme compound : heads rather man\', only 5-7 mm. high ; bracts
about 20, linear-acuminate ; the calyculatc ones minute and few :
rays 8-10, about 3 mm. long and 1.5 mm. wide, 3-nerved :
achenes angled and strigose.

This is nearest related to S. lobatus but smaller and characterized
by the many stems from the same root, the few segments of the
basal leaves, the small heads and the narrow bracts. Only
Wright's specimens were collected in the region here treated.
[Plate 5, f 8.]

Mexico : Santa Rosalia, Dr. Gregg (type in Torrey Herba-
rium).

Texas: Near El Paso, 1851-2, C. Wright, 1413 ; between
Frio and Nueces Rivers, 1880, E. Palmer, 754; Loredo to Frontera.
WrigJit (Mexican Boundery Survey), ^jp, in part.

3. Senecio millelobatus sp. nov.

Senecio Tampicmius A. Gray, PI. Wright. 2 : 99. 1852 ; not DC.

1837 ; S. mnltilobatiis A. Gray, Syn. Fl. i" : 394, in part. 1884 ;

not T. & G. 1 849.

Annual or biennial, perfectly glabrous or slightly floccose at
the base of the leaves : stem branched, 3-5 dm. high, very leafy :
all leaves pinnately divided, 3-7 cm. long, lanceolate or oblanceo-
late. in outline ; segments 13-23, obovate, 3-10 mm. long, lobed or
cleft into oblong or ovate lobes ; the upper segments more or less
confluent : cyme compound : heads about 8 mm, high ; bracts ob-
long-lanceolate, acuminate, 12-15 "I number ; calyculate ones
minute and few : rays 5-8 mm. long and 1.5-2 mm. wide : achenes
very scabrous on the angles.

This is nearest related to the preceding and to 5. lobattts but is
easily distinguished by the numerous and lobed or cleft segments.
It may be related to 5. Tampicanus which I have not seen, but that
species is described as having onl}^ 4 or 5 pairs of segments and
glabrous achenes. [Plate 5, f. 11.]

New Mexico: Hills on the Limpia, 185 1-2, C. Wright, 1287
(type in the Torrey herbarium) ; P. V. Le Roy.

172 RvDBERG : Studies on Rockv Mountain Flora

Texas: Lower Rio Grande, 1852, Pany (Mexican Boundary
Survey), 6^8.

4. Senecio multilobatus Torr. & Gray; Gray, PI. Fend. 109.

1849

The type of this species is in the Torrey herbarium and very
unhke the plant regarded by Gray as ^\ iiiultilobatiis. It is char-
acterized by its fleshy leaves and stands perhaps nearer to vS.
conipactus than to the group with which it was associated by Dr.
Gray. I have placed it in this group on account of its pinnatifid
basal leaves. The earliest of these are, however, entire, in the
same manner as they occasionally are in 5". rosjdatiis. This analogy
and the close relationship to 5. conipactus undoubtedly made Prof.
Greene name Baker, Earle and Tracy's specimens " Senecio com-
pactus R}'db., verging towards S. Fendleri." These specimens
differ from Fremont's plant only in the fact that the basal leaves
are shorter and less divided. Eastwood's specimens are exactly
like the type. The base of these specimens shows that the plant is a
perennial rather than an annual as stated in the original descrip-
tion. It grows at an altitude of about 2000 m. [Plate 6, f. 11.]

Utah : Uintah River (a tributary of Greene River), Fremont,
j^p (type in the Torrey herbarium) ; Ogden, 1871, Hayden Sur-
vey ; South Utah, J. E. Jolinson.

Colorado: Grand Junction, 1892, A. Eastivood ; Mancos,
1898, Baker, Earle & Tracy, 4-^6 ; Dolores, 1892, C. S. Crandall;
South Park, 1871, W. M. Canby.

5. Senecio Nelsonii Rydb. Bull. Torr. Club, 24 : 483. 1899

Prof Greene in Pittonia, 4 : 112, devotes over a page to this
species, claiming it to be the same as S. Fendleri, and criticises
both Prof. Nelson and myself. We had, however, both investi-
gated the matter thoroughly before the species was published. In
claiming that the two species are the same. Prof. Greene must either
not know one of the plants or both, or else do it for the purposes of
finding fault. 5. Nelsonii has the leaf form of 5". Fendleri, but
there ends the similarity. In the former the caudex is short, not
woody, and with numerous fibrous roots, placing it nearer to S.
nuiltUobatiis and Ji. conipactus, while 6". Fendleri has a very thick

RvDBERG : Studies on Rocky Mountain Flora 173

and woody rootstock. The heads of ^. Nelsonii are larger, 8—10
mm. high, and the bracts are thin and in anthesis almost equal
the disk, while in vS. Fendlcri they are thick and much shorter.
The former also lacks the fine tomentum of the latter ; it is merely
floccose when young in the manner of 5. nudtilobatiis and .S". com-
pactiis. S. Nchonii is dark green while 5". Fcndleri is more or less
yellowish. Neither of the species forms mats, as Prof Greene
states. Heller's specimens, from the type locality of .9. Fcndleri,
agree fully with Gray's description of that species, except that the}-
are more glabrate than the type. They represent a plant of several
season's growth, but show nothing to indicate a matted plant. I
think that Prof Greene has here confused 5. rosidatus with the
two. Prof Greene's conception of ^. Fcndleri must be very com-
prehensive, indeed, as he also includes in that such forms as 6".
siibciuicatus, S. canovire?is, and another species, nearly related to
S. fastigiatiis Nutt. This statement is founded on specimens de-
termined by Prof Greene only a year or two ago and found in the
herbarium of the New York Botanical Garden. By comparing
Plate 5, Fig. 6, and Plate 6,' Figs, i, 3, 4 and 9, with each other,
one can get an idea of Prof Greene's conception of 6". Fcndleri.
I hav^e seen no specimens of S. Nelsonii except the type.

AUREI

Slender perennials, over 2 dm. high, with a short erect caudex
or rootstalk, glabrous or slightly floccose when young, the wool
remaining in age only at the oase of the leaves : basal leaves entire,
merely toothed : stem leaves more or less pinnatifid and the upper
more or less reduced : heads small, cymose, with very small and
few calyculate bracts.

Heads radiate.

Leaves thick, more or less fleshy.

Basal leaves nan'owly oblanceolate.

Plant low, less than 2 dm. high, often with several stems from the base;
basal leaves short-petioled.
Cyme dense ; upperleaves generally pinnatifid with narrow lobes.

6. ^^ compactus.
Cyme more open ; stem leaves entire or merely crenate.

Basal leaves subentire or 3-toothed at the apex.

7. S. tridenticulatiis .
Basal leaves crenate. 8. S. oblanceolatus.

174 Rydberg : Studies on Rocky Mountain Flora

Plant about 4-6 dm. high, simple.

Leaves serrate or subentire ; basal leaves i cm. or more, slender-

petioled. 9. S. longipetiolatiis.

Stem leaves with long, triangular lobes ; basal leaves 5 cm. or less.

10. S. fulgens.
Basal leaves cuneate, spatulate or broadly oblanceolate, subentire at the base.
Lower stem leaves spatulate with a broad-winged petiole ; upper ones

sessile and with much enlarged bases. II. S. crocattts.

Stem leaves neither broad-winged nor with much enlarged bases.

Basal leaves crenate above the middle. 12. .S'. cyinbalarioidcs.

Basal leaves angularly dentate. 13. .V. Jonesii.

Basal leaves sharply dentate or serrate above the middle.
Lower stem leaves cuneate, plant 3-4 dm. high.

14. S. siibinneatus.
Lower stem leaves oblanceolate ; plant about 2 dm. high.

15. S. acutidens.
Leaves thin.

Basal leaves ovate or more commonly cordate, serrate. 16. S. psciidaiircus.

Basal leaves obovate or oval, crenate or sinuate-dentate.

Stem leaves ovate in outline, with broad segments.

17. S. platylobiis.
Stem leaves lanceolate or oblanceolate in outline, jj'ith narrow segments.

18. S. Jiavovirens.
Stem leaves spatulate or oblanceolate in outline, with short and broad

segments. 19. 5. aiirellus.

Head discoid.

First basal leaves reniform. 20. 5. nephrophylhts.

None of the leaves reniform.

Plant stout ; stem leaves with broad laciniate segments.

21. 6". Idahoensis.
Plant slender ; stem leaves with narrow segments.

Plant yellowish or light green ; bracts not purple-tinged.

17. S. Jlavovii'ens.
Plant dark green.

Basal leaves crenate ; head 8-10 mm. high ; bracts purple-tinged,
linear. 22. S. paticifloriis.

Basal leaves wavy ; heads about 6 mm. high ; bracts broadly lanceo-
late or oblong, not purple-tinged. 23. S. fedifolius.

6. Senecio compactu.s (A. Gray) Rydb. Mem. Torr. Club, 5 :

342. 1893
Senecio aureus var. coinpactus A. Gray, Syn. Fl. i' : 391. 1884.
Dr. Gray's description covers two if not three species. For
which of these .S". covipatus should be used is a matter of ques-
tion. I have adopted it for the plant of the plains with a dense
cyme and usually pinnatifid-dentate stem leaves. Dr. Gray evi-
dently had this plant in mind when he adopted the name coin-

RvDBERG : Studies ox Rocky Mountain Flora 175

pactits. The description seems to indicate mostly this species,
which also I had in mind when I raised the variety to specific
rank and it is this that is described and figured in Britton and
Brown's Illustrated Flora. On the other hand if synonomy and
the citation of specimens are taken in consideration, the name may
have been applied to the next, for the first synonym and the first
specimen cited belong to that species. [Plate 5, f 15.]

Nebraska : Valentine, /. M. Bates, j^ ; Lewellen, G. D.
SivcL'sey, 82 ; Fort Niobrara, 1888, T. E. Wilcox; Thedford,
1893, P. A. Rydbcrg, r^ii ; Platte Bottom, 1891, 211.

Colorado: Plains, \Z^2, Allen & Brcivster ; Colorado Springs,
I ^g2, Isabel Mil I ford.

7. Senecio tridenticulatus sp. nov.
Senccio aureus var. borealis A. Gray, PI. Wright, i : 125. 1852 ;

not T. & G. 1843 ; S. aureus var. coinpactus A. Gray, Syn. Fl.

I^: 391 ; in part.

Perennial with a branched caudex, in age perfectly glabrous,
or slightly floccose at the base of the leaves : stems very slender,
about 3 dm. high : basal leaves very narrowly oblanceolate, thick
with a slender petiole, slightly 3-toothed at the apex or wholly
entire, 4-8 cm. long and 4-5 mm. wide : stem leaves linear and
subentire : cyme open and corymbiform : heads 7-8 mm. high :
bracts lanceolate, acute 73 or 3/ as long as the disk : rays light
yellow, about 8 mm. long and 1-1.5 mm. wide, 3-nerved ; achenes
hispid-puberulent on the angles.

This species differs, from the preceding in the slender stem and
the open cyme, and also in the form of the leaves. The latter
character also separates it from the next following. It grows in
wet sandy soil. The type was growing at an altitude of 2400 m.
Wright's specimens are past blooming, and good characters could
not be taken from them, wherefore I have made Sheldon's speci-
mens the type. The latter were mixed with some of the next.
[Plate 5, f. 12.]

Colorado : Cottonwood Creek, Buena Vista, 1892, C. S. Shel-
don (type in the herbarium of N. Y. Botanical Garden).

Texas: Mountams beyond the Limpia, 1849, Wright, 40^.

8. Senecio oblanceolatus sp. nov.

Perennial with several stems from a strong root, in age glabrous
or rarely slightly floccose at the base of the leaves : stems short and

176 Rydberg : Studies ox Rocky Mountain Flora

stout, about 2 dm. high : basal leaves oblanceolate, thick and fleshy,
4-6 cm. long, obtuse, crenate with entire long tapering bases or
the first subentire : stem leaves similar or reduced and linear in
outline : cyme corymbiform, not compact : heads about 8 mm.
high ; bracts linear acute, yellowish green ; the calyculate ones
minute and few: rays 5-6 mm. long and about 1.5 mm. wide,
4-nerved : achenes minutely scabrous on the angles.

This species has also been included in 5. aureus compactiis, but
lacks the dense inflorescence of S. compactiis as here understood.
The stem leaves never show any indication of being pinnatifid with
narrow lobes as in that species. .S. covipactus is a plant of the
Great Plains, while 6". oblanccolaUis is a mountain plant growing
at an altitude of 1800-3000 m. [Plate 5, f 9.]

Colorado: Como, South Park, 1895, C. S. Crandall (type
in the herbarium of N. Y. Botanical Garden) ; El Paso County,
1897, A. A. & E. Gertrude Heller, 3308 ; Cottonwood Creek,
])uena Vista, 1892, C. S. Sheldou.

9. Senecio longipetiolatus sp. nov.

A tall, simple, perfect!}^ glabrous perennial, with a short erect
rootstock : stem strict, 3-6 dm. high, terete : basal leaves oblan-
ceolate, I -1. 5 dm. long, with a slender petiole, from serrate to
subentire : lower stem leaves similar ; the upper reduced, lanceo-
late, sessile, sharply serrate, or laciniate-dentate, often auricled at
the base : cyme dense, corymbiform : heads 8-9 mm. high :
bracts about 20, linear, acute : the calyculate ones few, minute,
subulate and crisp: rays dark orange, 4—7 mm. long and 1.5 mm.
wide, 3-4-nerved : achenes strongly angled, glabrous.

In the type the basal leaves are strongly serrate, but in the
Colorado plants they are indistinctly so or subentire. Baker,
Earle & Tracy's specimens were named by Professor Greene,
Senecio crocatus Rydb. but it is entirely distinct from anything
collected by Hall & Harbour, and has .nothing that fits any de-
scription of 6". aureus var. croceus, except the dark rays. See
further remarks under vS". crocatus. S. longipetiolatus grows at
an altitude of 2000-3000 m. [Plate 6, f. 10, loa.]

Wyoming: Spread Creek, Teton Forest Reserve, 1897, F.
Tzveedy, 383 (type in the herbarium of N. Y. Botanical Garden).

Colorado : Medicine Bow Range, 1891, C. S. Crandall ; Ha-
mor's Lake, north of Durango, 1898, Baker, Earle & Tracy, 623.

Rydberg : Studies on Rocky Mountain Flora 177

lo. Senecio fulgens sp. nov.

Simple and glabrous perennial with a very short caudex :
stem about 3 cm. high : basal leaves oblanceolate, thick and some-
what fleshy, with the petiole about 5 cm. long, dentate or suben-
tire : lower stem leaves spatulate with a winged petiole, coarsely
dentate ; upper stem leaves sessile with an auricled base, lobed
with triangular or triangular lanceolate lobes, acute : cyme corym-
bose and rather dense : heads about 8 mm. high : bracts about
I 5, oblong, acute, with a broad membranous margin ; the calyculate
ones few, lanceolate : achene striate, glabrous : rays 4-5 mm. long
and 2 mm. wide, dark orange.

Closely related to the preceding, but differing in the short basal
leaves and the lobed upper stem leaves and also in the fewer and
broader bracts. It grows at an altitude of about 2700 m. [Plate

^,^. 1 3-]

Wyoming: Grand Creek, Teton Forest Reserve, 1897, F.

Tivecdy, 584. (type in the herbarium of N. Y. Botanical Garden).

II. Senecio CROCATUS Rydb. Bull. Torr. Club, 24 : 299. 1897
Senecio aureus var. croceus A. Gray, Proc. Acad. Nat. Sci. Phila.

1863: 68, 1863 ; not 5. croceus DC. 1837.

When the above name was published I did not know that this
variety of Gray's was a complex one. Hall & Harbour's no. 332,
which is the type, consists of two different things ; but as one of
them is rayless it can not be taken as the type of the var. crocaius,
which was named from its orange rays. That I did not draw a
new diagnosis, I admit, was perhaps careless, but this blunder I
think was not worth a page and a half of discussion as it was given
by Professor Greene in Pittonia, 4 : 114-116. I committed just
the same mistake as Professor Greene himself in establishing An-
tennana media Greene, Pittonia, 3 : 286. What Professor Greene
says of Mr. E. Nelson in Pittonia, 4: 85 can be applied to himself.
Gray's description of the var. croceus in the Proceedings of the Phil-
adelphia Academy is perhaps not adequate, but this description is
supplemented in the Synoptical Flora and -elsewhere and I think
that Senecio aureits var. croceus Gray is amply published according
to all rules we have. If so, Senecio croceus Rydberg is not a nomen
nudum, whatever Professor Greene may say. Professor Greene in
1897 or 1898 accepted my name, for he named Baker, Earle &

178 Rydberg : Studies on Rocky Mountain Flora

Tracy, no. 625, ^'. crocains Rydberg. This mistake is just as un-
pardonable because that plant shows none of the characters, assigned
to 6". mircus var. croceiis Gray, except the color of the rays, and this
character Professor Greene, agrees with nie in regarding as of little
value. That Professor Greene and other botanists may know what
I now mean by S. crocatus I shall give a diagnosis, here following
Professors Greene's example in the case of Antcniiaria media.

A glabrous perennial with a short erect rootstock : stem
1.5-3 ^^^- high : basal leaves 2-3 cm. long, obovate or spatulate,
crenate or subentire with a winged petiole : lower stem leaves
similar but with broader, winged petioles which are somewhat
auricled at the base, or else oblong without distinction between
blade and petiole and then more auricled : upper stem leaves
ovate or triangular with very large and large-toothed aiiricles :
cyme small and compact with 2—5 mm. heads, which are 8—10
mm. high : bracts about 20, linear : rays 7-8 mm. long and
1.5-2 mm. wide, orange to pale yellow, achenes striate, glabrous.
[Plate 5, f. 13.]

Colorado : Middle Park, 1862, Hall & Harbour, j;j2, in part
(type) ; J2g, in small part; 1868, Geo. Vasej (Powell's Expedi-
tion), j^o i) ; South Cottonwood Gulch, 1892, C. S. SJieldon ;
Gray's Peak, 1872, John Torrey ; (?) Little Kate Mine, 1898,
Baker, Earle & Tracy, ^Sg;"^ South Park, 1871, Caiiby ;
Long's Peak, 1886, Lettcrman (depauperate).

Wyoming: La Plata Mines, 1895, Aven Nclso)t, lySg."

Senecio DiMORPHOPHYLLUS Greene, Pittonia, 4 : 109. 1900

I have not seen any specimens of this species, and have not
been able to include it in the key. It is described as being a foot
(about 3 dm.) high, light green and with long golden-yellow rays.
Otherwise the description reads much like that given above for
6". crocatus. The type was collected about Pagosa Peak, Colo., in
1899, by C. F. Baker.

12. Senecio cymbalarioides Nutt. Trans. Am. Phil. Soc. II., 7:

412. 1841
Senecio aureus borcalis Torr. & Gray, Fl. 2 : 442. 1843 ; vS\ aureus
obovatiis Eat. King's Exp. 5 : 190, in part. 1871.

* These specimens were named Senecio hctcrodoxus Greene n. sp., but I can
not find any published description. They differ from the rest in being cespitose, with less
marked differentiation in the leaves and with traces of floccose pubescence at the bases
of the leaves and heads. The species may be distinct.

Rydberg : Studies on Rocky Mountain Flora 179

This has sometimes orange rays and has then been confused
with S. crocatus, but it lacks the winged petioles of the lower
leaves and the large auricles of the upper. It often reaches a
height of 3 dm. and then many of the stem leaves are also spatu-
late. Such a form with more permanent woolliness is Watson's
no. 669; otherwise I cannot distinguish it from 5. cymbalariodes.
[Plate 5, f. I.]

British America: Mackenzie River, 186 1-2, /. S. Onion.

Montana: Park Co., 18S7, Tweedy, j^j, in part; Silver Bow
Co., Mrs. Moore ; Jack Creek, 1897, Rydberg &■ Bessey, ^266.

Wyoming: Beaver Caiion, 1895, Rydberg; Bacon Creek,
1894, Aveji Nelson, go6.

Idaho : Mt. Chauvet, 1897, Rydberg &- Bessey, j26y.

Utah : Wasatch Mts., 1869, 5. Watson, 66 g.

Washington: Mt. Paddo, 1882, W. N. Snksdorf ; Columbia,
Niittall (type).

13. Senecio Jonesii sp. nov.

A glabrous perennial : stem about 3 dm. high, slender, terete :
basal leaves rounded-obovate, sinuately toothed, sometimes with
a pair of small lobes at the base ; their petioles often tinged with
red : stem leaves oblanceolate in outline, pinnatifid with oblong
segments, generally short-petioled : cyme corymbiform : heads
about 8 mm. long: bracts 12-15, lanceolate, acuminate, about 2^
as long as the disk, membranous margined ; calyculate ones few,
linear : rays about 5 mm. long and 2 mm. wide, 4-nerved : achenes
very strongly striate, glabrous.

Nearly related to the preceding, but differing in the toothing of
the leaves and the form of the bracts. The type was collected at
an altitude of 3300 m. [Plate 6, i. 5.]

Utah: Alta, Wasatch Mountains, 1879, M. E. Jones, 11 25
(type in the Columbia Herbarium).'

14. Senecio subcuneatus sp. nov.

Perennial with a branched caudex, slightly floccose when
young, glabrate in age : stems 3-4 dm. high : basal leaves about
5 cm. long, spatulate or cuneate, dentate or serrate above the
middle, entire at the base and tapering into a short petiole, rather
thick : lower stemleaves narrowly cuneate, short-petioled, dentate
at the apex : upper ones linear and sessile, often sharply dentate :
cyme corymbiform : heads about 8 mm. high: bracts about 15,

180 Rydberg : Studies ox Rocky Mountain Flora

lanceolate, acute, shorter than the disk : rays 4-5 mm. long and
1.5 wide, 4-nerved : achenes striate, glabrous.

This species is nearest related to S. cymbalarioides but is taller,
with narrower basal leaves, which have sharper teeth ; the heads
are also much more numerous. It grows at an altitude of 2000-
2500 m. Baker, Earle and Tracy's specimens were labeled by
Prof. Greene Scnecio Fcndleri, approaching S. couipactiis. This is
strange from one that claims that he has known ^. Fendlcri since
1870.* This plant has little in common with that species or with
5. Nelsonii Rydb. [Plate 5, f. 6.]

Colorado : Grizzly Creek, 1896, C. F. Baker (type in the
herbarium of N. Y. Botanical Garden); Lake City, 1878. H. IV.
Pease (depauperate) ; Mancos, 1898, Baker, Earle & Tracy, 6j.

15. Senecio acutidens sp. no v.

Perennial, with a thick woody rootstock and short caudex, in
age glabrate or slightly floccose at the base of the leaves : stems
several, about 2 dm. high, angled, more or less tinged with red :
basal leaves about 5 cm. long, thick, fleshy and somewhat glau-
cous, obovate or spatulate : sharply dentate above the middle, at
the base entire and abruptly contracted into a slender petiole :
lower stem leaves similar or oblanceolate and acute ; the upper
reduced, linear, laciniate-dentate or somewhat pinnatifid : cymes
corymbiforni and rather dense: heads 8-10 mm. high: bracts
broadly linear, acute, 7^ or ^ as long as the disk : rays about
5 mm. long and 1.5 mm. wide, 3-4-nerved : achenes slightly an-
gled, glabrous.

Nearest related to ^. cymbalarioides but the leaves are thicker,
more glaucous and very acutely dentate. The perennial rootstock
and caudex are also thicker and more woody. [Plate 5, f 2.]

Wyoming : Union Pass, 1 894, Aven Nelson, 8j8 (type in the
herbarium of N. Y. Botanical Garden).

16. Senecio pseudaureus Rydberg, Bull. Torr. Club, 24: 298.

1897

This is nearest related to the eastern 5. aureus and has the
same subcordate, thin, basal leaves, but these are distinctly serrate,
instead of crenate. It is the most common species of the group
in the Rockies, growing in wet meadows at an altitude of looo-
3000 m. [Plate 5, f 10.]

* See Pittonia, 4: 112.

RvDBERG : Studies on Rockv Mountain Flora 181

Montana : Little Belt Mountains, 1896, Flodman, gi8 (type);
Madison Co., Mrs. McNulty ; Bear Gulch, 1887,/': Tweedy, 340;
Columbia Falls, Mrs. Kennedy, g ; Spanish Basin, 1897, Rydberg
&■ Bessey, 5263 ; Indian Creek, 3264.

Wyoming : Lone Star Geyser Basin, Yellowstone Park, 1 897,
Rydberg & Bessey, 3262.

Colorado: Grizzley Creek, 1896, C. F. Baker; Long's Peak,
1886, G. IV. Letterman ; Mancos, 1898, Baker, Earle & Tracy,

45-

New Mexico: Pecos River, 8 miles east of Glorietta, 1897,

A.A.&E. Gertrude Heller, 3682.

Utah: E. Humbolt Mts., 1868, S. Watson, 667.

17. Senecio platylobus sp. nov.
S. aureus var. croceus D. C. Eaton, King's Exp. 5 : 190, in part.

1871 ; not Gray. 1863.

A stout, perfectly glabrous perennial : stem about 4 dm. high,
rather leafy, striate and somewhat angled : basal leaves 6-10 cm. .
long, slender-petioled, obovate or broadly oval, sinuately crenate-
dentate, thin : lower stem leaves oblanceolate, petioled, deeply
lobed or divided with rounded lobes : upper stem leaves broadly
ovate in outline, sessile, pinnately divided into broadly oblong or
cuneate divisions which are more or less deeply sinuate-dentate :
cyme compound ; its branches corymbiform : heads about 8 mm.
high ; bracts broadly lanceolate, acute, membranous margined and
slightly shorter than the disk : rays 6--/ mm. long, and 3 mm.
wide, 5 -nerved : achenes striate, glabrous.

Probably nearest related to the preceding, but easily distin-
guished by the form of the leaves and the broad rays. It grows
at an altitude of about 1500 m. [Plate 6, f 8.]

Utah : Wasatch Mountains, 1869, S. Watson, 671 (type in the
Torrey Herbarium).

18. Senecio flavovirens sp. nov.
^. Balsamitae Rydberg, Mem. N. Y. Bot. Card., i : 446, mainly ;

1900 ; not Muhl. 1804.

Light or yellowish green, slender, perennial, in age glabrate or
slightly floccose at the base of the leaves : stem 3-4 dm. high,
striate, pale : basal leaves 3-8 cm. long, obovate or broadly oval,
generally tapering into the petioles, but sometimes truncate at the
bases, obtuse, crenate or sinuate, light green : lower stem leaves ob-

182 Rydberg : Studies on- Rocky Mountain Flora

lanceolate in outline and petioled ; the upper lanceolate or linear in
outline and sessile ; all deeply pinnatifid with narrow, oblong or
linear segments.: cymes contracted, corymbiform : heads 7-8 mm.
high ; bracts linear, acute, yellowish-green, and occasionally with
brownish tips, a little shorter than the disk ; calyculate ones
few, linear, small and crisp : rays pale yellow, about 6 mm. long
and 1.5 mm. wide, 4-nerved, or very often lacking : achenes hispid-
puberulent on the angles.

Nearest related to the eastern 5. Balsamitac, but characterized
by its yellowish green color and a more contracted cyme. It
grows at an altitude of 2000-3000 m. [Plate 5, f 4.]

Idaho: Beaver Canon, 1895, /?y<^(^^7'^ (type in the herbarium
of N. Y. Botanical Garden).

Montana: Deer Lodge, 1895, Rydberg, 28^0 (rayless) ;
Helena, 1887, F. D. Kelscy, ^01.

Wyoming: Buffalo Fork, 1897, Tivecdy, 3S6 ; Laramie Plains,
1889, E. L. Greene (rayless); Green River, 1894, Aven Nelson,
ioj6 (rayless).

19. Senecio aurellus sp. nov.

Perennial with a short rootstock, somewhat floccose when
young, soon glabrate : stem striate, 4-5 dm. high : basal leaves
spatulate or cuneate, dentate-serrate, about 5 cm. long, with a
short petiole, soon glabrous : lower stem leaves long-petioled,
cuneate or spatulate, lyrately lobed, the upper narrowly oblance-
olate and subsessile, slightly auricled at the base : cyme com-
pound but with rather few heads, which are about 8 mm. high :
bracts 12-16, glabrous, yellowish-green, lanceolate, thin ; caly-
culate ones minute, lanceolate : rays golden-yellow, 5-6 mm.
long and 2 mm. wide, 4-nerved : achenes scabrous hispidulous on
the angles.

In the form of the stem leaves this most resembles, perhaps,
.S. rosidatiis, but these are thinner and perfectly green, only slightly
floccose when young. Otherwise it is intermediate between the
eastern 5". Balsamitae and .S". niultUobatus. The type was deter-
mined by Professor Greene as " S. psciuimireiis Rydb., not typical."
To that species it does not have any close relationship. [Plate
6, f 12, 12a.]

Colorado: Mancos, 1898, Baker, Earle & Tracy, gg8 (type
in the herbarium of New York Botanical Garden).

Rydberg : Studies on Rocky Mountain Flora 183

20. Senecio nephrophyllus R\'dberg, Mem. N. Y. Bot. Garden,

I : 446. 1900

Related to the two preceding, but characterized by the ray-
less heads, the reniform first basal leaves which resemble those
of Oxyria digyna, and the pinnatifid stem leaves with their oblong
blunt segments. The Colorado specimens are much lower than
the type, being about 2 dm. high. [Plate 6, f. 15. J

Montana : Big Blackfoot River, 1883, Canby, 20j.

Colorado: South Park, 1872, T. C. Porter; Middle Park,
Hall & Harbour, jj2, in part.

2 1 . Senecio Idahoensis sp. nov.

Stout perennial, in age glabrate or slightly floccose at the
base of the leaves : stem 4-5 dm. high, branched, striate, tinged
with purplish : basal leaves obovate, serrate : lower stem leaves
oblanceolate in outline, about i dm. long, petioled ; the upper
ovate or lanceolate in outline, sessile and auricled at the base ; all
bluish-green, pinnately divided into oblong or oblique-cuneate in-
cised segments : heads numerous, about i cm. high ; bracts about
25, very narrowly linear, equaling the disk : rays none : achenes
striate, glabrous.

A species somewhat related to 5. aureus, but characterized
by its broad and laciniate leaves, rayless heads and numerous very
narrow bracts. [Plate 6, f 5.]

Idaho : Granite Station, 1892, Sanelberg, McDougal & Heller,
80J (type in the N. Y. Botanical Garden herbarium).

22. Senecio pauciflorus Pursh, Fl. Am. Sept. 529. 1814
This species, characterized by its few (2-6) rayless heads, linear
purple tinged bracts, equaling the disk in length, and its small
rounded-oval crenate basal leaves, has been collected several
times in Labrador ; but exactly similar specimens are in the
Torrey herbarium and were collected by Burke in the Rocky
Mountains, but the locaHty is not given. [Plate 5, f 3.]

23. Senecio fedifolius sp. nov.

A delicate glabrous perennial with a very ^hort caudex and a
clump of fibrous roots: stem slender, weak, about 1.5 dm high:
basal leaves 3-5 cm. long with a slender petiole ; blade 1-2 cm.
long, rounded-ovate or broadly oval, wavy or subentire : stem

184 Rydberg : Studies on Rocky Mountain Flora

leaves mostly small, pinnately divided into short oblong segments :
heads 2-3, about 6 mm. high, somewhat turbinate ; bracts 12-15,
broadly lanceolate, conspicuously membraneous margined : rays
none : achene glabrous.

The species is named from the resemblance the basal leaves
have to those of certain species of Valcrianclla which genus has
also been known under the name Fedia. [Plate 5, f 7.]

Colorado : South Park, 1871, W. M. Canbv (type in the her-
barium of the College of Pharmacy, New York).

SUBNUDI

A slender perennial with a long slender horizontal rootstock,
perfectly glabrous, sparingly leafy, monocephalous : basal leaves
broadly obovate, coarsely crenate-dentate : heads decidedly turbi-
nate with narrow linear-acuminate bracts ; calyculate bracts, if
present, few, and half as long as the main series. A single species.

24. Senecio subnudus DC. Prod., 7: 428. 1837
Senecio aureus vzx . subnudus GrSiy , Syn. Fl. i" : 391. 1884.

This species has been included as a variety of 6". aureus, but I
think it should be regarded as the type of a distinct group. The
long slender horizontal rootstock and the decidedly turbinate in-
volucre is not found in any of the aureus allies. The latter char-
acters would place it near S. frigidus. [Plate 6, f 2.]

Washington: Chiquash Mountains, 1892, Suksdorf, 2i6j ;
Cascade Mountains, 1882, Brandegee, iiS.

Oregon: 1882, T. Howell; 1871, Elihu Hall, J04..

California: Butterfly Valley, 1874, Mrs. R. M. Austin.

Montana: Park Co., 1887, Tweedy, j^^ ; Pony, 1897, Ryd-
berg & Bessey, 52 jo.

Wyoming: Wind River, 1842, Fremont; Buffalo Fork, 1897,
Tweedy, ^8 y ; Yellowstone Park, 1884, Tiveedy, 120.

TOMENTOSI

Perennials, over 2 dm. high, with a short caudex or root-
stock, which often is subligneous and cespitose, floccose when
young, becoming more glabrate in age, but with some wool
always remaining : basal leaves, except the very first ones, from
serrate or dentate to pinnatifid : stem leaves always present, but
often reduced, generally pinnatifid or pinnately lobed or toothed :
heads cymose, small, with few and very small calyculate bracts.

26.

5". Piatt en sis.

27-

S. Neo-Mexicaniis.

28,

S. salicinus.

29.

S. canovirens.

Rydberg : Studies on Rocky Mountain Flora 185

The pjroup is closely related to the Aurei and Lobati with
which it is connected through 5. Plattensis and 5. Fendleri respec-
tively. 5. Nclsonii and -S". miiltilobus in young state may also be
sought here.

Basal leaves mostly dentate or crenate.

Basal leaves suborbicular, ovate or cordate ; margins of the petioles arachnoid-

tomentose. 25. S. flavuhis.

Basal leaves obovate or spatulate.

Lower stem leaves acute ; plant dark green.
Lower stem leaves obtuse ; plant yellowish-green.
Basal leaves lanceolate or oblanceolate.

Basal leaves narrowly oblanceolate, seiTate.
Basal leaves lanceolate, sinuate-dentate.
Some of the basal leaves entire or dentate, the rest pinnatifid or pinnately lobed.
Plant conspicuously rosulate-stoloniferous ; stem leaves sinuately round-lobed.

30. S. rosulatiis.
Plant not stoloniferous ; stem leaves pinnatifid with toothed lobes.

26. S. Plattensis.
All leaves pinnatifid with toothed segments.

Caudex subligneous ; plant tomentose ; bracts shorter than the disk.

31. 6". Fendleri.
Caudex not subligneous ; plant glalirate.

Bracts almost equaling the disk. 5. S. Nelsonii.

Bracts much shorter than the disk. 4. S. viultilobus.

25. Senecio flavulus Greene, Pittonia, 4: 108. 1900
I have seen no specimens of this species, but from the descrip-
tion it must be most nearly related to the next, differing in the
more slender habit, the small leaves and their form and the peculiar
arachnoid tomentum on the margins of the petioles. The type
was collected by C. F. Baker at Aboles, Colo., in 1899.

26. Senecio Plattensis Nutt. Trans. Am. Phil. Soc. II. 7: 413.

1841
This species is exceedingly variable as to the leaf form as well
as to the pubescence ; usually the basal leaves are merely serrate
and obovate or oval in outline, but sometimes some of them are
like the lower stem leaves, more or less lyrately pinnatifid. It is
conspicuously floccose when young, but in age the leaves become
almost glabrous and the wool remaining only on the lower part
of the stem and petioles. The species belongs to the region of
tl e plains. [Plate 6, f 14.]

186 Rydberg : Studies on Rocky Mountain Flora

South Dakota: Brookings, 1897, L. W. Carter; Hot Springs,
\^(^2, P. A. Rydberg, 8 28.

Colorado: Fort Collins, 1895, C. S. Crandall.

Indian Territory: Sapulpa, 1895, B. F. Bush, I2jf.g ; False
Washita, 1868, Edi^'. Palmer, ^61.

Kansas: Riley Co., 1895, /. B. Norton, joj ; Manhattan,

1887, W. A. Kcllennan ; Osborn Co., 1894, C. L. Shear, 28 ;
Fort Riley, 1892, E. E. Saj/e, ^j2.

Nebraska: Lincoln, 1887, H. J. Webber; Fort Niobrara,

1888, T. E. ]Vilcox ; Mrs. Austin ; Creete, G. D. Szveesey.
Missouri: St. Louis, i'^a^^, RieJd ; McDonald Co., 1891, i>.

F. BiisJi.

Illinois: Oquawka, 1873, H. N. Patterson.

27. Senecio Neo-Mexicanus a. Gray, Syn. Fl. i": 392. 1884

The original publication of this species is generally given as
Proc. Amer. Acad. 19 : 55, but no description is there to be found.
As far as I can find, the one in the S}'noptical Flora is the first
published. Probably more than one species is included in that
description and in the specimens cited below there are appar-
ently three different types, but it has been impossible for me to
find any constant characters by which to distinguish them. In
one of them the stem leaves are more or less lyrate-pinnatifid and
even the basal ones occasionally have a few lobes on the petiole,
while in the others the stem leaves are narrow and merely toothed.
The tomentum is sometimes deciduous, sometimes almost persis-
tent. It grows at an altitude of 1200-2500 m. [Plate 6, fig. 7.]

New Mexico: Silver City, Pinos Altos Mts., 1880, E. L.
Greene ; between Santa Fe and Canoncito, 1897, A. A. & E. Ger-
trude Heller, 374^.; Organ Mts., 1895, E. O. JVooton ; P. V. LcRoy.

Arizona: Fort Huachuca, 1892, T. E. WUeox ; 1876, E. Pal-
mer, 614 ; Santa Catalina Mts, 1883, C. G. Pringle ; San Francisco
Mts., 1 88 1, H. H. Rusby, 212; Mogollon Mts., 1887, E. A.
Meavns, j8 ; Squaw Creek, 1887, E. A. Mearns, lyi ; Santa Rita
Mts., 1 88 1, C. G. Pringle.

28. Senecio salicinus sp. no v.

A floccose perennial, with a short cespitose'caudex : stems sev-
eral, about 4 dm. high, simple below, striate and floccose : basal

RvDBERG : Studies on Rocky Mountain Flora 187

leaves 5-10 cm. long, narrowly oblanceolate, the first ones sub-
entire, but most of them serrate exxept at the tapering base, finely
tomentose on both sides, but in age more glabrate above : stem leaves
small, linear, sessile, dentate, slightly auricled at the base : cyme
compound : the heads subumbellate at the ends of the branches,
y-S mm. high, slightly floccose at the base : bracts yellowish, thin,
lanceolate, shorter than the disk : rays light )-ellow, short and
broad, 3-4 mm. long and 2 mm. wide : achenes striate, glabrous.

A species related to 5. Fendlcri, but characterized by the nar-
row, merely serrate basal leaves, the small subumbellate heads in
a short, flat-topped inflorescence with widely spreading ultimate
branches. It grows in the foothills at an altitude of about 1800 m.
[Plate 6, f. 6.]

Colorado: J. Laramie Co., 1895, /. H. Cozven (type in the
herbarium of N. Y. Botanical Garden).

29. Senecio canovirens sp. no\'.

Stout perennial with a short erect rootstock, dark green, floc-
cose with more or less deciduous wool : stem 4-5 dm. high, loosely
floccose: basal leaves 5-12 cm. long, short-petioled ; blades lan-
ceolate or oblanceolate, sinuately toothed, dark green, firmly floc-
cose, in age almost glabrate above : lower stem leaves similar ;
upper ones linear, sinuately dentate with rather acute teeth : cyme
with erect or ascending branches : heads 7-8 mm. high, turbinate
campanulate ; bracts 12-14, oblong, linear, floccose at the base
and more or less villose, acute, brownish on the back and with
yellowish margins : rays 5-6 mm. long and 2 mm. wide, at first
orange, in age paler, 4-nerved : achenes glabrous, striate.

The type was determined by Prof. Greene as Senecio Fendleri,
but it is very unlike the typical form of the aggregate that has been
known under that name. The dark green color, the leaves which
are merely toothed, never pinnatifid and still less bipinnatifid as
they often are in ^. Fendleri easily distinguish it from that species.
The perennial caudex and rootstock are similar to that of 5. Fend-
leri but less thick and less woody and, as far as seen from the
specimens, not branched. It grows at an altitude of about 2000
m. [Plate 6, f. 9.]

New Mexico: White Mountains, 1897, F. 0. VVooton, 244
(type in the herbarium of N. Y. Botanical Garden) ; Organ Moun-
tains, -f-gj.

Colorado: Pike's Peak,' 1895, Mrs. S. L. Clarke.

188 Rydberg : Studies on Rocky Mountain Flora

Senecio mutabilis Greene, Pittonia, 4: 113. 1900
As I have seen no specimens of this species I have not been
able to include it in my key. It may be the same as the preced-
ing, but several characters given in the description do not agree
with it, especially the deeply tridentate rays. Professor Greene is
always criticizing other botanists for drawing vague descriptions.
No better example of just such a description can be given than the
one here made by himself.

30. Senecio rosulatus sp. no v.

Senecio aureus var. Balsaviitae A. Gray, Proc. Acad. Nat. Sci.
Phila. 1863 : 68, in part.

More or less tomentose perennial, with a branched rootstock
and generally numerous underground stolons producing leafy off-
sets : stems 2-3 dm. high, floccose or sometimes becoming almost
glabrate in age ; first leaves of the offsets spatulate and entire ;
other basal leaves and lower stem leaves oblanceolate, finely white
tomentose especially on the lower surface, occasionally becoming
more glabrate in age, sinuately pinnatifid with rounded lobes,
petioled : upper stem leaves lanceolate or linear, sessile, auricled at
the base also with rounded lobes : cyme corymbiform, open :
heads small, 5-6 mm. high, floccose at the base : bracts about 12,
broadly lanceolate, acute, yellowish with rather broad membranous
margins, much shorter than the disk : rays about 5 mm. long and
2 mm. wide, 4-5 -nerved : achenes strongly striate, glabrous.

The species has been confused with 6". Fendleri, but is distin-
guished by the more slender rootstock and the numerous branches
forming offsets, the entire first leaves not seen in that species and
the short rounded entire lobes of the stem leaves. The heads are
usually also smaller. It grows at an altitude of 2500-4000 m.
[Plate 6, f. 4, 4a.]

Colorado: Georgetown, 1885, N. H. Patterson yg (type);
Golden City, 1892, E. L. Greene; 1862, Hall &- Harbour, 333,
in part; 1871, W. M. Canby ; Twin Lakes, 1873,/. M. Coulter;
Pike's Peak, 1884, G. IV. Letterman, 260; Caribou, 1891, E.
Penard, 228 ; Fort Collins, 1896, C. F. Baker; Silver Plume,
Gray's Peak, and Georgetown, 1895, P. A. Rydberg.

31. Senecio Fendleri Gray, Mem. Am. Acad. II, 4 : 108. 1849

The typical S. Fendleri is a rather rare plant with a decidedly

woody rootstock and caudex. None of the specimens seen show

RvDUERG : Studies on Rocky Mountain Flora

189

any sign of entire leaves nor of subterranean stolons. The leaves
are often bipinnatifid or pinnatifid with oblong toothed, and acutish
segments. See also the remarks given under ^. Ah'/sonii. [Plate

6. f 3-]

New Mexico: Santa Fe Caiion, 1897, A. A. & E. Gertrude

Heller, ^774.

Colorado: i^y^, Brandegee, ^46.

Utah : Salt Lake City, 1869, S. Watson.

Kxplaiiatloii of Plates

Unless otherwise stated, the drawings represent basal leaf, stem leaf selected a
little above the middle of the stem and the head of each species. The leaves are re-
duced to one half and the heads are of natural size.

Plate 5

1. S. cymbalarioides Nutt.

2. S. acutidens Rydb.

3. S. pauciflorus Pursh.

4. S. Jlavovirens Rydb.

5. 3". Jonesii Rydb.

6. 5. subcuneatus Rydb.

7. S. fedifolius Rydb.

8. S. Greggii Rydb.

1. .S'. Nelsonii Rydb.

2. S. subnudus DC.

3. .S". Fendleri A. Gray.

4. S. rosulatus Rydb.
4a. An offset of the same.

5. S. Idahoensis Rydb.

6. S. salicinus Rydb.

7. S. Neo-Mexicanus A. Gray.

8. 5. platylobus Rydb.

9. S. canovirens Rydb.

9. S. oblanctolatus Rydb.

10. ^9. psettdaureus Rydb.

11. S. viillelohatus Rydb.

12. S. tridenticulatus Rydb.

13. S. crocatiis Rydb.

14. S. sanguisoiboides Rydb.

15. S. compactus (A. Gray) Rydb.

Plate 6

10. .S". longipetiolatiis Rydb. (from

type).
\oa. Basal leaf from Baker, Earle &^
Tracy, 62^.

11. S. multilobatus Torr. & Gray,

12. .S". aurellus Rydb.

12a. Lower stem leaf of the same.

13. S. fulgens Rydb.

14. S. P/at(ensis Nutt.

15. S. nephrophyllus Rydb.

Bui.i.. ToRR. BoT. Ci.rii, 27.

Pi,. 5-

^^^^^'
^^^%'"-

RYDBERG ON SENECIO.

Bi i.i.. Tour. Bdt. Cluu, 27.

Pi.. 6.

RYDBERG ON SENECIO.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN~No. 6

NOTES AND DESCRIPTIONS OF NORTH
AMERICAN PLANTS— MI

BY JOHN K. SMALL

NEW YORK
1900

TReprinted from the Bulletin of the Torrey Botanical Club, Vol. 25, No. 6, June, 1898,

and Vol. 27, No. 7, May, 1900.]

Notes and Descriptions of North American Plants.— 1.

By John K, Small.

Saxifraga Napensis.

Perennial by very short rootstocks, scapose, almost glabrous
or sparingly grandular-pilose above. Leaves basal ; blades thin-
nish (drying very thin), more or less obliquely oval or ovate, 2-7
cm. long, rounded at apex, undulate, abruptly or gradually nar-
rowed at base, ciliate, sometimes slightly pilose on both surfaces :
petioles slightly shorter than the blades or longer, broadly winged :
scapes erect, slender, 1.5-3 dm. tall, loosely-paniculate-corym-
bose : calyx glabrous or glabrate ; tube broadly turbinate, nearly
I mm. high ; segments oblong or ovate-oblong, slightly longer
than the tube, obtuse or acutish, gland-tipped : corolla white, 5
mm. broad ; petals broadly oval or suborbicular, 2 mm. long,
rounded or notched at apex, sessile or nearly so, 5-7-nerved :
ovaries united to calyx-tube, surrounded by a conspicuous lobed
disk : follicles short and stout, 4 mm. high, united to each other
to about the edge of disk : seeds dark red, 0.3 mm. long.

On hillsides, Napa Valley, California.

Collected by J. M. Bigelow (Whipple's Expedition) and George
Thurber, no. 496.

A species of lax habit, resembling Saxifraga claytoniaefolia
more closely than any other member of the subgenus Micranthes.
It differs from 5. claytoniaefolia by its broadly oval or suborbicular
5-7-nerved petals.

Saxifraga Van-Bruntiae.

Perennial, bright green, minutely glandular-pilose or glabrate
below. Stems tufted, 2-7 cm. long, simple, or sometimes corym-
bosely branched above, leafy to the top : leaves alternate ; blades
linear, 4-8 mm. long, leathery, blunt, with a thick apex, sessile,
turning black at the base of the plants : calyx glabrate in age ;
tube broadly turbinate ; segments oblong or ovate-oblong, obtuse,
longer than the tube, spreading : petals yellow, oblong, about
4 mm. long, obtuse, firm, more or less crisped, much longer
than the calyx-segments : stamens 10, filaments filiform.

The original specimens of this hitherto undescribed Saxifraga
were collected by Mr. and Mrs. Cornelius Van Brunt during the

(316)

317 Small: North American Plants

past season on the summit of Sulphur Mountain near Banff, British
Columbia. The species is related to Saxifraga serpyllifolia and
5. ckvysantlia ; the habit suggests those species but the stems are
copiously leafy to the top and the leaves are narrower. The
petals are smaller, of a much less brilliant yellow and oblong or
ovate-oblong instead of oval-orbicular or obovate as they are in
the two related species.

Galpinsia Toumeyl

Perennial from a shrubby base, slender, bright green, puberulent.
Stems branching near the base ; branches erect or ascending, wire-
hke, 1-3 dm. long, usually simple above, leafy, pale when young:
leaves numerous, sometimes clustered in axils ; blades linear-
spatulate to linear, 1-2 cm. long, acute, entire, with midrib promi-
nent beneath, lower ones short-petioled, upper sessile : spikes
few-flowered, leafy -bracted : calices very minutely pubescent ; tube
slender, 3-5 cm. long; segments about 1-5 cm. long, their free
tips 5-6 mm. long : corolla yellow ; petals orbicular-obovate,
1.5 cm. long, undulate : anthers linear, as long as the filaments :
capsules linear-prismatic, 2 cm. long.

Arizona: Chincahua Mountains, July 25, 1894, J. W. Tou-
rney, no. 197. Fort Huachuca, August, 1892, T. E. Wilcox.

The species just described is related to Galpinsia Hartivegii, but
is of a much more slender build. There are minor characters in the
foliage and habit, but one of the more crucial points of difference
lies in the calyx, where we find the free tips of the segments 5-6
mm. in length.

LiMONIUM LIMBATUM.

Perennial, bluish green or glaucescent. Leaves basal ; blades
leathery, spatulate or oblong-spatulate, 0.5-1.5 dm. long, obtuse
or notched at the apex, prominently nerved beneath in drying ;
petioles shorter than the blades or rarely longer, margined :
scapes erect, solitary or several together, corymbosely branched ;
branches zigzag, ascending ; spikes in dense terminal corymbs :
bracts suborbicular or sometimes orbicular-oval, obtuse, often
eroded at the apex, hyaline-margined : calyx trumpet-shaped with
a flaring limb, nearly 4 mm. long ; tube hirsute ; segments broadly
deltoid, apiculate : corollas bright blue.

In alkaline soil, Texas and New Mexico.

As far as I can learn there has been no attempt heretofore to

Small : North American Plants 318

separate the Texan plant referred to Limonunn Californicwii
either varietally or specifically. An examination of considerable
material both from the Texas and the Californian districts discloses
the fact that there have been two perfectly distinct species con-
fused under the old Limonhim Californiaim. As in the case of
all the species of the genus the two under consideration resemble
each other in habit. The diagnostic characters are contrasted

below :

Liinoniiim Calif ornicinn. Calyx narrowly funnelform ; tube
glabrous ; segments erect or nearly so, rounded and mucronulate.

Limoniiim limbatnni. Calyx trumpet-shaped ; tube hirsute ;
segments more or less spreading, broadly deltoid.

The following specimens belong here: Wright, no. 1435;
Woodhouse, Zuni Mts., N. M., Aug., 185 i ; Wooton, no. 172.

Androsace diffusa.

Annual, acaulescent, more or less pubescent. Leaves basal ;
blades oblanceolate to spatulate or nearly linear, 1-4 cm. long,
obtuse or acute, sharply serrate above the middle or sparingly
toothed near apex only, sessile or with short winged petioles :
scapes erect and spreading, often diffusely branched at base, 5-10
cm. long or shorter : bracts lanceolate : pedicels filiform, very vari-
able in length, often 1-8 cm. long in the same cluster : calyx cam
panulate to turbinate-campanulate, 3-3.5 mm. high; segments
triangular, acute, ciliate, about y^ as long as the 5 -ridged tube ;
corolla white or pink, included, sometimes equalling the tips of the
calyx-segments, 3-3.5 mm. broad; segments oblong, obtuse or
retuse at apex, about as long as the tube : filaments shorter than
the anthers : capsules subglobose, about 3 mm. in diameter.

In rocky soil, western Arctic America to the Dakotas, New
Mexico and Arizona. Spring and summer.

For some inexplicable reason the species here described as new
has always been associated with Androsace septentrionalis with
which it has not even a habital resemblance. Androsace septen-
trionalis is a plant with strict, conspicuously elongated scapes
which are surmounted by umbel-like clusters of pedicels of nearly
equal length, whereas Androsace diffusa, has comparatively short,
more or less diffusely spreading scapes, while the pedicels of the
clusters are exceedingly variable in length. A more tangible

319 Small : North American Plants

character exists iri the corolla. In Androsacc scptoitrionalis this
organ conspicuously surpasses its calyx while in the newly de-
scribed species it is shorter than its calyx or barely equals it.

Androsace subumbellata (A. Nelson).

Androsace scptentrionalis subumbellata A. Nelson, Bull. Wyom.
Exp. St. 28: 149. 1896.

Annual, diminutive, sparingly pubescent. Leaves basal ; blades
thick, oblong, 2-8 mm. long, obtuse, entire, sessile : scapes 1-5
mm. long, or wanting : bracts ovate-lanceolate or lanceolate, pedi-
cels solitary or several together, 5- 10 mm. long: calyx nearly
glabrous, turbinate-campanulate, 2.5 mm. high ; segments triangu-
lar, acute, slightly shorter than the 5 -ridged tube: corolla white
or pink, 2.5-3 "^n^- broad, surpassing the calyx ; segments oblong,
obtuse, or retuse at the apex, shorter than the tube ; filaments
much shorter than the anthers : capsule globose-pyriform, about
2 mm. thick.

On hillsides, near summit of Union Peak, Wyoming. Summer.

In order to treat this genus consistently, we should recognize
the above as a species. On the one hand Androsace siibiunbellata
is related to Androsace diffusa : this species it resembles in habit
and foliage, but it is more diminutive in all its parts. On the other
hand it is related to Androsacc septentrionalis in having the corolla
exserted beyond the calyx.

Primula serra.

Perennial, glabrous or nearly so, deep green. Leaves basal,
5-10 cm. long; blades narrowly oblong or spatulate, much
longer than the broadly winged petioles, rather regularly dentate,
acute or apiculate : scapes erect. 1-2 dm. tall, solitary or several
together : bracts scarious, ovate-lanceolate, acuminate, often mi-
nutely pubescent : pedicels 1-3 cm. long, glabrous in age : calices
6-7 mm. long ; tube campanulate ; segments lanceolate, granular-
ciliate, acuminate, as long as the tube or shorter : corollas lilac-
purple ; tube as long as the calices or somewhat longer ; seg-
ments suborbicular or obovate-orbicular, notched at apex, 7 8
mm. long, destitute of apiculations.

Primula serra resembles Primula Rusbyi in habit, but both the
foliage and the inflorescence furnish characters for distinguishing
the two species. In the case of the species just described we find

Small : North American Plants 320

more coarsely toothed leaf-blades and shorter petioles ; but more
prominent characters exist in the inflorescence : The calices are
twice as large as those of Privmla Rusbyi, the corolla-tubes are
comparatively stout and they never twice exceed the length of the
calices as do the very slender tubes of Privmla Rtisbyi. In addi-
tion, the lobes of the corolla-segments are destitute of the minute
but characteristic apiculations found in the related species.

The original specimens were collected by Mr. Pringle on damp
ledges, Santa Rita Mountains, Arizona, at an altitude of 8000
feet, on July 25, 1884.

Primula tenuis.

Perennial, glabrous, bright green. Leaves basal, 0.5-1.5 cm.
long ; blades oblong, oval or suborbicular, undulate or toothed,
obtuse or acutish, much shorter than the slender petioles : scapes
erect, wire-like, 5-10 cm. tall, solitary: bracts linear-subulate, 1-5
mm. long: calices 3-5 mm. high, tubes turbinate, segments narrowly
lanceolate to narrowly linear, longer than the tube, acute : cor-
ollas pink, 4-5 mm. broad ; tubes surpas.sing the calices, 4 mm
long, notched at the apex.

In moist places, Pastolic, Alaska.

A delicate species related to Primula borealis, but much more
slender ; easily distinguished by its flimsy leaves, shorter pedicels
turbinate calyx-tubes and narrow calyx-segments. The corolla
tube is further exserted than in Privmla borealis and the more
delicate lobes less deeply notched.

The original specimens were collected by W. H. Dall, on
June 25, 1871, or 1872.

Notes and Descriotions of North American Plants.— II

By John K. Smai.i,
I. NOTEWORTHY SPECIES

Habenaria Garreri Porter, Bot. Gaz. 5: 135. 1880
The original and second known stations for this interesting
orchid are both near Manatee, Florida. A second locality can
now be placed on record ; this is Orange County, Florida, where
Mr. F. L. Lewton discov^ered the species at several stations in
the summer of 1894. His specimens are essentially the same as
the type.

Habenaria macroceratitis Willd. Sp. PI. 4 : 44. 1805
This remarkable tropical Habcfiaria has been found native in
Florida, by Mr. Lewton. It is not rare in Sumter County, where
he first met with it in 1 894.

Thermopsis mollis (Michx.) AI. A. Curtis, Mem. Am. Acad. II.

3: 47. //. g. 1848
Heretofore this comparatively rare species has been reported
as growing in the mountains of Virginia and North Carolina.
But its range is wider than this; in May, 1869, Mr. Canby col-
lected it on Lookout Mountain, Tennessee, and on May 21, 1890,
Professor Scribner rediscovered it at the same locality.

Pluchea imbricata (Kearney) Nash, Bull. Torr. Club, 23: 108.

1896
Excellent specimens of this PhicJiea were collected in swamps
about Forest City, Orange County, Florida, by Mr. F. L. Lewton
in July, 1893. 'T^^ specimens of this collection agree almost
perfectly with the type.

Hieracium Scribneri Small, Bull. Torr. Club, 21 : 20. 1894
Professor Ruth has sent me almost typical specimens of this
rare member of Hieracium from near Knoxville, Tennessee, where
he collected the plant in 1897.

(275)

27G Small : North American Plants

Senecio MiLLiFOLiUM T. & G. Fl. N. A. 2 : 444. 1843

In 1887 Mr. E. R. Memminger rediscovered this rare Senecio
in Henderson County, North Carolina, where it was collected
■ many years ago by Buckley. In 1895 Mr. A. M. Huger sent
me specimens from Macon and Jackson counties, North Carolina,
where he found it growing plentifully on sloping cliffs at altitudes
ranging from i 100-1400 meters.

11. HITHERTO UNDESCRIBED SPECIES

Allium arenicola

Bulbs nearly i cm. long, with fibrous outer coats. Leaves
basal ; blades very narrowly linear, becoming almost filiform,
about as long as the scape or shorter : scapes erect, sometimes
several together, T-3 dm. tall, more or less curved : umbels erect,
iO-3o-flo\vered : pedicels 5-1 0 mm. long, slender : perianth deep
pink ; segments linear to narrowly linear-lanceolate, about 4 mm.
long, very delicate : filaments dilated below : capsules not crested.

In sandy soil, Mississippi. Spring.

This species has been confused with Al/iiim inutabile Michx.
for nearly three quarters of a century. It is much more slender
in habit and smaller in all its parts. The type specimens were
collected by Martha B. Flint at Brookhaven, Mississippi, April i ,
1888.

Ranunculus cuneiformis

Foliage hirsute below the inflorescence. Roots thickened,
clustered : stems usually several together, 2-3 dm. tall, erect or
ascending, rather slender : leaves mainly basal ; blades, at least
some of them, twice-divided into cuneate rather obtuse segments,
5-10 cm. long, about as long as the petioles ; upper stem leaves
with blades 3 -parted ; segments narrow, often incised : flowers
yellow, about i . 5 cm. broad, on strigillo.se peduncles : heads of
fruit subglobose or ovoid-globose, about i cm. long : receptacle
barely elongated : achenes 4 mm. long, conspicuously winged and
with a triangular beak.

On prairies, near Kerrville, Texas. Spring. Heller, PI. S.
Tex. no. 1688. It differs from its relative as shown below :

Small: North American Plants 277

Ranunculus cuxeiformis Ranunculus macranthus

Blades of lower leaves twice- Blades of lower leaves once-
divided : corollas 1.5 cm. broad : divided : corollas 3-5 cm. broad :
heads of achenes sub<^lobose or heads of achenes oblong to
ovoid-globose, receptacle barely cylindric : receptacle elongated :
elongated : achenes conspicu- achenes narrowly margined,
ously winged, with triangular with subulate slightly curved
beaks. beaks.

Ranunculus Mississippiensis

Perennial, stolonifcrous, fleshy. Stems stout, about 2 dm.
tall, more or less branched : leaves various ; basal or those on the
lower part of the stem with ovate or ovate-lanceolate sinuate-
dentate blades 1.5-4 cm. long, and elongated petioles, upper leaves
with oblong or linear remotely-toothed blades 3-8 cm. long :
flowers few : sepals oblong to suborbicular, sparingly pubescent :
corollas about 1.5 cm. broad; petals about 9, nearly oblong, deep
yellow and lustrous within.

In low grounds, Arkansas and Mississippi. Spring.

Arkansas: Varner, Lincoln Co., April 28, 1898; BiisJi, no.
12.

Mississippi: "Alluvions." 1840; Peck.

Related to Ranunculus oblongifolius, but more robust, with
truncate or cordate blades terminating the elongated petioles of the
lower or basal leaves and much larger corollas consisting of about
nine petals.

Thalictrum mirabile

Perennial, slender, glabrous, bright green. Stems erect, 1-3
dm. tall, wiry, dichotomously branched above : leaves various,
basal usually ternately compound, with petioles about 2 cm. .long ;
upper leaves gradually more simple and shorter petioled : leaflets
suborbicular or orbicular-reniform, 2-3 cm. broad, very thin, deli-
cately nerved, glaucescent beneath, broadly crenate or shallowly
crenate-lobed, truncate or subcordate at the base, longer than the
petiolules : peduncles hair-like : flowers white : sepals spatulate or
rhombic-spatulate, fully 1.5 mm. long : filaments fully 2 mm. long,
club-shaped by an abrupt thickening about the middle : fruit
spreading at right-angles to the peduncle ; body plump, about 2
mm. long, acute, not depressed along the upper side, as long as
the filiform stalk or shorter.

Resembles TJialictruni clavatum but more delicate and smaller
throughout, and with very short-petioled basal leaves. The fruit

278 Small : North American Plants

is only about one-half the size of that of T. clavatim and has a
plump barely ribbed body not at all depressed along the upper side.
The original specimens were collected by Prof. F. S. Earle
under sandstone bluffs on Little Mountain near Moulton, Alabama,
June 25, 1899, no. 2212.

Phyllanthus Avicularia

Perennial, bright green. Stems branched at the base and
throughout, 3-6 dm. long, puberulent, striate in age : leaves nu-
merous, ascending : blades oblong, or slightly broadest above the
middle, 8-18 mm. long, blunt or barely pointed, slightly paler
beneath than above, rounded or truncate at the base : petioles i mm.
long, or shorter : calices short-pedicelled ; staminate delicate, barely
2 mm. broad, sepals orbicular-obovate or suborbicular : pistillate
firmer, fully 2 mm. broad or barely 3 mm. broad at maturity ; se-
pals oblong or oval, scarious-margined, persistent : capsules sphe-
roidal, 3 mm. broad.

In dry soil, along the Brazos River, Texas. Type from Co-
lumbia, Texas, collected by B. F. Bush, October 26, 1899, no. 263.

Related to Phyllanthus polygonoides, but much more robust in
all its parts. The leaves, too, are of an oblong type. The cap-
sules conspicuously surpass the mature pistillate calyx, whereas
those of P. polygonoides are at least equaled by the mature sepals.

OEnothera nyctaginiifolia

Apparently annual or biennial, sparingly pubescent. Stems
branched at the base, branches spreading or decumbent, 2—5 dm.
long, more or less branched : leaves rather few ; blades lanceolate
to ovate-lanceolate, 2-5.5 cm. long, acute or slightly acuminate,
often somewhat crisped and twisted, undulate, ciliate, cuneate or
truncate at the base ; petioles 2-6 mm. long, pale, margined :
flowers axillary : hypanthium bristly and with very slender hairs,
especially about the ovary ; tubular portion about as long as the
ovary: sepals linear-lanceolate, fully 1.5 cm. long, thin and deli-
cate: capsules 4-5 cm. long, club-shaped by the sterile basal por-
tion which is slightly shorter than the fertile portion, about 4
mm. thick : seeds 1.5 mm. long, reticulated.

In dry soil, Flagstaff, Arizona, September 5, 1894,/. W. Tonmey.

More closely related to OEnothera laciniata than any other
species. It differs in the larger flowers and the club-shaped cap-
sules, besides the conspicuous character of the leaves. These
members are very suggestive of the leaves of Nyctaginca or the
broad-leaved species of Allionia.

Small : North Amkrican Plants 279

Phlox Brittonii

Perennial, deep green. Stems copiously branched ; branched
matted, forming wide tufts, glandular-pilose : leaves numerous,
small ones often clustered in the axils of the larger; blades subu-
late or narrowly linear-subulate, 5-10 mm. long, ciliate, especially
near the base : calices 5-6 mm. long, glandular-pubescent like the
branches ; segments subulate, about as long as the tube : corolla
white : tube curved, about i cm. long ; limb 12-13 mm. wide ;
segments cuneate, with 2 pale magenta spots near the base, cleft
by a V-shapcd sinus about 3 mm. deep, usually with a minute
tooth in each sinus, tips acute or acutish.

On dry mountain slopes, Virginia and West Virginia to North
Carolina. Spring and Summer.

A relative of Phlox siibu/ata but more delicate in all its parts.
The contrasting characters may be shown as follows :

Phlox Brittonii Phlox subulata

Stems or branches glandular- Stems or branches not glan-

pilose : leaf blades mostly 5-10 dular : leaf-blades mostly lO-

mm. long: calices 5-6 mm. long: 15 mm. long: calices 8-9 mm.

limb of corolla less than 14 long: limb of corolla over 15

mm. broad ; lobes usually with mm. broad ; lobes usually with

a minute tooth in each sinus. toothless sinuses.

The specimens upon which the species is based were collected
by Dr. N. L. Britton, at White Sulphur Springs, West Virginia,
May, 1898. Dr. Britton then introduced the species in the herba-
ceous grounds of the New York Botanical Garden where the plants
have become thoroughly established.

Vernonia interior

Perennial, finely and usually closely pubescent. Stems erect
or ascending, 1-2 meters tall, simple below the inflorescence :
leaves numerous ; blades elliptic to elliptic-lanceolate, 6-20 cm.
long, acuminate, sharply and rather finely serrate, sessile or nearly
so : heads numerous, rather crowded : involucres campanulate,
6-y mm. high, 4-5 mm. broad : bracts pubescent, sometimes
hoar}% acute or with short keel-like acuminations, the tips erect or
slightly spreading : achenes pubescent : pappus purple.

On plains or prairies, Missouri and Kansas south to Texas.
Spring to fall.

280 Small : North American Plants

The species just described has heretofore been confused with
Vcrnonia Baldn'inii and V. Druuimondii. It is readily separable
from its nearest relative, Vernoiiia Baldivinii by the smaller in-
volucres and their bracts which have erect or barely spreading
tips. The following cited specimens belong here :

Missouri: Jackson County, BusJi, no. 233A ; McDonald
County, Biisli, no. 232.

Nebraska: Lincoln, Webber, September, 1888.

Texas: Kerrville, Heller, PI. S. Tex. no. 1927.

This species has been raised from seed in the nurseries of the
New York Botanical Garden and is now established in the herba-
ceous grounds.

Vernonia maxima

Foliage glabrous or sparingly pubescent. Stems erect, I-3
meters tall, branching above : leaves rather numerous : blades
narrowly elliptic to lanceolate or linear-lanceolate, 1-3 dm. long,
acuminate, sharply serrate, narrowed into short petioles or the
upper ones nearly sessile : corymbs 1-4 dm. broad : peduncles
angled, barely enlarged upward : involucres hemispheric, 4 mm.
to nearly 5 mm. high, rounded at the base : bracts ovate to ob-
long, acute to mucronate, ciliate, appressed : achenes 3 mm. long,
upwardly barbed on the ribs : pappus light or deep purple.

In low ground, Ohio to Missouri, south to Alabama and
Louisiana. Summer and fall.

For many years Vernonia gigantea or V. altissinia has been an
aggregate. The campestrian plant that has been known under both
of those names is very distinct from the Carolinian and Floridian
plant to which both the above cited names were originally applied.

The campestrian plant may easily be separated from the south-
eastern species by the lower involucres with rounded bases and
their proportionately broader appressed and compactly arranged
bracts. The involucres of the related species are narrowed at the
base and have narrower loosely spreading bracts. The following
cited specimens belong here :

Missouri : Jackson County, Bnsh, no. 230.

Ohio: no locality, Riddell, 1834. Scioto, Merriain, Septem-
ber 28, 1 89 1.

West Virginia : Monongalia County, MillspangJi, no. 677.

Kentucky: no locality. Short, 1842. Harlan County, A'm;'-
ney, no. 188.

Small : North American Plants 281

Tennessee: Knoxville, Ruth, September, 1894.
Mississippi : Agricultural College, Pollard, no. 1267.

Lacinaria Halei

Perennial, glabrous or nearly so. Stems erect, 6-9 dm. tall,
simple or sparingly branched : leaves various ; lower with linear
blades 1-2 dm. long, upper narrowly linear and much shorter, not
ciliate near the base : heads short-peduncled or nearly sessile, not
densely crowded : involucres becoming narrowly turbinate, 7-9
mm. high : bracts lanceolate to oblong-lanceolate, acuminate, cil-
iolate, rigid : pappus plumose : achenes closely pubescent.

On prairies, Louisiana. Summer.

This species has heretofore been included in Lacinaria acidota,
with which it has little or nothing in common, and it may be sep-
arated by its fewer leaves and much smaller heads which are dis-
posed in elongated interrupted spikes. The bracts of the involucre
are much shorter than those of L. acidota and have less elongated
tips.

The species is founded on Hale's no. 334.

Lacinaria platylepis

Perennial, bright green. Stems erect, 8—9 dm. tall, simple,
glabrate below, pubescent with white hairs above : leaves not very
numerous, narrowly linear, 2-10 cm. long, or longer at the base
of the stem, glabrous or nearly so : heads rather approximate,
sessile, surpassing the subtending bracts : involucres cylindric-
campanulate, 7—9 mm. long ; outer bracts often ovate, acute, inner
larger and broader, broadest above the middle, rounded at the
apex, ciliolate : pappus not plumose, pale.

In sandy soil, Louisiana.

Plants belonging here have been referred to Lacinaria acidota.
although none of the several characters warrant such a disposition,
The fewer and shorter leaves, the elongated more or less inter-
rupted spikes and smaller heads and involucres with their broad
rounded inner bracts, are some of the characters that separate
Lacinaria platylepis from L. acidota. The pappus too is not plu-
mose.

The original specimens were collected in Louisiana by Dr.
Hale.

COXTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 7

VEGETATIVE REPRODUCTION AND MULTI-
PLICATION IN ERYTHRONIUM.

BY FREDERICK H. BLODGETT

NEW YOEK
1900

[Reprinted from the Bulletin of the Torkey Botanical Club. 27 : 305-315. 25 June, 1900.]

Vegetative Reproduction and Multiplication in Erythronium*

By Frederick H. ]]lodgett
(With Platks 16-18)

John Burroughs, in Riverby, speaking of the yellow ErytJiro-
niiuii as he found it in grass-covered meadows, calls attention to the
brittle white threads which often appear above the turf. These he
found were connected with the immature bulbs, from which they
penetrate the soil in various directions.

The nature and purpose of these threads were studied in 1893
and 1894.1 They are smooth, scaleless offshoots or subterranean
runners, heavily charged with starch. The tip encloses a bud
which will become a bulb upon the death of the parent bulb. In
this species, and in E. albidum the runners arise from the base of
the parent bulb, differing in this respect from E. propidlaiis Gray,
which produces the offshoots from the side of the stem above the

bulb.t

The bulbs formed at the distant end of the runners repeat the
process indefinitely, producing annual crops of runners and runner-
bulbs until the conditions are met which result in flowering plants.
When the necessary vigor, depth and size are reached which cause
the development of a flower bud in addition to the leaf bud within
the bulb, the annual runners cease to be produced ; but when the
flowering bulb is removed from those conditions, runners are again
produced. The runners are first produced from the bulbs which
are formed during the growth of the seedling ; the last crop gives
rise to flowering bulbs four years or more after the seed has
ripened. After the runners cease to be produced, annual bulbs
are formed within the mature bulb, resembling the runners in their
structure save that there is no lengthening between the terminal
bud and the point of origin.

* Read in abstract before the Society for Plant Morphology and Physiology, New
Haven, Dec. 27, 1899.

t Bot. Gaz. ig : 61. 1894; and 20 : 172. 1895; with illustrations.

J Am. Nat., July, 187 1. A new Species of Erythroniiini. Dr. Gray mentions also
the function of the offshoots in the species Dens-canis, Americamim ?in6. propullans.

( 305 )

306 Blodgett : Vegetative Reproduction

The mature bulbs are usually in good flower about the middle
of April, and the seeds ripen in June. They must be looked for
among the leaves upon the surface of the ground, as the entire
plant becomes withered and prostrate soon after the fall of the
flowers. The ripe seeds are about 6 mm. long, half as wide, and
in shape resemble, distantly, a segment of an orange, on account
of a prominent raphe and spur along one side (Fig. i). In tex-
ture the seed is firm and hard ; in color, brown darkening with age
and exposure. The surface is strongly convex, and smooth,
though hardly shining. The raphe and spur are absent in seeds
which have lain in the soil over winter. In Erytlironiuni Dens-canis
the raphe is less developed, the spur is present as a slender curved
hook or beak from the apex of the seed.*

The seeds remain dormant for nine months — from June to
April, when they germinate (Fig. 4), reaching their best develop-
ment as seedlings (Fig. 7), about the time that the flowers are in
their prime. The young plant elongates in opposite directions :
the upper end, the tip of the cotyledon, contains a gland (Fig. 6,^'-)
by which the food substance of the seed is absorbed. The lower
end is protected by a root cap during the first stage of its growth.
While the tip of the cotyledon advances the length of the seed, the
opposite end penetrates the soil for an inch or more. About the
time that the food is absorbed from the seed and the cotyledon
frees itself from the empty testa, the descending part changes also-
From one side of this portion, close to the tip (/', Fig. 5), the rad-
icle is produced, and penetrates farther into the soil. It is plenti-
fully supplied at its base with root hairs (Fig. 9).

The cotyledon is supplied, for the greater portion of its length,
with stomata, and functionally it is a cylindrical leaf. Starch is
scattered through it, being in all portions of the tissue at a (Fig.
23), mostly epidermal in the guard cells of the stomata at h, usu-
ally near the fibrovascular bundles at c, and general again at d.
Close to the tip id') there is a local deposit of starch, as if pro-
vided for the growing cells just within the root cap {S, Fig. 5).
The part from which the radicle springs enlarges, becomes charged
with starch, and forms a bulb (Fig. 9), which is usually forced

*Irmisch : Beitrage zur verg. Morphologie der Pflanzen. Abhand. d. Natur. Gesell.
zu Halle, 17^: 184-195. 1863.

AND Multiplication in Erythronium 307

deeper into the soil at the end of a short vertical runner, but
may develop without such elongation. Within this bulb the
plumule is formed (Figs. lo, ii) and with its formation the
first step in the v^egetative life of the plant is completed. The sec-
ond step appears the next spring with the first leaf — the plumule
leaf, and ends when the first crop of runner bulbs is formed in
May.* From the plumule-bulb only one or two runners are gen-
erally produced, but from the larger bulbs three is the more com-
mon number (Fig. 12), thus increasing three-fold the number of
immature bulbs at each successive crop of runners.

A flowering bulb cannot be produced from seed in less than
lour years. In the first year the parent bulb would bloom, and
ripen its crop of seeds, from six or seven to twenty or more. In
the spring of the second year some of these seeds would germi-
nate and form plumule-bulbs. From each of the plumule-bulbs
there would appear in the third year a single leaf, and the first
runners would be produced at the distant ends of which runner-
bulbs would be formed. Some one of these runner-bulbs might
be formed under the conditions necessary to produce a flowering
bulb, but this is very unlikely ; so that one or more years would
elapse before a blossom would be formed, thus making a cycle of
five or more years. This cycle is shown at Fig. 14, with the
forms assumed at each step. At Fig. 26 the multiplication which
takes place during the same cycle is shown diagrammatically : one
seed ; one seedling and plumule-bulb ; one runner bulb ; three run-
ners with their bulbs ; and from each of these three there are pro-
duced three more the fifth year, nine in all. Some of the nine
will probably produce a flower ; those which do not will continue
to produce runners in most cases, although a bulb is occasionally
found which is apparently recuperating, for the depth is that of the
mature bulbs, but only one leaf is produced, and no runners are
present.

The following table illustrates the number of plants of different
ages during each of five years, supposing that five seeds from each
fruit ripen and survive the cycle. Each step is one line lower

*No bulbs of plumule size were found this spring (1900) with runners. The stiff
clay soil in which most of them were found may have some influence upon the absence
of runners.

308 Blodgett : Vegetative Reproduction

than the next preceding one, thus the five " seeds " of 1899, become
" 5 plumule-bulbs " in 1900, giving a total of 75 plants in 1903.

1899 1900 1901 1902 1903

5 seeds 5 seeds 5 seeds 5 seeds 5 seeds

5 plumule-bulbs 5 plumule-bulbs 5 plumule-bulbs 5 plumule-bulbs
5 yearlings 5 yearlings 5 yearlings

15 2 years old 15 2 years old

45 flowers

Total 75 plants

There is very little chance that there will be forty-five blossoms
at the end of five years, but as some of the bulbs will probably
flower then it is convenient to call the entire number by one name,
when they are of the same crop or age.

The seed resembles that of Iris in structure.* The walls of
the cells are thickened ; at certain points the walls of adjacent
cells retain their normal thin nature, so that a thin membrane
only separates the cell cavities. The cells are arranged with
their longer axes nearly perpendicular to the surface of the seed,
and in this way form lines which extend to the central portion of
the endosperm. Through the center of the seed a mass of dense
reserve cellulose extends (Fig. 19) with a few cell cavities scattered
through it. In the apex of the seed, and at the beginning of this
mass of dense material lies the embryo (Fig. 20). At germination
the embryo elongates, pushing the tip of the cotyledon along the
center of the seed, absorbing the dense cellulose as it advances.
From the cavity thus formed in the center, the solvent action ex-
tends toward the surface of the seed, following the lines of the
cells, and producing a honey-comb effect (Fig. 8). This honey-
combing is the more easily produced on account of the cell
arrangement which in a longitudinal section appears as in Fig. \Z,a;
in a horizontal one, as at b. The thin wall separating the depres-
sions in the thickened walls of adjacent cells readily dissolves under
the action of the absorbing gland of the cotyledon, thus opening
a passage for the movement of food materials from cell to cell, as
the solvent action extends. The dissolved material moves along
the channels so made toward the tip of the cot}'ledon. The ab-
sorbing organ (Fig. 6, g) is quite similar in structure, and probably

* Haberlandt, Pflanzen Anatomie, 298.

AND Multiplication in Erythronium 309

in action also, to the glands in the hairs of Drosera.* One or two
layers of cells cover the end of the fibro-vascular bundle which is
slightly larger than in the body of the cotyledon. No starch is
present in the seed, except in the raphe and spur, where a few
grains are scattered through the thin-walled cells (Fig. i8, c).
During the growth of the seedling starch is found just within the
glandular tip of the cotyledon, and in other portions as already
stated.

When the food material has been exhausted the cotyledon
" elbows " its way to the surface and functions as a leaf. At this
stage it resembles an onion seedling, but unlike the onion no fur-
ther development of foliage occurs until a year later. The plu-
mule-bulb is now forming and is complete, as a bulb, about the
time the seedlings wither early in May. The plumule is formed
at the base of the swelling at the origin of the radicle (/, Fig. lo).
At first a mere line of division separates the plumule cells from
the other cells of the young bulb. Later the plumule appears as
a protuberance at the bottom of a small cavity in the base of the
bulb [p, Fig. ii). After the cotyledon withers the plumule con-
tinues to develop, until a small leaf is formed by the first of No-
vember. The plumule-bulb becomes heavily charged with starch
early in its development, and thereafter the subterranean portions,
except the shoots inside the bulbs are starch bearing.

Runners have their origin at the base of the stem, as a bud in
the axil of the inner bulb-scale. From this point (Figs. 13, 15)
they push out irregular distances, and at the completion of their
growth form bulbs from their terminal buds. Mature bulbs are
annually renewed from similar buds (Fig. 17), which develop in-
side of the parent, beginning as buds in December ; the new bulb
being full size just after the blossoms fall, in late April. Fig. 30
shows the new bulb at the time that the parent is in prime flower.

The runner and annual bulbs begin to form the buds for the
next spring's leaves and flowers, in May. The first evidence is a
discolored line or streak extending upward from close to the bot-
tom of the bulb ; soon there appears a cavity at the lower end of
this streak, in which a short sprout is visible in June or July (Fig.
16). This sprout continues to develop until in November and

*Kerner, Nat. Hist, of Plants, 2: 145. / 126.

310 Blodgett : Vegetative Reproduction

December the floral parts are all present and well developed in-
cluding the ovules and the pollen mother-cells, and the perianth
is often tinged with yellow. As the spring advances the bulbs
push their sprouts further and further toward the surface, and
when the frost is fairly out of the ground the short distance re-
maining is quickly passed, and the flowers open while there may
still be snow on shaded hillsides. The sprout penetrates the soil
in the manner of an awl, piercing leaves and decayed wood which
may lie in its path,* rather than growing around, or lifting them.
For this awl-like work the sprout is protected by a hard tip, which
forms the apex of the leaf when the latter expands.

In Fig. 22 the successive steps are shown, by which the
flower becomes freed from the enclosing leaves, and assumes its
nodding position on the pedicel. At the left {a) the sprout has
fully penetrated the soil and the leaves have begun to relax their
tight clasp upon the bud. The outer and larger leaf expands
gradually, while the inner one remains close to the bud as shown
at d. Next the two leaves bend downward toward their final posi-
tion, the bud still held by the margins of the inner leaf, c. When
the leaves have reached nearly their normal position, the bud is
released d, and it then stands erect between the leaves, as at e.
In a short time the stem begins to bend near the bud, as at/, and
as the bud begins to open it has assumed the position shown at g:

The last two positions alternate while the plant remains in
flower, position/ being assumed on bright days, when the open
flower, with strongly recurved petals, is turned upward, returning
to the nodding position at night and on less warm or less sunny
days. With the fall of the flower the enlarging fruit assumes an
erect position upon the stem, which is retained until the whole
plant is withered and lies prostrate on the ground ; the ripe seeds
remain quite close to the open capsule.

Erytlivoninvi Americamun in its mature development normally
bears two unequal leaves and a single flower. During the past
two or three seasons a number of unusual cases have been ob-
served, in one of which two flowers were borne on an otherwise
normal plant. More frequently one leaf only was present in a
plant otherwise normal. Over a dozen plants showing the latter

*Bessey, C. E., A vegetable Awl, Plant World, I : 132.

AND Multiplication in Ervthronium 311

form of variation were found in an hours' collection in New Jersey.
A plant collected at Washington, D. C, had one leaf (Fig. 28)
eight stamens and seven petals ; the flower is shown at Fig. 27.
Plants having three leaves, with one flower ; and four leaves, with
two flowers ; have been found at long intervals. Fig. 3 1 shows
also three rosettes formed by the root fibers, which indicates the
minimum age of the bulb at the depth at which it was found, that
is, three years.

Bulbs and runners also have unusual developments, such as
surface buds or stimulation growths. The latter seems to have
started after a rain or other stimulation, and bursting through the
partly formed runner bulb, has grown further into the soil. Fig.
31 shows one in which the nearly formed bulb is burst in two
directions, sending a continuation downward, and attempting to
produce leaves through the upper opening. These rudimentary
leaves showed a trace of yellowish green when first gathered. In
bulbs which have been injured, so that the sprout from the center
of the bulb fails to develop, a surface bud may be formed. Two
such bulbs were recently observed and are illustrated at Fig. 24.

The starch which is very abundant in the bulbs is very finely
banded, the fine lines being accompanied at intervals by coarser
ones. Fig. 21 shows some of the more common shapes assumed
by the starch. Occasional grains are found which are greatly
elongated. Often there is a crack or fissure running through the
nucleus of the grain but this is not constant, especially in small or
medium sized grains.

So far as known, a satisfactory explanation of the common
name of Erytlu'onhnn has not been given. Many plants have for
their common names the translation of their Latin designation, and
the common name will remain through many changes in nomen-
clature of the scientific descriptions. People seeing in a new coun-
try a plant closely resembling one familiar near their homes,
usually give the new plant the name of the one already known.
Hence when Erythronhnn Aincricamnn was seen, so closely re-
sembling the European species, it received the common name of
hat better known plant, dog's-tooth violet.

In the libraiy of the U. S. Department of Agriculture at
Washington, there is a copy of Leonard Fuchs' herbal : De his-

312 Blodgett : Vegetative Reproduction

toria stirpium insignes * * * Basileae, 1542. When examining
this and several other volumes in December, 1897, a description,
and excellent illustration were found of the European Erythroniwn,
named Viola Dens-canis. The Viola part of the name is probably-
due to coincidence in time and place of flowering : Dens-canis re-
fers either to the shape of the bulbs, or to the dentations at the
base of the petals. In another part of the same volume a number
of violets are figured bearing the generic name Viola as now.

During- the examination of old botanical works in the search
for the above reference the following names were found, applied to
what has been known since Linnaeus as Erythroniiun Dcns-canis.

Pliny : (Historia Mundi ; Froben, ed. Basillae 1554), lib. 26, ch.
X., p. 472, speaks of "another Satyrion (called Erythriacon), seeds
like vitcx but larger, smooth ; root hard, cortex red, enclosing
white within ; sweet juice ; occurring in mountains."

Dioscorides (Pedacii ; Saracenius, ed. 1598), lib. 3, ch. 144,

P- 232.

" Satyrion Erythroniiun or erythraiaim has seeds resembling
limim, but larger, firm, shining or dull. * * * The cortex of
the root rather delicate, and red ; the internal portion white, juice
sweet, pleasant to the mouth."

Dodoen's Erundeboeck, Antwerp, 1554, gives an illustration
of Satyrion erythroninm, which is probably meant to be one of the
Allinms as some of the herbalists consider that Allium nrsinuni is
referred to by Dioscorides' Satyrion erythroniiun.

Camerarius' Kreuterbuch, Frankfort, A. M., 1586, page 389.
In this herbal Hcrmodactybis and Pseiidohermodactyhts, are figured
side by side and represent Iris and Erythroniuni respectively. The
two names are often given by the old botanists as synonyms of
Dens-canis, or dog's-tooth, as they most frequently called it.

" Pseiidoherniodactylns is a beautiful plant, it brings two, or rarely
three leaves much like the lily. It is spotted with many brown
spots. Its flower grows upon a yellowish-brown stem, light pur-
ple with brown stamens (fasemen) and a white pistil (stiftlin), in the
center. The little leaves [petals] unfold when the sun shines
warm upon it, but when, however, they have fallen away, a three-
angled knob develops full of yellow seeds. The root is longish,
thicker below than above, which are often clustered together. * *

AND Multiplication in Erythronium 313

While it does not bloom it brings only one flower. It is called
Dentali by Apotheckers in Stiermarck."

In Lobelius' Stirpium, Antwerp, 1576, there is a good cut on
page 64 of Erjtlironium with the brief description : " Erythroniinn
with colored flowers and unequal leaves." Satyrion Dens canimis,
Hennodactylus and Pscndohcrinodactylus are given as sjaionyms
used in different regions. " * * * Flowers either purple-red or
white, or purple and white mixed."

Plunkenetius (Phytographia, 1692,3: 130).
''Dens caninns, flowers yellow, Virginianus." *
Salmon's Herbal, 17 10.

"Dogs tooth * * * the same as 'satyrion having red cortex
on the root,' in Dioscorides, lib. 3, ch. 144." Three color forms
are described having white, red and purple flowers respectively.
Roberto Morrison, of London, in his Plantarum Historia
Oxonienis, 171 5, page 343, says that the name " Dens canis is
from the shape of the oblong tuberous root imitating the figure of
a dog's tooth, the name is good." " It has a capsule of three dis-
tinct valves. * * * When the flower is red the leaves are red
spotted, when white the spots are milky."

Gronovius (Flora Virginica, 1739, 151, ist edition).
'' Erytlwoniuni, leaves oval or oblong, glabrous, dark -spotted,
flowers yellow." t (Cited in Linnaeus Spec. PI. 1753, as var. /-,
under E. Deus-canis).

Gronovius' Flore Virginia,' Lugdonum, 1762, 51 (2d edi-
tion), contains a description of our native species. " Erythroninm
with ovate oblong leaves, smooth, dark-spotted. Dens-caninus
agiiatalis, flower yellow, pendulous, leaves ovate-oblong, prostrate,
two upon a stalk, glabrous, dark -spotted. Blooming toward the
end of March."

The reproduction by offshoots or runners, is mentioned in
Miller's Gardener's Dictionary, London, 1754, Vol. I., " Dens-canis
— Dog's tooth. * * * These plants are propagated by seeds, as

* Virginia mis is italicized, as are authors cited on the same page, but here it may
be a typographical error for a reference to Virginia, since the European form has purple
or red flowers.

fThis is probably the first description of the American species; especially in-
teresting, as it has the modern genus name. See previous note.

314 Blodgett : Vegetative Reproduction

also by offsets from the old roots. * * * The offsets, which
these plants produce but sparingly, should be taken off at the
time that the old roots are transplanted, * * * toward the latter
end of May."

Linnaeus cites descriptions from a number of the early bota-
nists, giving them as synonyms of his E. Dcns-canis, or as varieties
of it. Thus Dodonaeus (Stirpium historiae pemptades, 1616, 203)
gives " Dens caninus' which Linnaeus regards as synonymous with
Evythroniuni Dcns-canis, as is also " Dcns-canis with broad and
rotund leaves," of Bauhinus (Pinax, Basiliae, 1671). But " Dens-
canis with narrow and elongated leaves," of the same author is
regarded as variety ^ under E. Dens-canis, and Gronovius's descrip-
tion given in a preceding paragraph, is ranked as variety y in Spe-
cies Plantarum, 1753.

Among names used or proposed for use as non-Latin designa-
tions of ErytJironiuin, Trout Lily and Fawn Lily, have been sug-
gested by John Burroughs in Riverby, on account of the spotted
leaves, and because of the alert position which the two leaves
often assume. Spring Lily has been proposed, on account of its
early flowering, and the lily-shaped blossom. But as a colloquial
name, " Roosters " is the least suggestive of the plant meant.
This name is used in central New York, among the children
especially, because of the custom of " fighting roosters " with the
nodding flowers or buds, as is done with violets in New Jersey.

Explanation of Plates

Plate 16

1. Surface of ripe seeds, June and March, X 3-

2. Longitudinal and cross sections of seed (June), X 3-

3. Cross sections of ovary, April and May, X 12 and X 5-

4. Successive steps in germination, yi l}4 and X 3-

5. Longitudinal section of tip of seedling. /, fibrovascular bundle, r, point of
origin of radicle, s, deposit of starch.

6. Tip of the cotyledon, showing the absorbing gland, X ^°-

7. Full grown seedlings and "elbows," X i-

8. Tip of cotyledon in partly absorbed seed, X ^■

9. Base of cotyledon showing the enlarged portion which becomes a bulb (plum-
ule-bulb), X 5-

10. II. Sections of young plumule bulb, plumule at/, X 5-
12. Runners, and the developing runner bulbs, 'X i}4-

13-15. Axillary buds, from which runners will develop. Slightly enlarged.
16. The first stages of the flower-and-leaves shoot (July l).

Bull. Torr. Bot. Club, 27.

Pl. 16.

ERYTHRONIUM AMERICANUM.

Bull. Torr. Box. Club, 27.

Pl. 17.

ERYTHRONIUM AMERICANUM.

Bull. Torr. Bot. Club. 27.

Pl. 18.

ERYTHRONIUM AMERICANUM,

AND Multiplication in Erythronium 315

Plate 17

17. Bud of annual bulb (May), X i-

18. Three sections of seed tissue, a, longitudinal, b, transverse of endosperm,
cells from raphe, containing scattered starch grains.

19. Longitudinal section of seed, X ^5-

20. Embryo in endosperm, X 5°-

21. Starch grains from bulb, X 225.

22. Successive steps in the liberation and orientation of the flower, X I-

23. SeedUng, showing the starch-bearing regions, X I-

Plate 18

24. Surface buds on bulbs after the sprouts have been destroyed, X ^^

25. Life cycle of five years, showing the successive fomis assumed.

26. Diagrammatic life cycle, showing multiplication of one seed in five years.

27. Abnormal flower, petals, 7; stamens, 8; leaf, I (Fig. 28).

28. Leaf of 27.

29. Abnormal bulb, twhmcd by surface bud, X ^■

30. Annual bulb turned aside to show the relation of old and new growth.

31. Mature bulb which has been three years in one place, rosettes of root fibers
indicate the age, X *•

32. Stimulation growth of runner, X ^-

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 8

TWO NEW SPECIES OF GRIMMIA
FROM MONTANA

BY R. S. WILLIAMS

NEW YORK
1900

[Reprinted from the Bulletin of the Toerey Botanical Club. 27 : 316-317. 25 June, 1900.]

Two new Species of Grimmia from Montana

By R. S. Williams
(With Plates 19, 20)

Grimmia (Eugrimmia) Brittoniae

Growing in dense hemispherical tufts up to 3.5 cm. high.
Stems slender, usually bearing long branches. Outer perichaetial
and upper stem leaves with blade i mm. in length, oblong, some-
what lanceolate pointed, concave, flat on the borders with nearly
smooth hair point up to 3 times the length of blade : lower leaves
a little smaller with hair point scarcely equaling blade, more con-
cave and somewhat recurved : inner perichaetial leaves minute,
triangular, with hair point 8 to 10 times the length of the blade :
upper cells irregular, roundish or quadratic, about .006 mm. in
diameter, gradually becoming elongated below, toward base 3 or 4
times longer than broad near costa and i ^ to 2 times longer than
broad near margin : cells but slightly sinuous walled when filled
with chlorophyl, later on the walls becoming distinctly sinuous
both above and below: sections of costa show two large cells on ven-
tral side with usually 5 or 6 surrounding cells, in single row, on
dorsal side : leaf lamina is of a single row of cells with sometimes
a doubling of one row in one or both margins : apparently dioi-
cous : fruit unknown.

Growing on shaded perpendicular walls, partly calcareous, in
rather dry places. Collected for several seasons in Bad Rock
Cafion, Flathead River, Mont.

This species is dedicated to Mrs. Elizabeth G. Britton, to whom
all students and lovers of our mosses will ever be indebted.

Grimmia (Eugrimmia ?) tenuicaulis

In compact tufts up to 6 cm. high. Stems very slender, often
thread-like, with few simple, mostly short branches : perichaetial
and upper stem leaves rather broadly ovate-lanceolate, concave,
revolute on borders, blade i ^ mm. long, with rough hair point
about 7< blade in length, the papillae of point spreading, often re-
curved : moistened leaves erect-spreading : upper cells irregular,
somewhat transversely or vertically elongated, mingled with
rounded cells .004—006 mm. in diameter : cells toward base
more or less elongated rectangular, those near margin from nearly

(316)

317 Two NEW Species of Grimmia from Montana

quadratic to twice longer than broad, towards costa becoming 2-
4 times longer than broad : cells apparently never distinctly sinu-
ous walled : sections of leaf show costa with two large cells on
ventral side and two rows of somewhat smaller cells on dorsal
side : lamina of leaf is of one thickness of cells, usually wnth one
or two rows doubled near costa, or sometimes somewhat distant
from costa, but not doubled in margin : occasionally the leaf is
hyaline nearly y^ down from apex, the hyaline cells always elon-
gated : evidently dioicous : sporophyte not seen.

Specimens of type collected near Neihart, Belt Mts., Mont.,
Sept. 21, 1 89 1, also obtained at Marsh Lake and Dawson on the
Yukon River.

Hxplanation of Plates

Plate 19. Grimmia Brittoniae

1. Plant about natural size.

2. Upper leafj X about 35 diam.

3. Lower stem leaf, X 35 diam.

4. Section of leaf, X ^75 diam.

5. Part of perichaetiuni.

6. Median cells, X 35° diam.

7. Basal cells.

8. Apex of leaf from upper side, X ^75 diam.

9. Inner perichaetial leaf.

Plate 20. Grinunia ienuicaulis

1. Plants about natural size.

2. Upper stem leaf, X about 35 diam.

3. Lower stem leaf, X about 35 diam.

4. Part of perichaetium.

5. Median cells, X 35° diam.

6. Basal cells, X 35° diam.

7. Apex of leaf from below, X ^75 diam.

8. Section of leaf, X ^75 diam.

9. Section of leaf lower down, X ^75 diam.

Bill. Tokr. Box. Clib, 27.

Pl. 19.

v^

8fc

WWm

GRIMMIA BRITTONIAE.

Ik LL. TOKR. BOT. Cl.lB, 27.

Pl. 20.

\(i

i.

mQm

CD^

gsmm

OOQo

■spss?

GRIMMIA TENUICAULIS.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— I

No. 9

STUDIES ON THE ROCKY MOUNTAIN

FLORA— II

BY P. A. RYDBERG

NEW YORK
1900

J

[Reprinted from the Bxjlletin of the Torbey Botanical Club, 27 : 528-53S. 26 Oct., 1900.

Studies on the Rocky Mountain Flora — II

Bv P. A. Rydukrg

THE ROCKY MOUNTAIN SPECIES OF MELANTHACEAE
There has been a great deal of misunderstanding regarding the
species belonging to the family Melanthaceae (perhaps more com-
monly regarded as a sub-family of Liliaceae). Especially is this
true with respect to the species growing in the Rocky Mountain
region. In order that some of the confusion may be cleared up
and a way may be opened to further study of the group my present
views are here offered.

The family is represented in the Rockies by at least five genera
and seventeen species. Of these I have found myself obliged to
describe one genus and seven species as new. The genera are as
follows :

TOFIELDIA Huds.
This is represented by three species, distinguished as follows :

.Stem glabrous, scapose ; seeds unappendaged. T. palustris.

Stem viscid-pubescent at least above ; seeds appendaged.

Bractlets broadly triangular, connate for two thirds of their length.

T. intennedia.

Bractlets lanceolate-triangular, connate half their length or less. T. occidentalis.

ToFiELDiA PALUSTRIS Huds. Fl. Aug. Ed. 2, I : 175

A subalpine species of wet places, growing in America from
Greenland to Alaska and from the Arctic coast south to Quebec
and British Columbia. It is also found in subarctic Europe. The
following specimen from the Rockies is in the New York Herbaria :

British Columbia: Rocky Mountains, 1890, JoJin Maconn.

Tofieldia intermedia sp. nov.

Tofieldia ghUinosa Hook. Fl. Bor. Am. 2 : 179, in part. 1838.
Wats. Bot Calif. 2 : 184. Not Pursh.

A slender perennial with more or less leafy stem, 1.5—3 ^'^•
high, viscid-pubescent above. Leaves 5-20 cm. long, 2-5 mm.
wide, obtuse or acute : raceme short, dense, 1—2 cm. long : bracts

(528)

529 Rydberg : Studies on Rocky Mountain Flora

broad, ovate : pedicels usually 3 together, very short, in fruit 1-4
mm. long: bractlets under the flowers 3, broadly triangular, con-
nate for about two thirds their length : flowers yellow : sepals
obovate, 4-5 mm. long : petals somewhat narrower and longer :
capsule ovoid, about 5 mm. long : beaks about i mm. long, spread-
ing : seeds appendaged.

This has been confused with the eastern T. ghitinosa Pers. which
has an elongated raceme, longer pedicels, and oblong subequal
sepals and petals, about 4 mm. long. All specimens from the
west referred to T. glutinosa may belong to T. intermedia. At
least all that I have seen do so. T. intemnedia grows in bogs to
an altitude of 2,700 m. from Saskatchewan to Alaska, British
Columbia and Wyoming. The following specimens belong to it :

Alaska: Sheh-Shooh Lake, 1895, M. W. Gorman, 76* (type) ;
Yes Bay, 1895, Thos. Hoivell, 1666 ; Khantaak Island, 1892, F.
Fiinston, ^j ; Sitka, Bongard.

British Columbia: Summit of Selkirk Mountains, 1890,
John Macoiin.

Washington : Totoish Mountains, 1897, O. D. Allen ^7-/.(?)

Montana: Flathead River, 1892, R. S. Williams, gij ;
Granite, 1892, Kelsey ; Upper Marias Pass, 1883, Canby, J28.

Wyoming: Cement Creek, 1897, F. Tweedy, jj6.

Tofieldia occidentalis Wats. Proc. Am. Acad. 14: 283. 1879
This species is distinguished from the preceding by the nar-
rower, less connate bractlets, longer pedicels, narrower sepals,
larger capsule, 8 mm. long, and longer ascending beaks. The
following specimen from the Rocky Mountain region is in the
New York Herbaria.

British Columbia : Avalanche Mountain, 1890, /. M. Macoun.

XEROPHYLLUM
This genus is represented by two species :

Petals and sepals 7-10 mm. long. X. tenax.

Petals and sepals 4-6 mm. long. X. Douglasii.

Xerophyllum tenax (Pursh) Nutt. Gen. Am. i : 235. 1818

Hclonias tenax Pursh Fl. Am. Sept. 243. 18 14.

X. Douglasii R)-db. Mem. N. Y. Bot. Gard. i : 92 ; not Wats.

Rydberg : Studies ox Rocky Mountain Flora 530

This has been greatly confused with the following. Watson
stated that A\ Doiiglasii was collected on the Hayden Surveys of
the Yellowstone Park, but does not cite any locality in the Rocky
Mountains for X. tcnax. Coulter in his Manual of the Rocky
Mountain Region included consequently a description of the for-
mer, but none of the latter. As this book is the one most used
by Rocky Mountain botanists, all specimens of XcropJiylbun from
that region have therefore been labeled X. Doitglasii. I made the
same mistake in preparing my Catalogue of the Flora of Montana.
All the specimens cited there belong to X. tcnax instead of X.
Doiiglasii. Besides the Montana specimens there cited, the fol-
lowing Rocky Mountain specimens belong to this species :

British Columbia: Toad Mountain, Kootenay Lake, 1890
/. M. Maco2in.

Idaho: Kootenay Co., 1888, J. H. Saunders; Wiessner's
Peak, 1892, Sandbcrg, MacDougal & Heller, j88.

Xerophyllum Douglasii Wats. Proc. Am. Acad. 14: 284.
1879.

I have not seen any Rocky Mountain specimen of this species,
but include it on the authority of Dr. Watson who referred to it the
specimens from the Hayden Collection metioned above.

Stenanthella gen. no v.

Erect bulbous glabrous herbs with few narrow leaves and race-
mose or paniculate perfect greenish, brownish or purplish flowers.
Petals and sepals each 3, subequal, withering-persistent, narrowly
lanceolate, acuminate, with reflexed tips, and at last involute, without
gland and distinct claw. Stamens 6, free, included ; anthers reni-
form, confluently i -celled. Ovary ovoid, superior; styles 3.
Capsule lance-ovoid, 3-beaked, septicidal to the base, wholly su-
perior. Seeds, oblong, winged.

Dr. Gray included the species belonging here in Stenantldum,
but the genus is better defined than many of the recognized gen-
era in the family. In Stenanthium, the flowers are polygamous,
open, not campanulate in outline, the base of the ovary is inferior,
the tip of the petals and sepals not reflexed and the general habit
different. The genus Stcnantliclla contains two known species,
one from the island Sachalin, S. Sachalinensis {Stenanihiiini Sach-
aliiiense F. Schmidt), and the following ;

631 RvDBERG : Studies on Rockv Mountain Flora

Stenanthella occidentalis (A. Gray) Rydb. n. n.

Stenantlmnn occidentale A. Gray, Proc. Am. Acad. 8 : 405.

The species ranges from Alberta and British Columbia to Ore-
gon and Idaho. The following specimens are from the Rockies :

Montana: Deer Lodge, 1892, il/m Emma Ware; Flathead
River, 1883, Can/?r, jj2 ; Big Blackfoot, Caiiby ; divide between
Hell Gate and Blackfoot, 1880, Watson ; Columbia Falls, 1894,
R. S. Williams.

Alberta and British Columbia: Rocky Mountains, 1858,
E. Boiirgcan ; Kicking Horse Lake, i^^y , Jolin Macoiin.

Idaho: Kootenay Co., 1890,/. B. Lcibcrg, ^^12.

VERATRUM L.

Flowers greenish, bractlets foliaceous, often equaling or exceeding the flowers.

V. viride.
Flowers white or yellowish white ; bractlets membranous, much shorter than the pedi-
cels and flowers. V. speciosum.

Veratrum viride Ait. Hort. Kew. 3: 422. 1789

V. lobeliajiiim /5 Eschsclioltziaiuim R. & S. S}'st. 7 : 1555- 1829.

V. Esclisclwltzii Gray, Ann. Lye. X. Y. 4 : 119. 1837.

If the western plant should be included in V. viride or not, is
doubtful. The eastern plant extends west to Minnesota and the
western is found first on the Pacific slope in Idaho and British Co-
lumbia. Both are very variable and no good character has been
found to separate the two. As a rule the western plant has shorter
stamens and bracts ; the former scarcely half as long as the petals
and sepals. The western range of V. viride ox V. Eschseholtsiamim,
whatever name it should bear, is from southern Alaska to Oregon
and Idaho. The following are the only specimens seen from the
Rockies.

Idaho : Packsaddle Peak, 1892, Sandberg, MacDotigal & Hel-
ler, 863.

Veratrum speciosum sp. nov.

Veratrum Californiciim Wats. Proc. Am. Acad. 14 : 277 ; in
part. 1879. Not Durand.

A stout, more or less pubescent, very leafy plant, 1-3 m. high.
Leaves, except the uppermost, broadly oval, 2-3 dm. long, 1-2

Rydberg : Studies ox Rocky Mountain Flora 532

dm. wide, obtuse or acute, glabrous above, finely soft-pubescent
beneath, sessile and sheathing : panicle many-flowered with ascend-
ing branches : bracts lanceolate, foliaceous : bractlets membranous,
yellowish, ovate, acuminate, shorter than the flowers and pedi-
cels : petals and sepals yellowish white, oval or broadly oblanceo-
late, mostly obtuse, 5-7-nerved, 8-10 mm. long, 4-5 mm. wide:
capsule oblong, about 3 cm. long and i 2 mm. in diameter : seeds
oblong, y—8 mm. long with a wide white wing-margin.

In looking over the specimens of V. Califoniicum in the Co-
lumbia Herbarium I found one specimen collected in Oregon on
the Wilkes' Expedition, which looked very unlike the rest, having
a narrower and denser panicle and narrower and more acutish
petals and sepals. I took it out and placed it in the cover used
for the specimens not named. I found in that cover a similar
specimen. On the label was given neither locality nor the col-
lector's name ; but on the sheet was pasted a paper with a tracing
of the basal leaves, the description o{ ]^. Calif orniciim E. Durand
and some remarks, among others the words "petiole 3-4 inches
long." The specimens are presumably a part of the type of V.
Califoniicwn or at least have been compared with Durand's speci-
mens and the notes made by Durand himself Durand in his de-
scription expressively states that the lower stem-leaves are petioled.
As this is never the case in the plant of the northern Rockies and
the Columbia Valley, I am certain that the plant generally re-
garded as V. Californicwn and from which Dr. Watson's descrip-
tion in his revision was mainly drawn is perfectly distinct from
Durand's plant. The latter is a rare plant judging from the fact
that I have not seen more than the two specimens mentioned above.

V . spcciosuin ranges from Montana to Washington, California
and Colorado, reaching a maximum altitude of 2,500 m.

Montana: Bridger Mountains, 1896, Flodinan, j^/ (type);
Little Belt Mountains, j-f-f.}^ / Bozeman, P. Koch ; Deer Lodge
Co., Ein?n2 Ware; Belt Park, 1886, R. S. Williams, 47^; Belt
Creek, 1883, Scribner, 28 j ; Lo-Lo Creek, 1880, Watson.

Idaho: Lake Pend d'Oreille, 1892, Sandberg, MacDoiigal &
Heller, 741 ; Kootenay Co., 1886, /. H. Sandberg ; Lake Waha,
1896, A. A. & E. Gertrude Heller, jj8o ; 1892, Isabel ATulford.

Utah : Heber Valley, 1869, S. Watson, ii6j ; American Fork
Canon, 1880, M. E.Jones.

533 RvDBERG : Studies on Rocky Mountain Flora

Washington: 1889, G. R. Vasey.

Oregon : /. 5. Nciobcny.

California: Modoc Co., 1893, M. S. Baker; i86s, B'. iV.
Bolander, 62jj ; Mt. Shasta, 1897, H. E. Broivn.

Colorado: Pagosa Peak, 1899, C. F. Baker, 2j8 Q). (This
has smaller flowers, petals and sepals being only 6-8 mm. long.)

ZYGADENUS Michx.

Much confusion has existed in this genus. The species of
Colorado, Wyoming and Montana have been variously named Z.
Nuttallii, Z. veneiiosiis and more rarely Z. panieulatiis, but much
complaint has been made that they do not fit Watson's descrip-
tions. Some have complained that those descriptions haye been
too narrow, others that there is no line to be drawn between Z.
venenosus and Z. panicidaUis and that both those species come to-
near Z. Nuttallii. Dr. Watson's descriptions of Z. venenosus and
Z. paniculatus are unusually good and correct. He knew them
both from the field. The common Rocky Mountain plants belong
to neither. Z. venenosus does not grow east of Idaho and Z. pan-
iculatus not east of Utah, while Z. Nuttallii is a species of the plains
and I doubt that it is found at all in the Rocky Mountain States.
The Rocky Mountain species are distinguished as follows :

Gland obcordate ; base of ovary inferior.

Petals and sepals 7-8 mm. long, 7-13-nerved. Z. elegans.

Petals and sepals 5-6 mm. long, 3-7-nerved. Z. Coloradcnsis.

Gland obovate or semi-orbicular ; ovary wholly superior.

Petals and sepals more or less clawed ; filaments adnate to the base of the claws.
Petals and sepals rounded or obtuse at the apex.

Upper leaves without sheaths at the base ; both petals and sepals long-
clawed and sub-cordate at the base*; gland with a thick margin.

Z. venenosus.
All leaves with distinct sheaths ; petals long-clawed and sub-cordate at
the base ; sepals short-clawed ; margin of the gland ill-defined.
Petals and sepals 4-5 mm. long ; petals ovate ; leaves 3-5 mm. wide.

Z. gramineiis.
Petals and sepals 6-8 mm. long ; petals oblong ; leaves 5-9 mm.
wide. Z. intermedins.

Petals and sepals acute or acuminate at the apex ; all leaves with sheaths ;
sepals cuneate at the base and short-clawed.
Leaves less than 5 mm. wide ; petals and sepals both cuneate at the base
and short-clawed. Z. acuttis.

Rydberg: Studies on Rocky Mountain Flora 534

Leaves over 5 mm. wide.

Raceme simple ; petals long-clawed and sub- cordate at the base.

Z. falcatttm.
Raceme branched ; petals short-clawed, not sub-cordate at the base.

Z. paniculatuiii.
Petals and sepals clawless ; gland obovate with a poorly defined margin ; fila-
ments free. Z,. Nuttallii.

Zygadenus elegans Pursh, Fl. Am. Sept. i : 241. 18 14
The typical Z. clcgans is a western plant, mainly belonging to
the Rocky Mountain region, where it grows in meadows up to an
altitude of 2500 m. If the eastern plant or Z. glmiais of Nuttall,
should be included in this species, is very doubtful. It is always
darker green, more glaucous, inflorescence more paniculately
branched, the petals and sepals narrower and more greenish. It
ranees from New Brunswick to Minnesota and south to Vermont.
The range of Z. elegans proper is from Saskatchewan to Alaska,
south to Colorado and Nevada.

Zygadenus Coloradensis sp. nov.

A rather slender glabrous plant, 2-4 dm. high. Bulb ovoid,
about 2 cm. long and i-l5 cm. in diameter: leaves narrow,
erect, about 2 dm. long, 3-5 mm. wide : flowers racemose, yel-
lowish white-tinged with brownish or purplish : bracts linear-
lanceolate, all equaling or exceeding the pedicels : petals and
sepals 5-6 mm. long, 3-7-nerved, oblong or narrowly obovate,
acute: capsule ovoid, 15-18 mm. long and 7-8 mm. in diameter.

Z. Coloradensis is closely allied to Z. elegans, differing in the
smaller flowers, greener foliage, long and narrow bracts equaling
or exceeding the pedicels and a brownish or purplish tint of the
inflorescence, bracts and flowers. It grows in the mountains at
an altitude of 2500-3500 m.

Colorado: Idaho Springs, 1895, Rydberg i\.y^€); Leroux
Creek, 1892, /. H. Coiven ; La Plata, 1873, Coidter ; Caribou,
1 89 1, Dr. E. Penard ; Empire, 1892, H. N. Patterson, 2g8.

Zygadenus venenosus Wats. Proc. Am. Acad. 14: 279. 1879

This species is characterized by its narrow leaves, of which the

upper lack the scarious sheaths, and by the thick, prominent

* When the petals or sepals are sub- cordate at the base and long- clawed the gland
is at the very base but when they are acute at the base the gland is a little higher up
on the blade.

535 Rydberg: Studies on Rocky Mountain Flora

glands. It is more slender and strict than all the species except
Z. acutus, which is easily distinguished by its acute sepals and
petals. Z. vencnosus grows mostly on hillsides at low altitudes,
scarcely ascending higher than 2000 m. It ranges from Idaho and
British Columbia to California and Utah. The following Rocky
Mountain specimens belong here :

Utah: Parley's Peak, 1869, S. Watson, ii6j.

Idaho: Keeley's Hot Springs, i^g2, Isabel Miilford; Big Pot-
lash River, 1892, Sandberg, MacDougal & Heller, Jig ; Little
Potlash River, ^o/.

Zygadenus gramineus sp. nov.

Zygademis vencnosus Rydb. Cont. U. Dep. Ag. 3 : 525. 1896.
Mem. N. Y. Bot. Card, i : 93 in part. Not S. Wats.

A slender yellowish-green plant, 2-3.5 elm. high. Bulb
elongated ovoid, 2-3 cm. long and 1-1.5 cm. in diameter: leaves
narrowly linear, scabrous on the margins and the midrib, 1-2 dm.
long, 3-5 mm. wide, conduplicate and somewhat falcate, all with
distinct scarious sheaths surrounding the stem : racemes rather
short : bracts scarious, lanceolate, long-acuminate : flowers light
yellow : sepals broadly ovate, obtuse at the apex, acute at the base
and very short-clawed : petals ovate, obtuse, subcordate at the
base and with claws about i mm. long : glands almost semi-orbic-
ular ; upper margin toothed, but thin and not well defined : cap-
sule elongated ovoid, 8- 10 mm.- long, 3-4 mm. in diameter.

Z. gramineus resembles somewhat the preceding, but is lower,
of a yellowish color ; its stem leaves are evidently sheathed, the
sepals are short clawed, not subcordate at the base and the gland
thin and without a distinctly thickened upper border. Z. gramineus
grows on hillsides up to an altitude of 2500 m. ; from Saskatchewan
and Alberta, south to western Nebraska and Idaho.

Montana: Spanish Basin, 1897, Rydberg &• Bessey, 3848
(type); 1896, Floduian, j^j ; Helena, 1891, F. D. Kelsey.

Idaho: Beaver Cafion, 1895, Rydberg.

South Dakota: Hot Springs, 1892, Rydberg, loji.

Wyoming: Yellowstone Park, 1888, Dr. Chas. H. Hall.

Saskatchewan: 1858, E. Boiirgeau.

Zygadenus intermedius sp. nov.
Zygadenus vencnosus Rydberg, Mem. N. Y. Bot. Gard. i : 93
in part. 1900. Not Nutt.

Rydberg: Studies on Rocky Mountain Flora 536

A rather stout light greeti plant, ^-6 dm. high. Bulb elon-
gated ovoid, about 3 cm. long and 1. 5 cm. in diameter: leaves
scabrous on the midrib and margins, 2 dm. or more long, 5-9
mm. wide, keeled and sometimes conduplicate ; all with conspicu-
ous scarious sheaths at the base : raceme rather long : flowers
light yellow ; petal and sepals 6-8 mm. long, obtuse ; the for-
mer broadly ovate, and acute or rounded at the base and short-
clawed ; the latter oblong subcordate at the base and with a claw
I mm. long : glands as in the preceding.

Like the preceding but taller and stouter, in habit resembling
mostly Z. panicidatiis, except that the inflorescence is seldom
branched. It is easily distinguished from that species by the ob-
tuse petals and sepals and by the distinct claws and the subcordate
bases of the petals. It grows on dry hillsides up to an altitude of
2000 m. in Montana, Idaho, Wyoming and Utah.

Idaho: Nez Perces Co., 1892,/. H. Sandbcrg, 10364 (type);
Peter Creek, 1892, Sandberg, MacDoiigal & Heller, 114; Lewis-
ton, 1896, A. A. & E. Gertrude Heller, 30^3.

Utah: Farmington, 1881, M. E.Jones, 2ogi.

Wyoming: Laramie Hills, 1894, Aven Nelson, 234.

Montana: Deer Lodge, 1895, F.N. Notestein ; Bridger Moun-
tains, 1897, Rydberg & Bessey,384g.

Zygadenus acutus sp. nov.

A very slender light green plant, 3-5 dm. high. Bulb rounded
ovate, 1.5-2 cm. long, 1-1.5 cm. in diameter: leaves narrowly
linear, scabrous on the margin, about 2 dm. long and 4-5 mm.
wide, keeled and often conduplicate : flowers pale yellow : petals
and sepals 4-5 mm. long ; both acute at the apex and at the base,
very short-clawed : glands obovate or cuneate ; upper margin
toothed, thin and not well defined.

In habit closely resembling Z. venenosns, but easily distin-
guished by the acute, short-clawed petals and sepals, which are
both cuneate at the base. The only specimens seen are the fol-
lowing :

South Dakota: Box Elder Creek, Black Hills, 1887, W. S.

Riisby.

Zygadenus falcatus sp. nov.
Z.Niittallii Porter & Coult. Syn. Fl. Colo. 133. In part.
1874. Not A. Gray.

537 RvDBERG : Studies ox Rocky Mountain Flora

A rather stout light green plant 3-4 dm. high. I^ulbs rounded
ovoid, 3-4 cm. long and about 3 cm. in diameter : leaves scabrous,
especially on the margins and midribs, 1.5-2.5 dm. long, 5-8 mm,
wide, keeled, conduplicate, and generally decidedly falcate, all with
scarious sheaths : raceme short, in fruit elongated : flowers yellow :
petals and sepals about 5 mm. long, acute ; the former deltoid
ovate, acute at the base ; the latter ovate, sub-cordate at the base
and with claws i mm. long : glands semi-orbicular, upper margin
toothed, thin and not well defined : capsule ovoid cylindrical.

All the specimens cited below have been named Z. Nuttallii
though the plant is more closely related to Z. paniciilatns, from
which it differs in the distinctly clawed petals which are subcordate
at the base. These characters, together with the more distinct
glands and the slightly adnate filaments, distinguish it from Z.
Nuttallii. Z. falcatus inhabits the foothills of Colorado at an alti-
tude of about 1500 m.

Colorado: Fort Collins, 1893, C. S. Cra ndall {tyT^t) ; 1896.
C. F. Baker ; Denver, 1873,/. M. Coulter.

Zygadenus paniculatus (Nutt.) Wats. Bot. King's Exp. 5 : 343,

1871

Helonias paniculata^nit. ]o\xn'\. Phila. Acad. 7: 57. 1834.

It is well characterized by Watson and easily distinguished by
its stout habit, generally branched inflorescence, and rhombic -
ovate acute and almost clawless petals and sepals. It grows on
hills up to an altitude of 1500 m., ranging from Montana and
Washington to New Mexico and California. The following Rocky
Mountain specimens belong here :

Utah: Ogden, Capt. Stansbury ; City Creek Canon, \Z%o, M.
E. Jones, i6j/j..

Idaho: Boise, \Z<^2, Isabel Mulford.

Montana: Grasshopper Valley, 1880, JVatson (?).

Zygadenus Nuttallii A. Gray, in Wats. Proc. Am. Acad. 14 : 279-

1879

Aniiantlmun Nuttallii Gray, Ann. Lye. N. Y. 4 : 123.

This is evidently a species belonging to the plains. All the
specimens in the Columbia and N. Y. Botanical Garden herbaria
are from Arkansas and Kansas. Dr. Watson included Texas and

Rydberg: Studies on Rocky Mountain Flora 538

Colorado in the range. The Texan plant referred here by him,
belongs to an altogether different plant. I think that Colorado
also should be excluded, believing that all specimens found there
and labeled Z. Nnttallii belong to Z. falcaijis, which resembles it
most in general habit, but has an altogether different flower.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 10

STUDIES ON THE ROCKY MOUNTAIN

FLORA— III

BY P. A. RYDBERG

NEW YORK
1900

1 Reprinted from the Bulletin of the Toeeey Botanical Club, 27 : 611,-636. 29 Dec, 1900. j

Studies on the Rocky Mountain Flora.— Ill

By r. A. Rydberg

SOME SMALLER GENERA OF COMPOSITES

STENOTUS Nutt.

This genus was established by Nuttall in 1840. It was re-
tained as a genus by Torrey and Gray in their Flora, but merged
with several others into Aplopappus by Gray in his Synoptical
Flora. A few years ago (1894), Professor Greene reestablished
the genus, removing from it, however, a member erroneously placed
in the genus by Torrey and Gray, viz., Stenotus pygmacus Torrey
and GiXZ-Y (^Aplopappus pyginaeiisGx2iy^,v^\\\z\\\\Q. referred \.o Macro-
nej/ia.^' If such a transfer was the very best is questionable, for
that species is aS" much a stranger in ]\Iacruneina as it is in Stenotus.
It has the outer bracts foliaceous and the style-appendages long
and attenuated which are characters found in Macroneina ; but the
habit is very unlike Macroneina and the outer foliaceous bracts are
numerous as in Pyrroconia. I had some transient thought of trans-
ferring it to that genus ; but the purely white pappus, the densely
cespitose habit, and the lack of the thick taproot, debar it from
Pyrrocoma. These characters ally it to Solidago ; but the differ-
ence in structure of the outer and the inner bracts makes it un-
natural to place it there as well. It is very hard to decide which
would be the best course to take, either to place it as an anoma-
lous member of one of these genera or to make it the type of a new
genus. Perhaps some other and better relationship may be found.

Of the other species included in Aplopappus § Stenotus by
Gray, A. Parryils, I think, rightly referred to Solidago. A. Lyallii
was altogether omitted by Professor Greene, when he made the
segregation in Pittonia. He may have overlooked this, but it is
more probable that he omitted it, because he did not know where
to place it. The relationship is without any doubt closest with
Solidago, notwithstanding the solitary head.

* In a recent distribution of plants from Colorado determined by Professor Greene,
this was distributed under its original name Stenotus pygmaeus.

6X4

615 Rydberg : Studies on the Rocky Mountain Flora

By removing Stenotus lineavifolins Torr. & Gray and S. interior
Greene, which constitute a good genus, the genus Stenotus be-
comes a very natural one, with only one somewhat aberrant mem-
ber, ^. lanuginosus which approaches Pyrrocoma imdoides and its
relatives in habit.

The Rocky Mountain species are distinguished as follows :

Plant glabrous or puberulent ; leaves firm and evergreen.

Leaves linear to filiform, i. S. sienophyllus.

Leaves mostly oblanceolate.
Bracts lanceolate, acute.
Plant puberulent.

Bracts in 3 series, broad, with broad scarious margins.

2. S. acaulis.
Bracts in 2 series, narrow, with narrow scarious margins.

3. .S*. Andersonii.

Plant glabrous.

Stem-leaves oblanceolate, 1-2 cm. long. 4. S. caespitostis.

Stem-leaves linear, 4-7 cm. long. 5. S. falcatus.

Bracts oval or oblong, very obtuse. • 6. S. aniierioides.

Plant floccose ; leaves softer, not evergreen. 7. S. hvuigino.us.

1. Stenotus stenophyllus (A. Gray) Greene, Erythea, 2 : 72.

1894
Aplopappns stenophyllus A. Gray, U. S. Expl. Exp. 17 : 347.

1862-74.

This species grows on stony hills and mountains and ranges
from western Idaho and Washington to California.

2. Stenotus acaulis Nutt. Trans. Am. Phil. Soc. II. 7 : 334.

1840

Chrysopsis acaulis Nutt. Journ. Phil. Acad. 7:33. 1834.
. Aplopappns acaulis A. Gray, Proc. Am. Acad. 7 : 353. 1867.

This is a rather common species, growing on dry rocky hills
and mountains at an altitude of 1000-2500 m., from Saskatche-
wan and Washington to Wyoming and California.

3. Stenotus Andersonii sp. nov,

A puberulent, cespitosc, but less woody perennial than the pre-
ceding and the three next following species. Flowering stems
about I dm. high, leafy at the base, few-leaved above : leaves nar-
rowly oblanceolate, not very rigid, distinctly 3-ribbed, 3-5 cm.
long, 3-4 mm. wide : bracts narrowly lanceolate, acute, glandular

RvDHERG : Studies on the Rockv Mountain Flora 616

puberulcnt, with a very narrow scarious margin, imbricated in 2
scries : achenes somewhat fusiform, about 4 mm. long, grayish or
white villous as in the other species.

This species is nearest related to the preceding, but character-
ized by the less rigid leaves, the fewer and narrower involucral
bracts, the involucre being more that of vS". /aniiginosits.

The type was collected on dry open hills.

Montana: Belt Mountains, 1886, /^ VV. Anderson, j ^6 1*

4. Stenotus caespitosus Nutt. Trans. Am. Phil. Soc. II. 7 : 335.

1840
Clirysopsis cacspitosa Nutt. Jour. Acad. Phila. 7 : 33. 1834.
Aplopappns acaulis var. glabratus D. C. Eaton, King's Exped.
5 : 161. 1871.

This grows on dry hills and mountains, at an altitude of 1000-
3000 m., from Montana and Idaho to Nevada, Arizona and
Wyoming.

5. Stenotus falcatus sp. nov.

A glabrous cespitose perennial, with a woody caudex, resem-
bling 5. armerioidcs in habit. Flowering stems 10-15 cm. high,
leafy, 1-3-cephalous : basal leaves oblanceolate, rigid, obtuse or
acute, 4-5 cm. long, 4-6 mm. wide, 3-ribbed ; stem leaves linear,
4-7 cm. long, 2-3 mm. wide, more or less falcate : heads about i
cm. high : bracts lanceolate, acute, with scarious margin : rays 6— 7
mm. long, 2.5-3 "im. wide.

This species differs from .S". arvierioides, which it closely re-
sembles in the narrower and acute involucral bracts. It grows in
barren soil at an altitude of about 1500 m.

Utah : Red Creek, 1877, Dr. E. Palmer, 202 (type) ; Milford,
1880, M. E.Jones, 1804.

6. Stenotus armerioides Nutt. Trans. Am. Phil. Soc. II. 7 :

335- 1840

Aplopappns armerioides A. Gray, Syn. Fl. i- : 132. 1884.

This species grows on dry hills and bad-lands, at an altitude of
1000-2000 m., from Manitoba and Assiniboia to Utah, New
Mexico and western Nebraska.

* Unless otherwise stated, the types of the new species described are preserved in the
herbaria of the New York Botanical Garden or of Columbia University.

617 Rydberg : Studies on the Rocky Mountain Flora

7. Stenotus lanuginosus (A. Gray) Greene, Erythea, 2 : 72.

1894.

Aplopappiis lanuginosus A. Gray, U. S. Expl. Exped. 17 : 347.
1862-74.

This species is a rather anomalous member of the genus, as
the leaves are not rigid, and scarcely evergreen. In habit it re-
sembles somewhat some species of Pyrroconia^ as for instance, P.
innloidcs ; but it has the thin involucral bracts and the white pap-
pus of Stcnoiiis. It grows on the mountains of Washington,
northern Idaho and Montana.

Stenotopsis gen. no v.

Low shrubs with fastigiate branches, narrow fasciculate glan-
dular punctate leaves and large peduncled heads. Involucre.broadly
hemispherical ; its bract subequal, almost in a single series, linear
lanceolate, thin scarious-margined, not at all hebaceous. Recep-
tacle naked, alveolar. Ray-flowers about 12, ligulate, yellow,
large, pistillate and fertile. Disk-flowers perfect : their corollas
tubular-trumpet shaped, deeply 5-lobed. Anthers obtuse at the
base. Style branches stigmatic their whole length, with ovate to
lanceolate-subulate appendages. Achenes densely silverj'-villous.
Pappus of white capillary bristles, rather deciduous.

The genus contains two known species. It differs mainly from
Stenotus in its shrubby habit and glandular punctate leaves.

Leaves 3-4 cm. long ; rays II-14 mm. long. I. S. Utiearifolius.

Leaves 1-2 cm. long ; rays 9-II mm. long. 2. .S". interior.

1. Stenotopsis linearifolia (DC.)

Aplopappus linearif alius DC. Prod. 5 : 347. 1836.
Stenotus linearif alius Torr. & Gray, Fl. N. Am. 2 : 238. 1842.
This species is a shrub 3-10 dm. high, and grows on dry
hills, from Utah to California and Arizona.

2. Stenotopsis interior (Coville)
Aplapappus interior Coville, Proc. Biol. Soc. Wash. 7: 65.

1892.

5/^«<3///.y /;//m^r Greene, Erythea, 2 : 72. 1892.

Its range is further southwest, from southern Utah to Arizona
and southern California.

RvDBERG : Studies ox the Rocky Mountain Flora 618

MACRONEMA Nutt.

This genus was also established by Nuttall in 1840 and re-
established by Greene. The latter included, as is stated before,
also Stcnotns pygmacus Nutt. [Ap/opappus pygmacus A. Gray).
As the relationship of that species is rather doubtful, I have not
included it in the treatment below. The species can be separated
as follows :

Heads radiate.

Heads small, fastigiate clustered ; disk 5-8 mm. in diameter.

Leaves broadly obovate, mucronale-cuspidate. I. M. obovatum.

Leaves oblanceolate, acute or pointed. 2. M. IVatsonit.
Heads larger, solitary : disk I cm. or more in diameter.

Leaves oblanceolate, acute. 3- ^^- szcffruticosum.

Leaves spatulate, obtuse or niucronate. 4- ^1-^- grindelifolium.
Heads discoid.

Leaves oblong-oblanceolate.

Outer bracts oblong, acute. 5- M- discoideum.

Outer bracts broadly oblong, obtuse. 6. M. obtusum.

Leaves linear. 7- ^^- H'leare.

I. Macronema obovatum sp. nov.

A glandular-puberulent undershrub, 2-3 dm. high, with light
yellow bark on the branches. Leaves broadly obovate, 1-2 cm.
long, 7-10 mm. wide, mucronate-cuspidate : heads usually 2-3
together, about 10 mm. high, and 8 mm. broad : its bracts firmer
than in the other species, oblong-linear, abruptly obcuneate-acute
at the apex, unequal, imbricated in about 4 series : rays short, 4-5
mm. long and 1-1.5 mm. wide, about 10 in number.

The species is nearest related to M. Watsonii, differing mainly
in the broad leaves and the abruptly acute bracts. The type was
collected at an altitude of 1600 m.

Utah : City Creek Canon, M. E. Jones, 108 1.

2. Macronema Watsonii (A. Gray) Greene, Erythea, 2 : 74.

1894
Aplopappus Watsonii K. Gray, Proc. Am. Acad. 16 : 79. 1881.
This grows on mountains at an altitude of 2000-21500 m., in
Utah and Nevada.

3. Macronema suffruticosum Nutt. Trans. Am. Phil. Soc. II.

7: 322. 1840
Aplopappus suffniticosus A. Gray, Proc. Am. Acad. 6 : 542.
1865.

619 Rydberg : Studies on the Rocky Mountain Flora

This species grows in the mountains of Oregon and California
up to an altitude of 3000 m. It has also been reported from
Wyoming, but possibly some specimens of the next have been mis-
taken for it.

4. Macronema grindelioides Rydberg, Mem. N. Y. Bot. Garden,

I : 384. 1900

The habitat of this species is rocky places on mountain-sides,
at an altitude of 2500-3000 m. It grows in Montana, Idaho and
northern Wyoming.

5. Macronema discoideum Nutt. Trans. Am. Phil. Soc. II. 7 ;

322. 1840

Aplopappus Macroncimi A. Gray, Proc. Am. Acad. 6 : 542,
1864.

This species grows on the higher mountains at an altitude of
2500—3000 m., from Wyoming and Colorado to California.

6. Macronema obtusum sp. nov.

A dense glandular-pubescent undershrub, 2—4 m. high, with
white-tomentose branches. Leaves oblong-oblanceolate, about
3 cm. long and 6 mm. wide, obtuse or mucronate : heads about
18 mm. high, 10-18 mm. in diameter: their bracts linear, acute,
scarcely more than half as long as the flowers, subequal, except
the outermost, which are foliaceous, broadly oblong, obtuse or mu-
cronate : rays none.

The species is closely related to the preceding, differing in the
stouter habit and the larger and broader, more obtuse outer bracts.
It grows on high mountains at an altitude of about 2500 m.

Colorado: South Cottonwood Gulch, 1892, C. S. Sheldon,
^8j8 (type); Twin Lakes, i?>y'i,,John Wolfe, 451.

7. Macronema lineare Rydb. Mem. N. Y. Bot. Garden, i : 384.

1900
In habit, this species resembles some species of CJirysotJiainmis
and Prof. Aven Nelson insists that it should be referred to that
genus. It has, however, the foliaceous outer bracts and long style-
appendages of Macronema and is clearly congeneric with the two
preceding species. It cannot very well be referred to Chrysotham-
mis, for it lacks the most essential character of the genus, viz., the

RvDBERG : Studies on the Rocky Mountain Flora 620

arran"-ement of the involucral bracts in distinct vertical rows. It
grows in gravelly places in northern Wyoming, at an altitude of
2000-2500 m.

SIDERANTHUS Fraser

The name SidcrantJius appears first in Fraser's Catalogue, but
there only as a nomen nudum. Pursh in his Flora on page 750 gives
Siderantlms integrifolius Fraser and 5. pinnatifidiis Fraser as syno-
nyms oi Amcllus villosus and A. spinulosiis described on page 564.
There may be a doubt as to which of these should be regarded as
the type of Siderantlms. Ajiwllns villosus with its relatives was
made a genus CJirysopsis by Nuttall in 1818 or by Elliott in
1824, according to different interpretations, long before Eriocavpwn
was established (1840). This leaves SiderantJius pi/iiiatifidiis as
the residue of the genus Sideranthus. Besides Nuttall, who was
the real author of Fraser's Catalogue, made Sideranthus a subgenus
of Dieteria containing the only species D. spinulosa {Aplopappiis
spinulosus DC. See Trans. Am, Phil. Soc. II. 7 : 301) ; and thus
shows that he regarded it as the type of Sideranthus.^

Leaves spinescent toothed, not pinnatifid.

Heads discoid ; perennial with woody caudex. i. S. grindelioides.

Heads radiate ; annual. 2. S. rubiginosus.

Leaves pinnatifid.

Stem more or less fioccose, or cinereous especially when young.

Plant cinereous pubescent, more or less glandular. 3. S. australis.

Plant more or less floccose, not at all glandular. 4. .S". spinulosus.

Plant neither floccose nor cinereous.

Plant perfectly glabrous or sparingly glandular puberulent.

5. S. glaberrimus.
Plant finely puberulent.

Heads hemispherical ; bracts slightly glandular. 6. S. puberulus.

Heads somewhat turbinate ; bracts densely glandular puberulent.

7. S. turbinellus.

I. Sideranthus grindelioides (Nutt.) Britton

Eriocarpujn grindelioides Nutt. Trans. Am. Phil. Soc. II. 7 :
321. 1840.

Aplopappus Nuttallii Torr. & Gray, Fl. N. A. 2 : 240. 1842.

* Professor Greene (Pittonia 2 : 115) seems to think Sideranthus is derived from
the Latin sidiis, star, and the Greek, avi?of, flower. It is better to regard the first part
also as Greek, ai6r/pog, iron. Why accuse Nuttall of making a hybrid word, which
we would not permit ourselves ?

621 Rydberg : Studies on the Rocky Mountain Flora

The species is not uncommon on barren hills and in bad-lands,
up to an altitude of 1500 m. Its range extends from Saskat-
chewan and western Nebraska to Colorado and Arizona.

2. Sideranthus rubiginosus (Torr. & Gray) Britton
Aplopappus rubiginosum Torr. & Gray, Fl. N. Am. 2 : 240.
1842.

Eriocarpiini riibiginos2ini (Torr. & Gray) Britton, Mem. Torr.
Bot. Club, 5: 316. 1894.

The species was first described as a perennial. Our plant is
evidently only annual. Possibly two different species have been
confused under the name E. rubiginosinn. It grows in sandy soil
from Texas to Colorado and western Nebraska, reaching an alti-
tude of 1500 m.

3. Sideranthus australis (Greene)

Eriocarpwn aiistrale Greene, Erythea, 2: 108. 1894.

This grows on the plains of western Texas, Colorado, New
Mexico and Mexico.
4. Sideranthus spiNULOsus (Pursh), Sweet, Hort. Brit. 227. 1826

Avielliis spimilosus Pursh, Fl. Sept. Am. 564. 18 14.
Sideranthus pinnatifidus Fraser; Pursh, Fl. Am. Sept. 750. 1 8 14
Aplopappus spimdosus DC. Prod. 5 : 347. 1836.
Eriocarpum spinulosiini {Vuvsh) Greene, Erythea, 2: 108. 1894.
The species is rather common on plains and dry prairies, from
Saskatchewan, Nebraska and Texas to Mexico, Arizona and Idaho.

5. Sideranthus glaberrimus sp. no v.

Perennial, with a short woody caudex, perfectly glabrous, or
very sparingly glandular-puberulent. Stems several, branched
above, very leafy, 1-3 dm. high : leaves rather rigid, about 2 cm.
long, bluish green, glabrous, pinnatifid ; segments oblong, 2-5 mm.
long and i mm. or less wide, spinulose-tipped : heads numerous,
corymbose, depressed hemspherical, 8-10 mm. high and 8-15 mm.
broad : bracts imbricated in 6-7 series, the outer gradually shorter,
firm, appressed, with a herbaceous tip, acute ; the inner tinged with
purplish : rays about 20, 4-5 mm. long and about i mm. wide.

The species is closely allied to the preceding, differing mainly
in the total lack of tomentum, even when young, the bluer color
and the smaller, generally purple-tinged bracts. It grows on
plains and hills, from Iowa and Indian Territory to New Mexico
and Wyoming.

RvDBERG : Studies on the Rocky Mountain Flora 622

Kansas : Osborn City, 1894, C.L. Shear, 116 ; Harper, 1888,
W. A. Kellcnnan ; Hamilton Co., 1895, Hitchcock, 222.

Nebraska : 1893, Rydbcrg, 14-Oja (type) ; G. D. Szveezey, ^^.
Indian Territory : 1868, Edivani Palmer, 44.2.
Colorado : Platte, Dr. James.
Wyoming: Platte, Fremont, Dayton, 1899, Tzveedy, 2oy6.

6. Sideranthus puberulus sp. nov.

A lo\v densely puberulent perennial, less than 1.5 dm. high :
Stems ascending, branched above : leaves 2-3 cm. long, pinnatifid
or bipinnatifid ; segments 1-5 mm. long, less than i mm. wide,
spinulose-tipped : heads 2-4 in a small corymb, hemispherical,
8-9 mm. high, 10-12 mm. broad: bracts narrowly linear-lanceo-
late, appressed, acute, with small herbaceous tips, finely puberu-
lent, but only slightly glanular : rays about 20, 8-9 mm. long and
about 1.5 mm. wide.

This is perhaps nearest related to E. australe ; but is easily dis-
tinguished by the fine pubescence.

Colorado : Salida, Mrs. C. B. Clarke, ly^..

7. Sideranthus turbinellus sp. nov.

A low and bushy puberulent perennial, 1.5-2 dm. high. Stems
corymbosely branched and very leafy : leaves . 5-2 cm. long, pin-
natifid or the upper reduced and merely toothed ; segments narrow,
1-5 mm. long, about .5 mm. wide : heads numerous, small, hemi-
spherical-turbinate, 7-8 mm. high, and scarcely as broad : bracts
imbricated in 6-7 series, linear-lanceolate, acute, appressed gland-
ular-puberulent : rays 10-15, 7-8 mm. long and 1.5 mm. wide.

Perhaps nearest related E. spumlos?im, the species is easily
known by the small, somewhat turbinate heads and the dense and
fine pubescence.

Idaho: Pocatello, 1892, A. Isabel Mnlford.

PYRROCOMA Hook.

This genus was established by Hooker in 1840, and based on
one species, P. carthavwides. In 1 894, Professor Greene extended
the genus so as to include the genus Homopappus of Nuttall, the
difference between the two genera being only that the ligules of
the ray-flowers in the former are very small and inconspicous or
wanting.

623 Rydberg : Studies on the Rocky Mountain Flora

Heads apparently discoid, the sterile rays being concealed in the pappus.

Inner bracts recurved cuspidate. I. P. subsquarrosa.

Bracts not recur%'ed cuspidate.

Bracts oblong or oval ; heads hemispherical.

Leaves and bracts more or less spinulose-toothed, the latter with a nar-
row scarious margin. 2. P. carthamoides.
Leaves and bracts not spinulose-toothed, the latter with a broad scarious
erose margin.
Pappus sordid ; plant almost glabrous. 3. P. rigida.
Pappus brownish red ; plant villous. 4. P. erythropappa.
Bracts lanceolate ; head campanulate-turbinate. 5- P- Ctisickii.
Heads distinctly radiate.

Bracts except the innermost obtuse, obovate or oblong. 6. P. crocea.

Bracts mostly acute or acuminate.
Plant not glandular.

Heads large ; disk 2 cm. or more in diameter ; bracts in about 3 series.
Upper part of stem and involucre decidedly villous ; bracts wholly
foliaceous.
Bracts oblanceolate, abruptly acute. 7. P. dementis.

Bracts lanceolate, long-acute. 8. P. villosa.

Stem and involucre almost glabrous ; bracts chartaceous at the base
with foliaceous tips. 9. P. integrifolia.

Heads smaller ; disk less than 2 cm. in diameter.
Bracts in 2-3 unequal series.

Leaves 5-15 mm wide ; disk 1-2 cm. wide.

Inflorescence corymbiform ; bracts long-acute.

10. P. lanceolata.
Inflorescence racemifomi ; bracts abruptly acute,

11. P. Vaseyi.
Leaves 2-4 mm. wide ; disk I cm. or less wide.

12. P. teiiuicaulis.
Bracts nealrly of the same length.

Stem-leaves oblanceolate ; bracts long-acuminate.

13. P. acuminata.
Stem-leaves lanceolate ; bracts acute or short-acuminate.

Plant villous. 14. P. imdoides.

Plant, except the. upper parts, glabrate in age.

15. P. unijlora.
Plant decidedly glandular. 16. P. hirta.

I. Pyrrocoma subsquarrosa Greene, Erythea, 3: 22. 1895
I have seen no specimen of this species ; but the description
indicates a plant wholly unlike the other species of the Rocky
Mountains, that I do not hesitate in accepting it as a good spe-
cies. The type of P. sidnqiiarrosa was collected in northern
Wyoming, by Dr. J. N. Rose in 1893.

2. Pyrrocoma carthamoides Hook. Fl. Bor. Am. i : 307. 1833
Aplopappus carthamoides A. Gray, Proc. Acad. Sci. Phila.
1863 : 65. 1864.

Rydberg : Studies on the Rocky Mountain Flora 624

As treated by Dr. Gray, A. cartliamoidcs comprises more than
one species, and probably both of the two following species are
covered by his description. There is no doubt, however, as to
which species shall bear the name, for Hooker's description and
figure point clearly to the plant with more or less spinulose-
toothed leaves and bracts. The latter are also longer, narrower
and more acute than* in the other two species. P. cartJiauioidcs
ranges from Alberta and British Columbia to Oregon and Idaho.

3. Pyrrocoma rigida sp. nov.

A perennial with a thick woody tap-root. Stems I-3, erect
or ascending, finely pubescent when young, glabrate in age, 2—4
cm. high, leaves pale bluish green, firm, oblanceolate, acute, entire-
margined, finely puberulent, or glabrate ; the lower petioled, 8-15
cm. long, 1.5-3 cm. wide; the upper smaller and sessile: heads
apparently discoid, 15-20 mm. high, 15-25 mm. broad: bracts
very firm, imbricated in 3-4 series, broadly oblong, acute or mu-
cronate ; the inner at least with broad scarious or semi-chartaceous
margins, which generally are erose, but not spinulose-dentate :
ligules of the ray-flowers erect, slightly longer than the light
brownish or sordid pappus : achenes glabrous, shining.

In general habit, this species resembles closely P. carthanioides
but has broader leaves and bracts, paler foliage and no indication
of spinulose toothing. It grows in sandy places and meadows in
Idaho, Montana and Washington.

Idaho : Granite Station, Kotenay County, 1 892, Sandberg,
MacDoiigal & Hclkr, ySj (type).

Montana: Columbia Falls, 1894, R. S. Williams.

Washington: Loomiston, 1897, A. D. E. Elmer, 6oj.

4. Pyrrocoma erythropappa sp. nov.

A finely villous pubescent perennial. Stem 1.5—3 ^"i- liigh>
with 1—4 heads : stem-leaves 3-6 cm. long, i — 1.5 cm. wide, firm,
oblanceolate or oblong, acute, subsessile, finely pubescent, or
glabrate on the upper surface : heads very short-peduncled in the
axils of the upper leaves, 10-15 mm. high, about 15 mm. in diam-
eter : bracts imbricated in 4-5 series, pubescent, broadly oval or ob-
long, obtuse, mucronate, or the outer acute, with a thin erose mar-
gin : pappus intensely brownish red : otherwise as the preceding.

Idaho : Clear Water, Rev. Spalding (type in Torrey Her-
barium).

625 Rydberg : Studies on the Rocky Mountain Flora

5. Pyrrocoma Cusickii (A. Gray) Greene, Erythea, 2: 59. 1894
Aplopappus cartliamoides var. Cusickii A. Gray, Syn. Fl. 2' :

126. 1886.

The range of this species is limited to Oregon and western
Idaho.

6, Pyrrocoma crocea (A. Gray) Greene, Erythea, 2 : 69. 1 894

Aplopappus croccHs A. Gray, Proc. Acad. Sci. Phila. 1863 : 65.
1864.

This species is not uncommon in the mountain regions of
Colorado at an altitude of 1800-3000 m.

7. Pyrrocoma Clementis sp. nov.

A perennial with more or less villous ascending stem, 1.5-4
dm. high: lower stem-leaves linear-oblong, about i dm. long,
somewhat fleshy, glabrous except the ciliate margin, saliently den-
tate ; the upper lanceolate or ovate-lanceolate, more or less auricled
at the base, dentate or entire-margined ; heads solitary ; disk about
15 mm. high and 2.5-3 cm. broad : bracts all foliaceous, imbricated
in 3-4 series, but the outer almost equaling the inner, oblanceolate,
abruptly acute, villous : rays 10-12 mm. long and 2-3 mm. wide,
bright yellow : achenes sparingly strigose-hirsute : pappus yellow-
ish white.

Colorado : Mt. Harvard, 1896, Frederick Clements, /j./}..

8. Pyrrocoma villosa sp. nov.

A low perennial with a thick woody tap-root. Stems 1-3, as-
cending, 1-1.5 dm. high, purplish, more or less villous, especially
above : basal leaves oblanceolate, 7-10 cm. long, 1-1.5 cm. wide,
somewhat glaucous and rigid, entire-margined or occasionally
slightly spinulose denticulate ; stem-leaves linear-lanceolate, 2-4
cm. long : heads solitary : disk about 2 cm. broad : bracts foliace-
ous, imbricated in 3-4 series, but the outer fully as long as the inner,
broadly linear or lanceolate, long-acute, more or less villous : rays
about I cm. long and 2 mm. wide : achenes glabrous : pappus
dirty white.

In habit this species resembles most P. iinijlora and P. imdoidcs ;
but is easily distinguished by the larger heads and the foliaceous
bracts in several series. It grows in meadows at an altitude of
about 2700 m.

Wyoming: Willow Creek, Big Horn Mountains, 1899, F.
Tweedy, 2o6j.

Rydberg : Studies on the Rocky Mountain Flora 626

9. Pyrrocoma integrifolia (Porter) Greene, Erythea, 2: 69. 1894
Aplopappus integrifolitis Porter; A. Gray, Proc. Am. Acad.

16: 79. 1881.

This species grows in meadows, at an altitude of 1000-2500

m., from Saskatchewan to Idaho and Wyoming.

Pyrrocoma integrifolia pumila var. nov.

Pyrrocoma Hozvellii Rydb. Mem. N. Y. Bot. Garden, I : 382.
1900. Not A. Gray.

Stem low, I dm. or less, monocephalous ; bracts shorter and
less acute.

Montana : Butte, 1895, Rydberg, 2808.

10. Pyrrocoma lanceolata (Hooker) Greene, Erythea, 2: 69.

1894

Donia lanceolata Hook. Fl. Bor. Am. 2 : 25. 1834.

Aplopappus lanceolatus Torr. & Gray, Fl. N. Am. 2 : 241 . 1 842.

The range of this species is from Saskatchewan to British Co-
lumbia, Nevada and Wyoming. It ascends to an altitude of

2000 m.

1 1. Pyrrocoma Vaseyi (Parry)

Aplopappus lanceolatus var. Vaseyi Parry ; D. C. Eaton, King's
Exped. 5 : 160. 1871.

I believe that this deserves a specific rank, as the racemose
disposition of the heads is accompanied with shorter, closer and
oblanceolate, abruptly acute bracts. It ranges from Saskatchewan
(according to Gray) south to Utah and Colorado.

12. Pyrrocoma tenuicaulis (D. C. Eaton) Greene, Erythea, 2:

69. 1894

Aplopappus tenuicaulis D. C. Eaton, King's Exped. 5: 160.
1871.

Aplopappus lanceolatus var. tenuicaulis A. Gray, Syn. Fl. 2^ :

129. 1884.

This grows in alkali meadows of Utah, Nevada and Oregon,
at an altitude of about 2000 m.

13. Pyrrocoma acuminata sp. nov.

A low finely villous perennial with a woody tap-root. Stems
several, slender, about i dm. high, monocephalous : leaves nar-

G27 RvDBERG : Studies ox the Rocky Mountain Flora

rowly linear-oblanceolate, with strong midrib, 2-4 cm. long, 2-4
mm. wide, entire-margined, acute, finely villous on both sides :
heads on naked peduncles, which are 3-4 cm. long : disk about
I cm. high and 12-15 mm. broad.: bracts in about 2 series, broadly
lanceolate, contracted into a long slender tip : rays about i cm.
long and 1.5 mm. wide : achenes pubescent: pappus tawny.

In habit this species resembles most P. imiloidcs, but differs in
the smaller leaves, the shorter pubescence and the acumination of
the bracts.

Wyoming: Fort Bridger, 1873, Dr. J. V. Carter i\.y^& in Co-
lumbia Herbarium).

14. Pyrrocoma inuloides (Hook.) Greene, Erythea, 2 : 60. 1894

Donia inuloides Hook. Fl. Bor. Am. 2: 25. 1834.

Aplopappus imdoides Torr. & Gray, Fl. N. Am. 2 : 241. 1842.

Dr. Gray included this in A. uniflorns, but I agree with Pro-
fessor Greene that it ought to be kept distinct. It ranges from
Montana to Idaho and Wyoming.

15. Pyrrocoma uxiFLORA (Hook.) Greene, Erythea, 2 : 60. 1894

Donia nniflora Hook. Fl. Bor. Am. 2: 25. 1834.

Aplopappus nniflorus Torr. & Gray, Fl. N. Am. 2: 241.
1842.

This species grows in alkaline meadows, up to an altitude of
2500 m., and ranges from Saskatchewan and Montana to Utah
and Colorado.

16. Pyrrocoma hirta (A. Gray) Greene, Erythea, 2: 69. 1894

Aplopappus hirtns K. Gray, Syn. Fl. 2': 127. 1884.
The range of this species includes eastern Oregon, Washington
and western Idaho, where it reaches an altitude of 1000 m.

BALSAMORRHIZA Hook.

Leaves entire or bluntly toothed, never pinnatifid.
Plant white-tomentose.

Leaves with entire margins or slightly undulate, oblong-cordate to hastate.

I. B. sagittata.
Leaves more or less distinctly toothed, ovate-lanceolate, with subcordate
base. 2. B. tomentosa.

Plant hirsute puberulent ; basal leaves cordate.

Rays linear, deciduous ; achenes glabrous. 3. B. dcltoidea.

Rays oval, becoming papery, and more or less persistent : achenes puberulent.

4. B. Careyana.

Rydberg : Studies on the Rocky Mountain Flora 628

Leaves mostly pinnatifid or at least incisedly toothed.
Plant canescent or white-tomentose.
Plants loosely white-tomentose.

Stem 1-3 dm. high ; segments of the leaves 1-3 cm. long, ovate, entire

or slightly toothed. 5- B. iiicana.

Stem 3 dm. or more high ; segments of the leaves 3-5 cm. long, lanceo-
late, coarsely toothed. 6. B. Jloccosa.
Plant finely canescent, tomentose only on the involucre ; .some of the leaves
merely toothed. 7. B. terebinthacea.
Plants more or less hispid, neither canescent nor tomentose.
Disk 3-4 cm. broad ; segments of the leaves mostly entire.

8. B, macrophylla.
Disk 2-2.5 cm. broad ; segments of the leaves mostly toothed.

9. B. Iiirsuta,

I. Balsamorrhiza sagittata (Pursh) Nutt. Trans. Phil. Soc. II.

7: 350. 1840

Bupthabiiiuvi saggittattun Fl. Am. Sept. 564. 18 14.

Growing on hillsides at an altitude of 1000-2500 m., this spe-
cies is not uncommon from Alberta and British Columbia to Cali-
fornia, Colorado and the Black Hills of South Dakota.

2. Balsamorrhiza tomentosa sp. nov.

A white-tomentose perennial with thick root ; but the tomen-
tum is shorter and finer than in B. sagittata. Basal leaves with
long petioles ; blades about 1 2 dm. long, ovate-lanceolate with
subcordate bases, i 5-20 cm. long, acute, coarsely toothed ; stem-
leaves generally two, including the slender petioles about i dm.
long, lanceolate to linear elliptic, acute at both ends : stem 3-4
dm. high, involucre densely floccose, over 2 cm. broad : outer
bracts half longer than the inner, reflexed : rays about 3 cm. long
and I cm. wide ; achenes glabrous.

Closely related to B. sagittata this species differs mainly in the
toothed leaves, shorter tomentum and longer outer bracts.

Wyoming : Headwaters of Tongue River in the Big Horn
Mountains, 1898, F. Tzvccdy, lo."^

3. Balsamorrhiza deltoidea Nutt. Trans. Am. Phil. Soc. II.

7: 351. 1840
The name of this plant is rather unfortunate, as the leaves are
rarely deltoid, but on the contrary usually broadly cordate. B.
deltoidea ranges from British Columbia to California and Idaho.

*A specimen collected by Tweedy on Teepee Creek in .1899 [no. 21 14), may
also belong here. It has smaller heads, not reflexed bracts and lanceolate leaves.

629 Rydberg : Studies on the Rocky Mountain Flora

4. Balsamorrhiza Careyana a. Gray, PI. Fendl. 81. 1849
This species grows on sandy plains of Idaho and Washington.

5. Balsamorrhiza incana Nutt. Trans. Am. Phil. Soc. II. 7:

350. 1840
Balsamorrliiza Hookcri var. incana A. Gray, Syn. Fl. 2' : 266.

1884.

This species grows on dry stony hills, up to an altitude of 2500
m., from Montana and Washington to California and Wyoming.

6. Balsamorrhiza floccosa sp. nov.

Balsamorrhiza Balsamorrhiza Rydb. Mem. N. Y. Bot. Garden,
I : 417. 1900. Not Heliopsis Balsamorrhiza Hook.

A loosely w^hite or gray-tomentose perennial with very thick
tap-root. Leaves mostly basal, floccose on both sides, 2-3 dm.
long, regularly pinnately divided to near the midrib : some some-
times only coarsely toothed : segments lanceolate, acute, 3-5 cm.
long coarsely toothed : stem-leaves 2, near the base, similar but
smaller, about i dm. long : stem scapiform, 3-5 dm. high, villous
or the upper portion densely floccose : involucre about 3 cm.
broad, densely floccose ; bracts numerous in several series, lanceo-
late ; the outer often spreading with recurved tips ; rays 3-4 cm.
long, about i cm. wide ; achenes glabrous, cuneate oblong, with
truncate apex.

This species has been mistaken for B. Balsamorrhiza (Hook.)
Heller or B. Hookeri Nutt., but the latter has much finer dissected
leaves and its pubescence is quite different. The latter is very
short and appressed, never consisting of long villous hairs, and
there is never dense wool at the base of the head as in this species.
B. Balsamorrhiza ranges from Washington to California ; but is
evidently not found in the Rocky Mountain region. All that I
have seen from the region and referred to B. Balsamorrhiza by
Heller, Holzinger and myself belong to B. floccosa. Those col-
lected by Parry, and if I am not mistaken, named by Gray, be-
long; to B. incana.

B. floccosa grows on hillsides at an altitude of 1000-2500 m.
The following specimens are in the New York herbaria :

Montana : Spanish Basin, Gallatin Co., 1897, Rydberg & Bcs-

so', 5175 (type).

Idaho : Lake Waha, 1896, A. A. & E. Gcrtrndc Heller, 3298 ;

RvDBERG : Studies on the Rocky Mountain Flora 630

Craig Mountain near Lake Waha, 1892, Sandbcrg, MacDougal &
Htilcr, 24S.

Wyoming : Headwaters of Tongue River, Big Horn Moun-
tains, 1898, F. Tiveedy, 11.

7. Balsamorrhiza terebinthacea (Hook.y Nutt. Trans. Am.

Phil. Soc. II. 7 : 349. 1840

Heliopsis? terebinthacea Hook. Fl. Bor. Am. i: 310. 1833.

In this species, as well as in the next, it often occurs that some
of the leaves are not pinnatifid, but merely coarsely dentate. It
is a rare plant growing in stony soil in eastern Oregon and western
Idaho.

8. Balsamorrhiza macrophylla Nutt. Trans. Am. Phil. Soc.

II. 7: 350. 1840

This grows on rocky hillsides in Wyoming, Utah and Idaho.

9. Balsamorrhiza hirsuta Nutt. Trans. Am. Phil. Soc. II. 7 :

349. 1840

This species is found at an altitude of 1000-2000 m, in the
dry regions of Utah to British Columbia and CaHfornia.

THELESPERMA Less.

Heads radiate.

Leaf-segments linear-filiform, i mm. or less wide.

Annual or biennial ; outer bracts subulate-linear, more than half as long as

the inner. I- T trifidum.

Perennial from a rootstock : outer bracts linear-lanceolate, half as long as the
inner or less. 2. T. teniie.

Leaf-segments linear, over I mm. wide ; plant perennial or the first only biennial.
Plant with tap-root, leafy throughout. 3. T. intennediiitn.

Plant with creeping rootstock or woody caudex ; leafy only near the base.
Involucre not cleft below the middle ; throat of the disk-flowers cam-

panulate, shorter than the lobes. 4- ^' ambigiium.

Involucre cleft below the middle ; throat of the disk-flowers cylindra-
ceous, longer than the lobes. 5. T. subHudum.

Heads discoid ; perennials with rootstock or woody caudex.

Plant less than 2 dm. high ; involucre with very broad scarious margins.

6. T. marginatitiii.
Plants 3-6 dm. high ; involucre with very narrow scarious margins.

7. T. gracile.

631 RvDBERG : Studies on the Rocky Mountain Flora

I. Thelesperma trifidum (Poir.) Britton, Trans. N. Y. Acad.

Sci. 9 : 182. 1890

Coreopsis trifida Poir. Suppl. Lam. Encycl. 2: 353. 181 1.

Thelcspenna filifoliuui A. Gray, Kew Journ. Bot. i : 253.
1849.

This species grows in dry soil from Nebraska to Texas and
eastern Colorado.

2. Thelesperma tenue sp. nov.

A slender, glabrous plant with perennial rootstock. Stems
1—3, slender, less than 2 mm. in diameter, light green, mostly sim-
ple and leafy to near the top, 1.5—4 dm. high : leaves twice pin-
nately dissected into linear filiform segments, mostly appressed to
the stem: peduncles 1-3, 5-10 cm. long : outer bracts 6-8, linear-
lanceolate, half as long as the inner or less ; the inner united to
the middle, broadly scarious-margined : rays 8—12 mm. long, 4—6
mm. wide : awns of the pappus very short, about half as long as
the width of the summit of the dark shining achenes.

This species resembles most the preceding but has a simpler
stem and perennial rootstock. It grows in sandy soil at an alti-
tude of about 2800 m.

Colorado : Veta Pass, 1900, Rydberg& Vreclaiid, S47J (type);
1870, Dr. G. IV. Hulsc : Plains, 1871, Win. M. Cimby.

3. Thelesperma intermedium sp. nov.

A glabrous bushy plant with a biennial or perhaps perennial
tap-root. Stems several, much branched and very leafy, 2—6 dm.
high : leaves once or twice pinnately divided into linear segments,
1-3 mm. wide: peduncles very numerous, 1-1.5 dm. long: invo-
lucre about I cm. broad : outer bracts very narrowly linear-lanceo-
late, about half as long as the inner; these united to about the
middle, scarious-margined: rays 10-12 mm. long, 6-8 mm. wide,
rounded, 3-toothed at the apex : teeth of the pappus longer than
the width of the achenes.

This has been confused with T. anibigiium, but is of a quite

different habit. T. interinedimn is much branched and very leafy

throughout, has a vertical tap-root of short duration, numerous

heads, rather long outer bracts and the inner with narrow scarious

margins. T. auibigiiuui is leafy only at the base, has a creeping

rootstock, very short outer bracts and the inner with a very broad

scarious margin.

Rydberg : Studies on the Rocky Mountain Flora 632

T. iiitcnncdiinn grows in sandy or loose soil, often taking pos-
session of old fields and acts much like a weed. The following
specimens are at hand.

Nebraska : Banner County, 1 890, Rydberg, ig2 (type) ;
Crawford, 1889, H.J. Webber.

Wyoming: Pine Bluffs, 1897, Aven Nelson, Jjoj.

Colorado : 1862, //a// & Harbour, 2S0, at least in part.

Colorado or Wyoming : 1843, Fremont.

New Mexico : Between Santa Fe and Canoncito, 1897, A. A.
& E. Gertrude Heller, 3747.

4. Tiielesperma ambiguum a. Gray, Proc. Am, Acad. 19: 16.

1883
This species seems to be confined to the plains of western
Texas, New Mexico and southern Colorado. The specimens re-
ported from Nebraska and Wyoming belong mostly to the preced-
ing, and those from Montana to T. marginatum. The latter
resembles T. ambiguum in many respects, especially as to the in-
volucral bracts ; but it is always without ray-flowers.

5. Thelesperma subnudum a. Gray, Proc. Am. Acad. 10 : 72,

1874

This is an inhabitant of the more arid regions of New Mexico,
Arizona, Utah and Colorado.

6. Thelesperma marginatum Rydb. Mem. N. Y. Bot. Garden, i :

421. 1900

This grows on dry plains of Alberta and Montana.

7. Thelesperma gracile (Torr.) A. Gray, Kew Journ. Bot. i :

253. 1849

Bidens gracilis Torr. Ann. Lye. N. Y. 2 : 215. 1827.

This species is the most common and most widely distributed
species of the genus. It is common on dry plains from Nebraska
to Montana, Arizona, Mexico and Texas.

HYMENOPAPPUS L'Her.

Throat of the corolla 1-I.5 mm. long, not over twice as long as the lobes.
Pappus over l mm. long, equaling the corolla-tube or nearly so.
Stem 3-6 dm. high, leafy throughout ; heads numerous.

633 Rydberg : Studies on the Rocky Mountain Flora

Plant sparingly and loosely floccose ; leaves glabrate in age.

1. //. tenuifolius.
Plant densely tomentose ; leaves permanently tomentose.

2. H. tomentosiis.
Stem less than 3 dm. high.

Stem-leaves much reduced ; stem white-tomentose or nearly so ; heads

few. 3. //■ scapostts.

Stem-leaves not much reduced ; stem sparingly grayish tomentose.

5. H. cineretis.
Pappus I mm. or less long, shorter than the corolla-tube ; stem-leaves and heads
few.
Pappus not hidden by the hairs of the achenes.

Stem permanently densely white-tomentose ; achenes silky.

4. H. nrenosus.
Stem sparingly grayish tomentose, glabrate in age ; achenes loosely villous

5. H. cinereus.
Pappus hidden by the hairs of the achenes, or sometimes none.

Leaflets glabrate in age, at least above ; ultimate segment 5-30 mm.

long. 6. H.filifolins.

Leaves permanently densely white-tomentose ; ultimate segments short,
1-5 mm. long. 7. H. luteus.

Throat of the corolla 3-4 mm. long, 3-4 times as long as the lobes. 8. H. inacroglottis.

I. Hymenopappus tenuifolius Pursh, FI. Am. Sept. 742. 18 14

This is generally described as a biennial ; occasionally the
root is of a longer duration and the plant becomes a short-
lived perennial and then hard to distinguish from a large specimen
of H. filifoliiis except by the pappus. H. temiifolius grows on
prairies, from Nebraska and Wyoming to Texas.

2. Hymenopappus tomentosus sp. nov.

A densely and permanently white-tomentose plant, apparently
biennial. Stem 3-4 dm. high, leafy and branched above : leaves
5-7 cm. long, bi-pinnately divided into linear segments, 4-10 mm.
long, about i mm. wide : heads many, corymbose-paniculate, about
8 mm. high and broad ; involucre densely woolly, somewhat tur-
binate : flowers yellow: corolla tube and throat each a little over
I mm. long ; the latter broadly campanulate, of about the same
length as the lobes ; achenes silky strigose ; scales of the pappus
a little shorter than the corolla-tube.

Nearest related to the preceding, this species is easily distin-
guished by the dense permanent tomentum and the yellow flowers.

Utah: St. George, 1877, Dr. E. Palmer, 2jo (type in the
Columbia Herbarium).

Rydberg : Studies on the Rocky Mountain Flora 634

3. Hymenopappus scaposus sp. nov.

Hymenopappiis lutcus A. Gray, PI. Fendl. 97. 1849. Not
Nutt. 1 84 1.

An almost scapose perennial with a (espitose caudex. Stem
almost naked, with 1-2 reduced leaves, more or less densely to-
mentose : leaves mostly basal, 5-7 cm. long, bipinnately divided
into linear segments, 3—10 mm. long, more or less densely tomen-
tose, especially at the base : heads few, cor}'mbose, hemispherical,
about I cm. high and broad : bracts obovate with yellowish or
purplish scarious margins : corollas yellow ; tube and throat each
nearly 2 mm. long ; the latter campanulate and longer than the
lobes : achenes densely silky, scales of the pappus fully 2 mm.
long, longer than the tube of the corolla.

This species has been confused with H. liitctis Nutt. but is easily
distinguished by the long pappus and also by the longer segments
of the leaves. H. scaposus grows in dry soil up to an altitude of
2200 m. from New Mexico and Arizona to Utah and Nevada.

Arizona: Vicinity of Flagstaff, 1898, D. T. MacDougal, izg
(type).

New Mexico: Santa Fee, 1897, A. A. &■ E. Gcrinide Heller,
3555 > 1847, A. Fendler, 436 ; 1869, E. Palmer, 11.

Utah : Southern Utah, 1 874, C. C. Parry, loj.

Nevada: 1868, ^^ Watson, 612.

4. Hymenopappus arenosus Heller, Bull. Torr. Club, 25 : 200.

1898

This resembles somewhat the preceding, but is taller, more
leafy, and has much shorter pappus. It grows in sandy soil, up
to an altitude of 2200 m., in New Mexico and southern Colorado.

5. Hymenopappus cinereus sp. nov.

A grayish tomentose perennial with a cespitose caudex. Stems
about 2 dm. high, branched, with 2-4 leaves : these bi-pinnately
divided into linear segments 1—2 cm. long and about i mm. wide,
sparingly grayish tomentulose : heads corymbose, 8—10 mm. high,
10-12 mm. broad, hemispherical or somewhat turbinate: bracts
oblong-obovate, with narrow scarious margins : corollas yellow :
tube and throat each about 1.5 mm. long; the latter broadly cam-
panulate, longer than the lobes : achenes loosely villous ; scales of
the pappus variable, in the type fully i mm. long and nearly equal-

G35 RvDBERG : Studies on the Rocky Mountain Flora

ing the corolla-tube, but often shorter, yet never hidden by the
hairs of the achenes.

This species has been confused with H. filifolius and H. fiaves-
ccns. Baker, Earle & Tracy's specimens were determined as the
latter by Professor Greene. The species is, however, much nearer
related to H . filifolius , from which it differs mainly in the lower
habit and the longer scales of the pappus. These characters, to-
gether with the longer lobes of the leaves distinguish it from //.
litteus. It grows on dry hills, at an altitude of i 500-3000 m,

Colorado: Walsenburg, 1900, Rydbcrg & Vreeland, S479
(type); Mesas near Pueblo, 5^77; Cuchara Valley, 5^76"; Du-
rango, 1898, Baker, Earle & Tracy, 1028 ; Garden of the Gods,
near Pikes Peak, 1895, E. A. Bessey.

6. Hymenopappus filifolius Hook. Fl. Bor. Am. i : 317. 1833

This species is common on plains and prairies to an altitude of
2000 m., and ranges from Saskatchewan and Montana to Colorado
and Nebraska.

7. Hymenopappus luteus Nutt. Trans. Am. Phil. Soc. (H.) 7:

374. 1 841

Dr. Gray in his Synoptical Flora referred this species to H.
filifolius, which, however, does not fit Nuttall's description. Many
years before he had named Fendler's specimens H. luteus. These
agree fairly well with Nuttall's description, except as to the pap-
pus, which in them is much more prominent than in any of the
other species. Nuttall states that the scales of the pappus of H.
luteiis are very short and hidden by the hairs of the achenes.

In 1897 Prof. Aven Nelson collected a Hymenopappus on
Green River, which he intended to describe as new, especially as
the type specimen was without pappus. A closer examination re-
vealed, however, that some of the specimens really had some
small scales. These agreed perfectly with Nuttall's description.
Nelson's specimens and my own, collected two years before, also
on Green River, are the only ones that perfectly agree with Nut-
tall's description of H. luteus. His type was collected on tiie
Ham's Fork of the Colorado of the West. What the present
name of Ham's Fork is or its exact location I have been unable

Rydberg : Studies on the Rocky Mountain Flora 636

to find ; but from the route that Nuttall took it is undoubtedly-
some stream of the Green River system.

8. Hymenopappus macroglottis sp. nov.

Shghtly tomentose perennial with a woody caudex. Stems
several, 3-4 dm. high, striate, slightly floccose when young, with
1-3 leaves : leaves mostly basal, about i dm. long, bi-pinnately
divided into linear lobes 8—25 mm. long, 1-2 mm. wide ; stem-
leaves generally simply pinnate or the uppermost simple : heads
few, corymbose, 12 mm. high and 12-15 mrn. broad, hemispher-
ical : bracts oblong, with very narrow yellowish scarious mar-
gins : corolla yellow ; tube about 2 mm. long ; throat deeply
campanulate, 3—4 mm. long, 3—4 times as long as the lobes :
achenes elongated obpyramidal, about 5 mm. long, hirsute: scales
of the pappus about i mm. long.

The long and broad lobes of the leaves and the long throat of
the corolla distinguishes it from other species of the Rocky Moun-
tains. The latter character it has common with only H. higens
Greene from southern California. This has, however, much more
finely dissected leav^es. //. macroglottis grows in arid regions
from Texas to Arizona and north to Colorado or Utah.

Arizona : Oak Creek, 1883, H. H. Rusby (type).

New Mexico: 1851-52, Wright, 12^2.

Texas: Camp 42, 1853, Bigeiozo, yjo.

Colorado or Utah : 1843, Fremont.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 11.

LIFE HISTORY OF SCHIZAEA PUSILLA

By Elizabeth G. Britton and Alexandrina Taylor

NEW YORK
1901

[Reprinted from the Bulletin of the Tokrey Botanicj4L Club. 28 : 1-19. 31 Jan., 1901.]

Life History of Schizaea pusiila

By Elizabeth G. Britton and Alexandrina Taylor
(With Plates i-6)

The material on which these studies were based was collected
at Forked River, New Jersey, on the third of July, 1900. The
plants were abundant, but only half grown, the sporophylls being
only five centimeters high. They were found around the base
of small white cedars {^Chamaecyparis tliyoides) kept moist by
hummocks of SpJiagnum, and surrounded by Lycopodmm Caroli-
nianuin, Juncus pelocarpus, Drosera rohindifolia and Utriadaria
cleistogmna. Young plants were found, ranging from two to ten
millimeters in height, growing in depressions of moist sandy loam, or
even perched upon the roots of sedges and Sisyrincldtim Adanti-
ciini. Several sods were taken with the plants in various stages,
and a large number of young plants were collected and preserved
in alcohol. With a magnification of fifteen diameters, it was dis-
covered at the time of collection that they originated from a fila-
mentous protonema, consisting of a tangled mass of dark green
filaments, spreading around the base of the young circinate leaf,
and that these filaments were persistent, even after some of the
leaves were 10-15 mm. high. Entangled with the filaments, in
such a manner as to render it necessary to clean them with a
camel's-hair brush, there were three species of hepatics {Odonto-
schisma sphagni (Dicks.) Dumort ; Lophozia inflata (Huds.) M. A.
Howe ; and Cephalozia catenulata (Hiibn.) Spruce ; also a slender
fresh-water alga, Rhizoclonium hieroglyphiaim (Ag.) Kiitz.
[Issued 31 January.] 1

2 Britton and Taylor : Life History of Schizaea pusilla

A complete or correct description of 5. pusilla cannot be found
in any manual or monograph. In several the spores are said to be
smooth, and the glandular hairs borne by the leaves are not men-
tioned, though they are known on other species of this genus.

The gametophyte is composed of numerous, erect, branch-
ing, dark green protonemal filaments ; monoicous, bearing from
5-12 archegonia, usually on a slightly thickened and expanded
series of cells in the nature of an archegoniophore (?) or directly on
the filaments ; antheridia more numerous, often on separate
branches and nearer the extremities of the filaments ; radicles sel-
dom borne on the filament but produced from specially modified,
large spherical cells, apparently in symbiotic relation with a fun-
gus. Sporophyte perennial, from a short erect or horizontal
rootstock, 5-10 mm. long, sterile leaf 2-5 cm. long by 0.5 mm.
broad, circinate, bearing small club-shaped hairs, nearly i mm.
long, occurring in three longitudinal rows on the dorsal surface,
alternating with two rows of stomata. Sporophyll 3-13 cm. long,
divided at summit into 14-16 fertile pinnae ; sporangia ovoid, with
a terminal ring : spores reniform, pitted, 76-84 11, maturing in
autumn.

On low wet banks with sphagna or in sandy swamps, in the
shade of larger plants ; known from numerous scattered stations
in the Pine Barrens of New Jersey, in Newfoundland (De La Py-
laie, Waghorne), and in Nova Scotia (E. G. Knight). The station
credited to New York by Prantl from the Berlin Herbarium, is
probably a mistake, though there is no reason why it should not
be found on the sandy plains of Long Island and Rhode Island.

According to Prantl there are nineteen species of Schizaea, of
which five are Polynesian, eight are found in Brazil, and five in the
West Indies ; all are of restricted distribution, and in most species
they are known from few stations. Of Lygodiiun he records twenty-
two species, of which five are Mexican and Central American, four
West Indian and only one from northeastern North America, L.
palmatimi. Of the forty- six species of Omithoptcris and Anemia
known, Brazil has thirty-five, Mexico nine, the West Indies six
and only two extend into the United States, 0. adiantifolia (L.)
Bernh., and 0. Mexicana (Kl.) Underw.

The Schizaeaceae are represented in the Tertiary by several

Britton and Tavlor : Life I Iisiokv of Schizaea pusilla 3

species of Lvgodiion, and by Anemia in the Cretaceous. Thus far
Schizaea is unknown in the fossil state. We may safely conclude,
however, that its maximum development in North America must
have been reached previous to the Glacial period, and that it is in
a degenerate condition and retrograding distribution in the only sur-
viving species, Schizaea pusilla, whose larger and more highly de-
veloped relatives exist now only in the tropics.

Spores

The spores of Schizaea pusilla measure 76—84 a, are nearly
reniform (Fig. i), and have a cuticularized exospore which is al-
veolate (Figs. 2, 3) ; on the concave side there is a ridge extend-
ing nearly two thirds the length of the spore, formed by the exo-
spore having a fissure nearly its whole length (Fig. 4). It is
through this slit in the exospore that the young tube emerges
when the spore germinates.

The development of the gametophyte from the spore to the first
archegonium could not be followed in the laboratory ; but from
the laboratory cultures and the different stages of spore germina-
tion found in the soil brought up from New Jersey a fairly good
idea of the manner and rate of growth may be drawn.

Spores, from the plants collected in July and matured in the
greenhouse, were sown on September 5th ; on the 14th they were
found to be slightly green ; the first signs of germination were
seen on the 27th, when the spores contained some chlorophyl,
and two had started to send out a tube which extended 27 n
beyond the aperture (Fig. 5); chlorophyl was visible in the lengthen-
ing tube on the 8th and rhizoids were also found on that date ;
the first cross-wall was formed on the loth ; on October ist the
second wall was formed making a filament of two cells (Figs. 9, 10).

On August 28th a spore was found in the sod of young plants
which had germinated and formed a small rhizoid, slightly brown,
with a curved apex and contained some chlorophyl, and a filament,
115 n'm length, of two cells (Fig. 19), the cell at base was shorter,
about twice as long as broad, containing chlorophyl with no special
arrangement ; the other cell was nearly four times as long as
broad, the chlorophyl denser at the apex and a newly formed wall
(Fig. 19, a). The density of the chlorophyl increased at the

4 Brixton and Taylor : Life History of Schizaea pusilla

apex. On August 30th the chlorophyl showed a tendency to as-
sume its final arrangement in the filament, being very dense in the
center, radiating toward the walls in rather thick bands and con-
necting with a layer next to the wall through the whole length of
the cell ; on August 31st the filament had lengthened to 126//
and one more cell had been cut off (Fig. 21), no further change
in the rhizoid having taken place. The filament consisted of five
cells by September 2d. On September 4th the filament had in-
creased to six cells, and a partial division of the contents of the
apical cell had taken place (Fig. 22). The tip of the filament
was very much curved and densely packed with chlorophyl ; the
basal cell of the filament had become slightly swollen near its
apical end. The filament measured 146 //.

On September 5th the filament consisted of six cells (Fig. 23) ;
the first walls formed were very nearly as thick as the cross-walls
of the older filaments.

The older filaments generally grow erect, and this tendency
toward an upward growth is plainly shown even as early as the
third cell of the filament ; the rhizoid also showed geotropic curva-
ture. One tube, issuing from the fissure of an exospore, was di-
rected downward ; but soon began a curvature which was continued
until the filament occupied a vertical position ; the rhizoid, first
directed horizontally, soon curved downward. The filaments, for
the most part, did not show the tendency to upward growth until
two or three cells had been formed, but the rhizoid took a down-
ward direction much earlier.

On September 5th another spore was found in the soil consist-
ing of one filament of six cells (Fig. 24) ; at the base of the fila-
ment, at its connection with the spore, there was a cell which had
evidently been the basal cell of another filament. The remaining
filament had given rise to two antheridia, which though not dried
up were empty ; one antheridium arose from a short branch from
the second cell, occupying the terminal cell of the branch ; the
other originated from the terminal cell of the filament.

A spore with a healthy filament of four cells (Fig. 26) had borne
an antheridium in which the mother cells of the antherozoids could
be easily seen.

The attachment of the spore appears to be of long duration, as
antheridia are formed while the filament is still attached.

Britton and Taylor : Life History of Schizaea pusilla 5

A sporangium (Fig. 27) filled with spores was sown at the same
time with the free spores on September 5th, and a great number
of the spores germinated inside the sporangium sending out fila-
ments through a basal break ; when the spores were removed from
the sporangium they were found to be more advanced than those
germinating outside, the rate of growth of the spores in the sporan-
gium in a given time being almost twice that of the other spores.

Branches were given off from the basal cells of the filaments,
sometimes from apical portions of young filaments, and in a few
cases the spore cell was found to divide into three primary cells

(Fig- 17)-

Intermediate stages between the earliest developed filament

from the spore and the much-branched protonema are lacking.

Protoxema

The protonema occurs on the substratum, or on rootstocks of
other plants, as small tufts of a dark green color, growing to a
height of 2 mm. and a breadth of 4 mm.

The protonemal filaments are larger than the protonema of
mosses. A comparison was made with Pogonatutn breidcatile and
MniiDi! punctatiiiii with the following results :

Length Breadth

Cells of Pogonatum brevicaule \ l,c, ^

15 // 15//

96 a 23 //

Cells of MniiDH punctatuni \ '

CqWs oi Schizaea pusilla < 173 n 92//

(134 yf 38/^

It is also seen from the above that the cells of the filaments of
Schizaea pusilla are fairly uniform in dimensions. They are
densely filled with chlorophyl ; starch is present. The cells are
cylindrical, sometimes flattened near the base, in the region of the
archegonia. Some few of the cross-walls were found to be per-
forated.

The protonema is copiously branched, the branches being
generally single from each of the cells of the filament, generally
near the upper end of the cell (Figs. y2, 73). Occasionally three
or four in succession will give rise to two branches from opposite

6 Brixton axd T/.ylor : Life History of Schizaea pusilla

sides, but maintain the same relative position (Fig. 30). The
branches, which give rise to the spherical cells to be described
below, divide in the same manner as the main filaments. The
division of other blanches is very irregular (Figs. 29, 30, 31).
The rhizoids are not usually formed directly from the ordinary
cells, but from specially modified cells (Fig. 38,^); in three in-
stances only were rhizoids found directly on the filaments, and in
one case one cell gave rise to two rhizoids. They arise as lateral
branches, at right angles to the long axis of the filament and tak-
ing the place of branches (Figs. 38, 39). There were two cases
found (Figs. 32, 33) where the cells of a branch, near the apex,
had formed partition walls. In Fig. 32 the third cell from the
apex had divided up into four cells, showing a tendency to form a
flat prothallus. Three cells showed signs of division (Fig. 33) :
these two instances were the only ones found. Bower speaks of
flattened expansions on the filamentous protonema of Tricliomanes
alatiini and Tvichonianes sinuosum as described by Mettenius.
These are much more rudimentary in Scliizaea pusilla.

Some cells of the filament have been found to undergo division
in the later stages, into a number of disk-shaped cells which do
not increase in the axial diameter. Constrictions sometimes follow
such divisions at the older cross walls ; the cell walls were a light
brown and showed signs of decay. Fig. 35 shows the cells of the
filament undergoing the same process, but these were as healthy
as the rest of the filament and densely filled with chlorophyl.
Bower refers (Ann. Bot. i : pi. i. f. c? ) to a similar develop-
ment in Trichoinancs pyxidifcnnn ?i\\^s^.ys that " * * * possibly the
moniliform development is merely a pathological condition ; its
appearance, however, is suggestive of that segmentation of the
protonema into spherical cells which is recorded as a mode of vege-
tative propagation for the protonema of Fun aria hygrouietrica."

After some of the filaments have formed several cells the apical
cell cuts off a new cell, which, after the first partition wall, that
is transverse septum, divides longitudinally, forming two cells
(Figs. 36, 37). These cells become large and round, each cell
containing chlorophyl, and giving rise to one, or generally two,
rhizoids (Fig. 37, a). The rhizoids also contain chlorophyl and
early take on a dark yellow color. The original cell of the fila-

Britto.v and Taylor : Like History or Sciiizaea pusilla 7

ment from which these cells arise either continues its errowth
normally (Fig. 36), or by a lateral innovation (Fig. 37). This new
filament, after the formation of two or three cells, may form spher-
ical cells at its apex (Fig. 39), or continue for some time before
doing so, or it may send off a branch at once, which in its turn
forms spherical cells. Generally a filament forming the spherical
cells once does so at intervals throughout its whole length (Figs.
72, 73). These branches early bend to the substratum. If ex-
amined at this stage they will be found to have lost their former
contents and to be filled with fungal hyphae (Fig. 43). This fungus
does not injure its host, but sets up a symbiotic connection by
which it functions as an absorbing organ to supply the gameto-
phyte. So the .spherical cells are undoubtedly formed by the
gametophyte for the reception of the fungus, which enters as soon
as the rhizoids touch the substratum (Fig. 41). While above
ground and filled with chlorophyl they do not show any evidence
of the presence of a fungus. The rhizoids wither early and ab-
sorption is carried on almost entirely by the fungal hyphae. The
lack of rhizoids on the filament is thus explained by the presence
of this fungus symbiont. The rhizoids formed from these spherical
bodies appear to be the only channels through which the fungus
enters the chambers built for it.

As to the nature of the fungus it is at present impossible to
give it a permanent place in any of the series because of lack of
evidence in regard to its method of reproduction. Perithecia
have been found with asci and also what was probably a conidial
stage ; fruit bodies of other forms have also been found among the
filaments ; however, none of these were connected with the fungal
hyphae under discussion. The young filament shortly after ger-
mination was in some few cases attacked by a fungus (Fig. 17, d),
and this fungus is found wrapped around many of the cells of the
older filaments, several instances having been found where haus-
toria had penetrated into the cells. Some of the plants are so in-
fected by fungi after the growth of the sporophyte has begun that
all the filaments have thickened walls and are pierced by three
or four haustoria in each cell ; they have turned brown and lost
their contents. This fungus not only clings to the gametophyte
of Schizaca pusilla but attacks the sporophyte also, though it does

8 Britton and Taylor : Life History of Schizaea pusilla

not appear on the sporophyte until the first leaf has reached a
height of I mm. It resembles the one referred to by Bower in his
work on Trichomanes. No connection has yet been made between
these hyphae and those of the fungus symbiont. As far as is
known at present, the fungus which wraps itself around the pro-
tonemal filaments is the same or at least bears a very close resem-
blance to that which attacks the sporophyte both on the rhizome
and leaf.

The hyphae of the symbiotic fungus penetrates the rhizoid
generally a short distance back from the tip (Fig. 41, <r). These hy-
phae sometimes branch in the rhizoid and their cross-walls are
more numerous, and in many cases the hyphal threads appeared
narrower. They enter the large spherical cells where they form
bladders or granulated swellings in these cells, sometimes nearly
filling them (Figs. 42, 43). The fungus sometimes penetrates the
ordinary cells of the filament, entirely changing the shape of these
cells. Pale brown bodies were found of irregular outline attached
to the hyphae which were apparently sporangia.

Antheridia

The antheridia are produced laterally on the protonema, occu-
pying the terminal cell of the lateral branch or more rarely the
terminal cell of a filament, which continues its growth laterally
or ends with the formation of the antheridium. They are found
either singly or in groups ; and may be considered as metamor-
phosed branches as in Fig. 44. They may either be formed from
the terminal cell of a short lateral branch (Fig. 44) or from the
terminal cell of a branch giv^en off from the basal cell (Figs. 44,
48) or from the second or any other cell of a lateral branch (Fig.
44) or rarely they may be formed on the terminal cell of a short
lateral branch given off from the same cell of the filament as a
previous antheridial branch (Fig. 46). Occasionally these groups
are borne on a branch of the filament which bears only antheridia
(Fig. 48) and for the most part in groups, sometimes from both
sides of a branch. These branches which bear the antheridia,
whether it is a branch of two cells with the apical one becoming
an antheridium, or one of the groups, occupies the same relative
position on the filament as do the branches of the main filament.

Brixton and Taylor : Life History of Schizaea pusilla 9

One or two were found to start out from the center of a cell of a
filament. The antheridia are produced in great quantities but a
great number of them are aborted, and the majority of antheridia
terminate a branch of two cells. They occur nearer the apex of the
filaments than the archegonia, and sometimes on the same filament
with the archegonia (Fig. 57). They also occur on a branch from
the filament which gives rise to the archegonia ; when they are
borne on a separate filament they are generally formed in larger
quantities. The cells of a filament which give rise to the anthe-
ridial clusters are often broader and sometimes shorter than the
ordinary cells.

The antheridium is formed by a cell of a filament sending out a
cell which divides by a partition wall near the apex (Figs. 49, 50).
This apical cell enlarges and soon cuts off another cell by a wall
parallel with the first (Fig. 51). This small cell does not elongate
but always remains short and forms the pedestal for the antherid-
ium ; the apical cell becomes large and globular and cuts off a
cap cell at the summit, with the wall oblique (Fig. 52) ; the large
cell divides up into the mother cells of the antherozoids (Figs.
52, 53, 54) and one ring cell. In some cases there appear to be a
single layer of two or three peripheral cells. The ring cell (or
cells) contain chlorophyl though they lose thfs before the anther-
ozoids are matured. Dehiscence takes place by the swelling of the
ring cell and the rupture of the cap cell. The antherozoids appear
to be surrounded by a A'ery fine membrane when they escape from
the antheridium (Fig. 55); they are spirally coiled, with cilia at
their anterior ends. Very few ripe antheridia were found. The
antherozoids do not seem to be produced in large quantities.

Archegonia
The archegonia occur nearer the base of the filaments than the
antheridia (Fig. 57) on cells of the filaments which have become
more than one cell wide through division. They generally bear
the same relation to the original cell of the filament as do the
branches from other cells ; they are borne singly or in pairs (Fig.
63), sometimes in groups of three or four (Fig. 58) often on both
sides of the protonema. One filament was found which gave rise
on six consecutive cells to two archegonia each ; two cells above

10 Brittox and Taylor : Life History of Schizaea pusilla

the sixth cell there was another archegonium formed, and a cell
above this another one, and branches also gave rise to archegonia.
There is no filament that is specially reserved for the formation of
archegonia as is sometimes the case with the antheridia. Each
archegonium is derived from a single superficial cell.

The archegonia are formed by the division of the initial cell
into three cells ; the basal cell forms the venter which may or may
not be imbedded in the cell of the filament. Some of the division
cells of the original cell of the filament grow up around it in
such a way as to make it appear as if imbedded (Fig. 65). From
the neck cell arises the neck of the archegonium, consisting of four
rows of cells, of four cells each (Figs. 60, 62, 63) ; a uniformity
which produces a straight neck to the archegonium.

Occasionally the cells of the two rows on the posterior side,
though they do not increase in number, become larger than those
on the anterior side (Fig. 62) thus slightly bending the neck
toward the anterior side. From the middle cell of the superficial
mother cell arises the central cell and the canal cell ; the middle
cell becomes sharply pointed on the upper end and forces itself
between the neck cells ; this point is cut off, forming the canal cell ;
the larger cell divides again into two cells of unequal size : the
smaller and upper one forms the ventral canal cell, the lower and
larger one forms the egg cell (Fig. 59). When this is mature the
canal cells dissolve into mucilage. When the archegonium opens,
the four stigmatic cells, which in this species are very large, are not
thrown off but fold back (Fig. 63). It is at this stage that the
curve in the neck oqcurs in some archegonia due possibly to the
fact that as the filaments grow erect or nearly so, the archegonia
occupy the portion of the filaments below the antheridia, and by
bending the neck they bring the canal to the oosphere in a more
direct line for the capture of the antherozoids, an adaptation tend-
ing to secure fertilization. Generally several archegonia are pres-
ent, but only one seems to give rise to a sporophyte.

The cushion of cells on which the archegonia are borne can
hardly be called an archegoniophore as some of these cells give
rise to vegetative branches (Fig. 65). Three cases were found
where an archegonium arose directly from a cell of the filament
without any partition other than that of the formation of the ar-

Brixton and Taylor : Life History of Schizaea pusilla 11

chegonium. Bower (/. c, Figs, ii, 12) says in his description of
the archegonia of Trichomancs pyxiciifertnn,"T\\Q diYch&gornd. are
borne on massive growths (archegoniophores). * * * jp one
case, however which has a special interest the mass of tissue on
u/hich the archegonia are produced is obviously the result of par-
tition of a single cell of a filament, without any marked increase
i 1 size having taken place (Fig. 13). This may be regarded as the
simplest form of an archegoniophore hitherto described in any
fern or even in any Bryophyte and it approaches near to that sug-
gested by Goebel as the simplest possible, in which the sexual or-
gans would be inserted directly on the protonemal threads."

The act of fertilization was not seen, nor the direction of the
first wall, but, from later stages they appear to be formed as is
common with most ferns ; a wall is formed parallel with the long
axis of the archegonium, then a cross-wall is formed.

Foot
The foot is in most cases extremely large, and is in every case
a well-defined organ, consisting of a great mass of cells which for
the most part contain chlorophyl (Figs. 69, 70, 71). It remains
attached to the protonema for some time, having been found in
connection with it after the formation of the third leaf (Fig. 75,
a), and even here it appeared to be in a healthy condition as did
also that part of the protonema on which the foot was borne. It
grows down into the cushion of the gametophyte (some of the
cushion cells appear to grow up around it). The venter cells
grow and form a calyptra around the embryo covering it for some
little time ; remnants of it were found still clinging around the first
root after the formation of three leaves (Fig. 75, b).

Root

The primary root is a prolongation of the main axis of the
sporophyte (Figs. 71, 72, 73), while the ultimate roots are always
adventitious and produced in acropetal succession. They arise
from all sides of the erect rhizome (Fig. 79), the epidermis of which
sends out rhizoids. The primary root is persistent and becomes
quite long. The second and third roots have a vestigial struc-
ture which covers them as the coleorhiza of some endogens

12 Hrittox AND Tavlok : Life History of Schizaea pusilla

(Fig. 76). The roots are fully formed and have root caps (Fig.
75, d). This root cap (Fig. 76) consists of four large pear-shaped
cells, inflated on one side ; the inflated side is away from the root,
the concave side rests on the root tip. They are developed before
the root sheath splits. The cells are replaced from the tip and, as
the older ones do not always fall off when the new ones have been
formed, there have been seen as many as five series (Fig. yj),
though they show their age by the partial discoloration of their
walls.

The epidermal cells are large and thin-walled ; the outer walls
often bend into the cavity of the cell and frequently break. The
cortex consists of two layers ; the cells of the inner layer are very
large and have the walls that lie next to the endodermis thickened ;
but in no instance was the thickening found to be as great as that
fio-ured for ScJiizaea Pennida. There is an endodermis of two
layers, and the central cylinder (Fig. 78) is like that described by
Prantl (Untersuch. Morph. Gefasskrypt., />/. jf) for Schizaea Pennida.

Sclerosis takes place in all the layers without any marked in-
crease in the thickness of the walls.

Rhizome

The rhizome is erect (Fig. 79), occasionally creeping. It always
forms a protective covering of trichomes over the growing end
(Fig. 74, d^\ these trichomes consist of from two to five cells
(Fig. 75, Ji), measuring i mm. in length which soon turn brown
and are persistent. The internodes are of varying lengths. One
rootstock (8 mm.) had borne nineteen sterile and two sporophylls,
all dead except the five sterile leaves last formed. The fertile
leaves measured 6.5 cm., the longest sterile leaf 4. 5 cm. Another
rhizome of the same length had twenty-two sterile leaves, six
green and two nearly brown, with two fertile ; these were 7 cm.
high, the base of the fertile leaves was green, the sporangia brown
and mature. There were twenty -two roots — six short and young ;
one root was 25 mm. long and had branched; the branches were
5 mm. long.

A cross-section near the young tip shows a central bundle with
a well developed endodermis (Fig. 80). Sclerosis takes place in
the entire cortex ; the different stages are beautifully shown in

Britton and Taylor : Life History of Schizaea pusilla 13

young sections ; these cells, including the epidermis, are filled with
starch. The epidermis and cortex are often invaded by fungal
hyphae.

The vascular bundle is concentric ; the xylem portion is en-
veloped in the phloem. The central cylinder is surrounded by a
well-defined endodermis and phloem sheath, the radial walls of
which are thin and fragile ; the phloem elements are represented
by two or three imperfect rows of narrow parenchymatous cells
and sieve tubes ; the xylem consists of scalariform tracheids with
occasionally small spiral tracheids close to the phloem. The
medullary parenchyma is composed of large, thick-walled cells,
pitted, and early showing sclerosis, but not as early as the funda-
mental tissue outside the bundle.

Sterile Leaves

The sterile leaves are linear, slender and tortuous. The de-
velopment of the leaf is very slow, the lower portions having long
been fully formed while the apex is still unfolding. The vernation
is circinate (Fig. 79). Owing to the more rapid growth of the cells
on the dorsal side than those on the ventral, the leaf is rolled up
on the ventral side. When fully developed they bear on their
dorsal side two rows of stomata alternating with three rows of
glands (Fig. 81), sometimes four or more rows of glands. The
glands seem to originate from special cells cut off from the epi-
dermis ; these epidermal cells frequently do not lengthen, keeping
very nearly an isodiametric shape ; when they do lengthen the
glands remain at or near the upper wall.

The young leaves and the tip of the stem are more or less
completely clothed with trichomes early turning brown. These
are not to be confounded with the glandular hairs. They are
composed of two or more cells and are extremely long, measur-
ing in some instances i mm. to 3 mm. or perhaps more. The longest
glands of the leaf measured nearly 100 ij., others 76 /y. and in
width 3 I p.. Some are composed of one cell, others of two cells ;
they are all club-shaped and contain granular protoplasm. These
glands were rarely found on the ventral surface, and sometimes
they did not appear to follow any law as to their formation on the
dorsal surface, though, for the most part, they were formed in al-
ternate rows with the stomata.

14 Britton and Taylor : Life History of Schizaea pusilla

The stomata are restricted to two rows of epidermal cells and
almost every epidermal cell in these rows gives rise to one (Fig.
82). A cell of the epidermis before it has lengthened forms a U-
shaped wall at the upper end of the cell (Figs. 86, a, b) ; the points
of the U meet the radial wall which separates this cell from the
one next above (Fig. 86, b) ; this cell becomes the mother cell of
the stoma, and by growth presses the partition wall back a short
distance into the upper epidermal cell (Fig. 86, c). This cell
divides by a tangential wall into two cells of equal size ; these
become the guard cells (Fig. 86, c, d) each containing abundant
chlorophyl. These guard cells enlarge considerably (Fig. 86,^, e)
so that they are raised above the epidermis as shown in an oblique
view (Fig. 83). The wall between the guard cells splits along its
central portion making an opening to the space below ; the epi-
dermal cell meanwhile has lengthened and the cell above has
formed a stoma in the same way. The leaf bundle is more nearly
collateral than that of the stem ; the xylem faces the ventral, the
phloem the dorsal surface of the leaf (Figs. 88, 90). There is a
two-rowed endodermis around the bundle ; the epidermal cells are
large and in some instances occupy one half of the cross section.
The ground tissue is made up of thin-walled parenchyma with
numerous air spaces (Fig. 89) and the cells contain chlorophyl.

Sporophyll

The sporophyll is very similar to the sterile leaf with the excep-
tion of the formation at its apex of pinnae bearing the sporangia ;
these have been carefully studied by Prantl and others in several
species of Schizaea, the descriptions of which, from present obser-
vations, appear to hold good for Schizaea pusilla. One of the
largest sporophylls measured 1 3 cm. from base to apex, the por-
tion bearing the pinnae was 6 mm. long and eight pinnae were
formed on one side and seven on the other ; the longest pinna
measured 4 mm., of this 2^ mm. is the portion which bore the
sporangia. The lowest pinna on each side had formed four spo-
rangia each, the others eight each. Prantl figures six sporangia
for Schizaea dichotoma, and sixteen for Schizaea Penmda. The
edge of each pinna rolls up over the sporangia, forming an in-
dusium, and the end cells at the summit and also along the mar-

Britton and Taylor : Life History of Schizaea pusilla 15

gin produce trichomes which also cover the sporangia. These
trichomes are often composed of more than one cell, and resemble
the trichomes formed by the rhizome and are in some instances as
long as 134 /i, 345 /i, 461 ix, the width being 38 jx, and 30 }x.

A surface view of the dorsal side of the sporophyll shows two
rows of stomata alternating in some cases with rows of glands,
though these sometimes are not in rows, and occasionally only
two glands were found. These glands are smaller than those
generally found on the sterile leaf measuring only 38 fi.

The stomata appear sunk below the epidermis, but a cross-section
showed them to be the same as in the sterile leaf. The two rows
of stomata continue up the leaf from the base to the pinnae, where
they are lost in a great number of stomata which cover the dorsal
surface of the pinnae with no special arrangement. No glands
were found on any of the pinnae examined.

The warts or swellings from the epidermal cells are far more
numerous in the sporophyll, though they are found on the sterile
leaf (Fig. 93, a). They do not appear to follow any law as to their
arrangement on either leaf, though they appeared to be more
numerous on the ventral side. The epidermal walls are thicker
than the epidermal walls of the sterile leaf, some of which had ex-
tremely thin walls and a rudimentary bundle (Fig. 90).

The bundle has a well-marked endodermis ; it appeared from the
cross-section to be collateral as did all the bundles with the excep-
tion of that of the rhizome (Fig. 94). The elements of the bundle
were not traced out, but reticulated and ring tracheids were
found in the xylem. The mesophyll tissue of the sporophyll (Fig.
89) is composed of thin-walled cells with numerous air spaces :
these cells seem to be branched in a stellate manner in both sterile
leaves and sporophylls.

Summary

The spores are small, are nearly reniform, and have a cuticu-
larized exospore which is alveolate. There is a ridge along the
concave side having a fissure nearly its whole length through
which the young tube emerges when the spore germinates.
Out of a great number sown at one time only two had germi-
nated by the end of the third week, the others taking a longer

16 Brittox and Taylor : Life History of Schizaea pusilla

time. The spore remains attached for some time after the forma-
tion of antheridia.

The gametophyte is a filamentous protonema, irregularly
branched, bearing both antheridia and archegonia on the same
filaments ; and producing rhizoids from specially modified cells
which are inhabited by a symbiotic fimgus.

The antheridia occur singly, or in groups on special branches
bearing antheridia alone. They are produced in great numbers
though but few ripen. They are simple in their structure and
the first wall formed in the antheridium is parallel with the wall
cutting it off from the lateral branch, forming a pedicel. A small
number of antherozoids are produced in an antheridium, which
are enclosed in a membrane when they escape from the antheridium.
The archegonia arise at, or near, the base of the filaments,
either directly on the filament or, more often, on cushions formed
by the division of the cell of the filament. They are character-
ized by the uniformity of the neck rows and the large size of the
stigmatic cells.

The foot is a large, well-defined organ, remaining attached to
the protonema for some time after the formation of the third frond,
carrying nourishment from the gametophyte to the embryo which
is far advanced before it breaks through the calyptra.

The primary root is persistent. The second and third roots
have a vestigial sheath through which they do not break until
after the development of the root-cap. The root-cap consists of
four large pear-shaped cells inflated on the side away from the
root tip.

The rhizome is erect, always forming a protective covering
over the growing end ; the trichomes are large, turn brown early,
and are persistent.

There is a central concentric bundle with a well-marked endo-
dermis. Sclerosis takes place in the entire cortex, the cells of
which, with the epidermis, are filled with starch. The epidermis
and cortex are often invaded by a fungus hypha.

The sterile and fertile leaves have two rows of large stomata,
on the dorsal side, alternating with two or more rows of glands ;
these glands are small and sometimes wanting on the fertile leaf.
The young leaves are more or less completely clothed with

Britton and Tavi.ok : Liik Historv or Schizaka i'L'sili.a 17

trichomes. Warts or swellings occur from the epidermal cells on
both surfaces, though more numerous on the ventral side.

The bundles appear collateral with a well-marked endodermis.
The mesophyll tissue is composed of thin-walled cells, branched

in a stellate manner.

Explaiiailoii of I'lates

Plates I, 2, 3 and 4 were drawn from a niagnilkation three limes as great as ex-
pressed in the numbers which represent the magnification of the figures as they stand

in the reproduction.

Pi.ATi: I

1. Different views uf the spore, X 80.

2. Spore, X 140-

3. Portion of exospore, X 333h-

4. Ridge and fissure in exospore seen from above, X 195-

5-17. Different stages in the germination of the spore, X S^K- ".filament;
/', rhizoid ; f, c, new branch.

18. Germinating spore of Botrychium obliqunni, two weeks and five days.X h'^Vi-
Sown at same time as Schizaea spores. The last-named did not start to germinate un-
til after three weeks.

19. Germinating spore found in soil on August 28, X ^"- "> Indication of cross-
wall.

20. The same on August 29. a, cross-wall formed.

21. The same on August 31, X jO-

22. On September 4, X 3°-

23. On September 5, X 3°-

24. Spore found in soil on September 5, X 3°- ". antheridia ; b, swelling at base
of terminal antheridium.

25. The same with wall formed at a.

26. Filamentof four cells with antheridium showing mother-cells of antherozoids

(rt). h, Rhizoid, X 80.

27. Sporangium with spores germinating inside, X 3°- Owing to position in

which the sporangium fell when sown— the filaments from the spores are not sent out

though the regular fissure at a.

Plate 2

28. 29, 30 and 31. Methods of branching of the protonemal filaments.
32 and I';,. Cells of the filament dividing.

34 and 35. Cells of the filament becoming moniliform.

36. Voung spherical cells with the longitudinal wall («) just forming, X 80.

37. Older stage of the same showing young rhizoids {a, a) and young branch
starting from filament at base of spherical cells /', X 80.

38. Portion of filament showing spherical cells, antheridia, and archegonia. a,
spherical cells ; /', rhizoids ; c, fungus in spherical cells ; d, antheridia ; e, archegonium,

X30.

39. Shows position of spherical cells, X 3°-

40. Abnormal condition. One cell of the filament giving rise to one spherical
cell, and a cell of the filament next above giving rise to two, X 3°-

41. End of rhizoid showing fungus penetrating into the cell. Shaded portions
are hyphae which are inside the rhizoid, X 195-

18 Bkitton and Taylor : Life History of Schizaea pusilla

42. Upper part of same showing portion of spherical cell with the bladder-like
hyphae (i). A hypha in rhizoid, >; 195-

43. Spherical cells filled with fungus a, X I40-

Pl.ATIC 3

44. 45, 46 and 47. Different ways of branching of anlheridial filament.

48. One of the antheridial branches with antheridia in different stages of comple-
tion, a shows one filament giving rise to three, X jO-

49, 50, 51, 52, 53, 54, 55 and 56. Stages in formation of the antheridium. Fig.
49, X 8o> first cell sent out from main filament. Fig. 50, X 30, later stage showing
first wall cut off near tip. Fig. 51, X^o, older stage. Terminal cell a becomes the
antheridium ; cell d the pedestal. Fig. 52, >< 140, <?, cap cell ; d, mother cells of the
antherozoids. Fig. 53, X 140, later stage. Fig. 54, X 140, ripe antheridium before
splitting cap-cell. Fig. 55, X SSS'i) antherozoids still in membrane. P'ig. 56, empty
antheridium.

57. Branch showing antheridia and archegonia. All the antheridia but «~are
aborted, X 162^.

58. Young archegonia, X ^4'^-

59. Young archegonium, X 30. «, canal cell ; /', ventral canal cell ; c, egg cell.

60. Older stage of same before opening, X ^O-

61. Looking down on the four stigmatic cells of the archegonium, X ^^^

62. Archegonium opening, X ^o.

63. Showing large stigmatic cells (a) folding back, X 80.

64. Cell of the filament dividing up before the formation of the archegonia, X ^4^-

65. The same with an archegonium, X 30-

Plate 4

66 and 67. Upper and under view of egg cell after fertilization enclosed in the
calyptra, X 3°-

68. Young embryo, X 3°-

69. a, garaetophyte with archegonium. i>, foot; c, leaf; a', stem of sporophyte,
X30.

70. Young sparophyte. a, foot; />, leaf; c, stem; </, trichomes ; e, gland on
rond ; /, calyptra, X 80.

71. Young sporophyte. a, foot ; /', root ; c, frond ; d, calyptra.

72 and 73. Two stages in the growth of the sporophyte showing curled tip of
frond. Marking the same in both, o, gametophyte ; />, sporophyte ; r, calyptra,

X 12^.

74. a, Rhizome ; />, root ; c, first leaf ; d, trichomes.

Plates 5, and 6 were drawn from a magnification twice that expressed in the num-
bers which represent the magnification of the figures as they stand in the reproduction.

PL.VfE 5

75. Sporophyte still attached to gametophyte after the formation of the third leaf.
a, foot ; 6, portion of calyptra ; c, root ; d, young root, the dotted lines indicate root-cap
which can be seen through the vestigeral covering ; /, leaf; g, rhizome; A, trichomes
which cover tip of young leaf and rhizome ; some have been removed to show glands
on leaf ; /, gametophyte, X 45-

76. Young root just emerging from its covering, X ^7/4-

Brittun AM) Tavi-or : Life Historv of Schizaea pusilla 19

77. Root-cap ; here shown in five series.

78. Cross-section of root. </, thickened inner walls of cells of the ground tissue
next to endodermis (A) ; c, phloem ; </, xyleni, X 292^2.

79. Showing rhizome with four leaves and five roots ; <r, rhizome ; /', roots ; c, tri-
chomes (the internode here represented is unusually long and distinct), > 8-

80. Cross-section of rhizome not far from tip, X 120. ir, cells tilled with starch ;
d, endodermis ; r, phloem ; ,/, xylem ; e, fungus hyphae entering epidermal cells.

Plate 6

81. Surface view t)f dorsal side of sterile leaf, showing the two rows of stoniata
with the glands alternating with them, X 120.

82. Portion of epidermis of sterile leaf with three stomata, a, b, c. a, shows the
chlorophyl grains ; in /' the contents have been removed to show the original cross-wall
((/) between the epidermal cells, and the way the guard cells {e) rest on epidermis at

f, X 210.

83. Oblique view of sterile frond showing raised stomata (<;).

84. Longitudinal section of same. </, epidermal cells ; /', one guard cell of stoma ;
c, air cavity.

85. Cross-section of same, a, guard cells ; b, air cavity, X 210.

86. Five diagrams showing development of stoma. I b, cross-wall between epi-
dermal cells ; 2 /', mother cell of stoma with U-shaped wall ; c, formation of longi-
tudinal wall through mother cell dividing it into the two guard cells ; </, shows curve in
original cross-wall, and the splitting of the longitudinal wall. The dotted line indi-
cates the relative size of the guard cells which have started to swell. In c, the stoma is
complete ; i c, original cross-wall ; 2 c, guard cells.

87. Diagram of cross-section of sterile leaf showing the two rows of stomata (t).
a, bundle ; b, endodermis, X 45-

88. Bundle from sterile leaf, a, endodermis; c, phloem; d, xylem, X 210.

89. Mesophyll tissue from sterile leaf, X 210.

90. Cross-section of a young sterile leaf with a rudimentary bundle, marking as
in Fig. 88, X 210.

91. Diagram of cross-section of fertile leaf, marking as in Fig. 87, X 45-

92. Stoma seen in cross-section of sporophyll, X 210. «, stoma ; b^ air space.

93. Two epidermal cells from cross-section of sporophyll showing warts («), X
210.

94. Cross-section of bundle in sporophyll, marking as in Figs. 88, 90, X 210.

Bull. Torr. Bot. Club, 2S.

PL. I.

THE HELIOTYPE PRINTING CO.. BOSTON.

Bill-. ToRR. BoT. Club, 28.

Pl. 2.

Bull. Torr. Bot. Club, 28.

Pl. 3.

Bii.L. ToRK. lUn . CiA'i), iS.

Pi.. 4.

Bull. Tokr. Bot. Club, 28.

PL. 5.

/

Bull. Torr. Bot. Club, jS.

Pl. 6.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 12.

STUDIES ON THE ROCKY MOUNTAIN

FLORA-IV

BY p. A. RYDBERG

NEW YOEK
1901

[Reprinted from the Bulletin of the Toerey Botanical Club. 2S:. 20-S8 31 Jan., 1901.J

Studies on the Rocky Mountain Flora.— IV

]'>V p. A. RvDliKKC

Arnica tomentulosa sp. nov.

A leafy perennial with slender horizontal rootstock. Stem
villous, about 4 dm. high : leaves oblanceolate, obtuse, sessile or
the lower with short-winged petioles, denticulate, finely villous-
pubescent, almost tomentulose, with two pairs of stronger veins,
the larger 13 dm. long, somewhat )'ellowish : heads few, hemi-
spherical ; disk about i 5 mm. high and 2 cm. broad : bracts ovate
or ovate-oblong, obtuse, villous, 12-16 in number: rays light
yellow, over i cm. long, 4 mm. wide.

This species is nearest to A. inollis, but differs in the broad
obtuse involucral bracts. It grows at an altitude of about 2700 m.

t>

Wyoming: Buffalo Fork, 1S97, P. Tivccdy, 52^.

Arnica tenuis sp. nov.

A low slender perennial with horizontal rootstock. Stem spar-
ingly villous pubescent, about 2 dm. high, monocephalous : leaves
usually 3 pairs, sparingly villous when young, entire ; the lower
two pairs oblanceolate or spatulate, the upper lanceolate : involucre
turbinate-campanulatc, villous, about 13 mm. high, 10-12 mm.
broad: bracts linear, 1-1.5 mm. wide, green; rays orange, 16-
18 mm. long and 4 mm. wide.

This species resembles A. gracilis in the size of the plant and
form of the head, but the leaves are much narrower and the heads
solitary. It is intermediate between that species and A. fulgciis.
It grows at an elevation of about 2200 m.

Wyoming : Big-Horn Mountains, 1899, F. Tiveedy, 2og^.

Artemisia diversifolia sp. nov.

A white tomentose perennial with horizontal rootstock. Stems
.simple, leafy, white-tomentose, 5- 10 dm. high : leaves densely to-
mentose on both sides, subsessile, 5-10 cm. long : the lower pin-
nately cleft into 3-5 narrowly lanceolate acuminate lobes, which are
directed forward : the upper entire, linear-lanceolate, passing into
the bracts of the inflorescence : inflorescence a narrow panicle,
1.5-3 <^n^- long: heads numerous, conglomerate, sessile, 3-4 mm.
high and 3 mm. broad : bracts oblong or ovate-oblong, scarious-

20

21 RvDBERG : Studies on the Rocky Mountain Flora

margined, densely villous-tomentose : flowers heterogamous, but
all fertile, light yellow : receptacle glabrous.

This species belongs to the A. Ludovici(xna group, and in many
respects agrees with the original description thereof. It is, how-
ever, a western species, not growing near the region from which A.
Ludoviciaiia was described. What the latter really is I have been
unable to settle. The one that I think is the only one that has
any claim for the name, is a lower plant from Missouri to Colorado,
with shorter leaves, more green above, with more divergent lobes
and brownish flowers. A. diver sifolia grows on sandy beaches up
to an altitude of 2200 m.

Idaho: Priest River, 1900, D. T. MacDougal, igo (type);
Farmington Landing, Lake Coeur d'Alene, 1892, Saiidberg, Mac-
Dougal & Heller, jog.

Washington: 1889, G. R. Vasey, ^yg.

Wyoming: Yellowstone Lake, 1899, Aveu & Elias Nelson,

6603.

Picradenia helenioides sp. nov.

A comparatively tall, finely pubescent plant with apparently
only biennial root. Stem leafy, about 5 dm. high, with several to
many erect branches : leaves rather firm, distinctly ribbed, finely
pubescent ; the lower petioled and with half clasping bases :
basal leaves entire, very narrowly linear-oblanceolate ; middle
stem-leaves erect, fully i dm. long, parted into 3-5 linear divi-
sions : upper stem-leaves linear, entire : heads corymbose : invo-
lucre somewhat tomentose, 8-10 mm. high and often 15 mm.
broad ; outer bracts united only at the base, lanceolate, longer
than the inner, 14-18 in number: rays orange, about i cm. long,
2-3 mm. wide, 3-toothed at the apex: achenes silky: scales of
the pappus broadly lanceolate, acuminate.

It is nearest related to P. biehins (A. Gray) Greene ; but differs
in the yellowish green herbage, the erect branches, the broader
segments of the leaves and the darker flowers. It grows in moun-
tain valleys at an altitude of about 2700 m.

Colorado: On Sangre de Christo Creek, 1900, Rydberg &-
Vreeland, 54.^5.

Antennaria Piperi sp. nov.

Somewhat surculose-rosuliferous : basal leaves 2-4 cm. long,
obovate or oval with a short petiole, densely floccose on the lower
surface, only slightly so on the upper surface when young, but

Rydberg : Studies ox the Rockv Mountain- Flora 22

soon <;labi-atc, i -ribbed or indistinctly 3-ribbed, <,^enerally distinctly
mucronate : flowering stems 1-1.5 dm. high, somewhat glanduli-
ferous above ; its leaves small and bractlike, oblanceolate or linear ;
heads 5-7 in a short raceme : fertile heads 7-8 mm. high and
6-7 mm. in diameter ; its bracts imbricate in about 4 series,
slightly floccose, green at the base, purplish in the middle and
with a light brownish scarious margin above ; the outer short and
ovate ; the inner lanceolate, acute : pappus very slender, filiform,
dirty white ; sterile heads about 5 mm. high and about as broad ;
its bracts broadly oblong, more floccose, brown and with a broader
scarious margin of the same color as in the fertile head, obtuse or
truncate ; pappus only slightly broadened above, white.

It is nearest related to A. raccmosa, but differs in the more co-
pious and more persistent tomentum, in the broader and brighter
colored bracts of the fertile head and the broad scarious margins
of those of the sterile one.

Oregon: Olympic Mountains. 1895, C. V. /'//^v (fertile plant
in herbarium of Washington E.xperiment Station, Pullman) ; moun-
tains of northern Oregon, Wilkes Expedition (sterile plant in the
Torrey herbarium).

Aster mollis sp. nov.

A strict perennial with a horizontal rootstock. Stem densely
villous, almost tomentose, or in age more glabrate, 4-8 dm. high,
simple below, with short flowering branches above ; leaves obovate
to oblanceolate, 5- 10 cm. long, densely and softly grayish pu-
bescent on both sides, sessile, sHghtly clasping, and occasionally a
little auricled at the base : heads terminating the short branches,
hemispherical ; disk about 1 5 mm. high and broad ; bracts
herbaceous, oblong, obtuse, villous-pubescent ; rays numerous,
bluish purple, about i cm. long and 2 mm. wide.

This species reaches an elevation of 2200 m.

Wyoming: Big-Horn Mountains, 1899,/^. Tweedy, 202g {\.y]^€).

Washington: Pullman, 1893, C. V. Piper, 1604.

Townsendia Vreelandii sp. nov.

A dark green biennial, branching near the base. Stems and
branches simple, erect, 1.5-4 dm- l^igh. mostly erect, sparingly
villous. Leaves numerous, all oblanceolate, short-petioled,
glabrous, mucronate, 4-8 cm. long, 5-8 mm. wide : involucre
about 15 mm. high and 25-30 mm. broad: bracts lanceolate,
acuminate, scarious, light green with dark green center : rays

23 RvDBERG : Studies on the Rocky Mountain Flora

bluish purple, 12-1 5 mm. long, 2 mm. wide : achenes thin, oblong,
truncate : pappus of 2 subulate bristles, with a few intermediate
squamellae.

It is nearest related to T. eximia, but has larger heads, is more
leafy, and even the upper leaves are oblanceolate. It grows at
an altitude of 2500-3000 m.

Colorado : Side ot Veta Mountain, 1900, F. K. Vreeland,
6jg (type); Veta Pass, 1900, Rydberg & Vrcc/aud, J404 and
S405 ; West Spanish Peak, j^oO.

Erigeron leucotrichus sp. nov.

Perennial with a somewhat branching rootstock. Stems about
2 dm. high, sparingly villous below, more copiously so above :
basal leaves oblanceolate orspatulate, 3-5 cm. long, short-petioled,
bright green and sparingly hairy : stem-leaves oblanceolate to
linear, 1-2 cm. long ; head solitary ; involucre white-villous with
multiceptate hairs ; bracts very numerous, lanceolate, almost
black, acuminate with spreading tips : rays very numerous, about
6 mm. long and i mm. wide, light purplish pink or white.

This species is nearest related to E. mdanocephalus, but easily
distinguished from it by the white, not black, hairs of the involucre.
It differs from E. simplex in the taller habit, the larger heads and
the darker bracts of the involucre. It grows at an altitude of
about 2500 m.

Wyoming: Big-Horn Mountains, July, 1899, Frank Tzveedy,

200J.

Erigeron incanescens sp. nov.

(?) Erigeron glabellas var. mollis A. Gray, Proc. Acad. Phil.
1863 : 64, in part.

A densely and finely cinereous pubescent plant with perennial
rootstock. Stem simple, 1.5-3 ^^- ^^'gh- mostly ascending, striate
densely pubescent, leafy : basal leaves oblanceolate or spatulate :
stem-leaves about 5 cm. long and i cm. wide, oblong, oblanceo-
late or the upper lanceolate, sessile and somewhat clasping : heads
1-4; disk about 8 mm. high and 15 mm. broad: bracts very nu-
merous in one series, narrowly linear, cinereous as the rest of the
plant ; rays pale blue or violet, very numerous, about i 5 mm. long
and less than i mm. wide.

From Dr. Gray's description, this species must have been in-
cluded in E. glabellas wdiY. mollis; but all specimens that I have

RvDBERG : Studies on the Rocky Mountain Flora 24

seen under that name belong to E. subtrino-vis Rydb., which is its
nearest relative. From this it differs in the dense grayish pubes-
cence, the narrower stem-leaves, which are not 3-ribbed, and the
lower habit. E. iiicancsccns grows in rocky places at an altitude
of about 3000 m.

Colorado : West Spanish Peak, i goo, Rydberg & Vree/and,

5415-

Erigeron viscidus sp. nov.

A low cespitose perennial. Stems ascending, a little over i
dm. high, more or less hirsute and glandular-pubcrulent, especially
above ; leaves 3-5 cm. long, oblanceolate, dark green, sparingly
hirsute and ciliate on the entire margins : heads few : disk about i
cm. high and i 5 mm. broad : bracts very numerous, subequal in
one series, narrowly linear, fuscous, acuminate, glandular-pubcru-
lent, but not hirsute : rays very numerous, pale blue, about 8 mm.
long and .5 mm. wide.

This species is intermediate between E. glandnlosus and E.
macrantliiis. It has the habit of the former, although larger, and
the large heads and numerous narrow rays of the latter. It grows
in wet ground, at an altitude of about 2700 m.

Colorado: Near the Gray-Back Mining Camps, 1900, Ryd-
berg & Vreeland, 541 6.

Valeriana acutiloba sp. nov.

A bright green plant with horizontal or ascending rootstock
and polygamo-dioecious flowers. Fertile plant 4-5 dm. high :
basal leaves entire with a short wing-margined petiole, 5-7 cm.
long ; blade spatulate or obovate, acute : stem-leaves usually 3
pairs, pinnately divided ; lateral divisions lanceolate to Hnear, long-
acuminate ; the terminal one large, oblanceolate or of the upper-
most very small leaves linear-lanceolate, entire or saliently toothed :
cyme dense, contracted, 2-5 cm. long and about as wide ; gland-
ular-puberulent : bracts linear subulate, about i cm. long : flowers
perfect; corolla funnelform, about 4.5 mm. long; tube proper
about I mm.; fruit broadly ovate, about 4 mm. long : pappus
about 7 mm. : staminate plant lower, 3-4 dm. high, with more
sterile shoots, which have much longer leaves : stem-leaves usually
only 2 pairs, less divided, with only 1-2 pairs of lateral divisions :
cymes denser, flowers all or nearly all staminate : their corollas
larger, 5-6 mm. long, more oblique ; the limbs about 5 mm. wide.

This species has the dense cyme of V. capita/a ; but the plant
is taller, the corolla shorter and is easily distinguished from all its

25 Rydberg : Studies on the Rocky Mountain Flora

American relatives by the narrow, long-acuminate divisions of its
stem-leaves. It grows in wet places, especially under snowdrifts,
at an altitude of 2500-3300 m.

Colorado : Near Gray- Back Mining Camp, Sangre de Christo
Range, 1900, Rydberg & Jlreland, ssy6 (type, fertile plant);
Mountain near Veta Pass, JJ/i (staminate plants) ; Bear Creek
Cafion, 1895, Ernst A. Bessey {^^xW\€)\ Pagosa Peak, 1899, C. F.
Baker, 620.

Campanula MacDougalii sp. nov.

A slender glabrous perennial, about 3 dm. high with slender
horizontal rootstock. Leaves very thin : the basal ones and lower
stem-leaves with slender petioles 2-4 cm. long : blades broadly
ovate, 2-3 cm. long, coarsely sinuate-dentate : upper stem-leaves
lanceolate to linear, entire ; the largest 5-6 cm. long : sepals linear
subulate, 10-12 mm. long, at last reflexed : corolla 13-15 mm.
long, nearly of the same shape as that of C. roinndifolia : style
exserted, straio-ht : fruit not seen.

The species is nearest related to C. Scoulcri ; but is easily dis-
tinguished from that species by the lack of the sharp toothing of
the leaves. The lower leaves resemble somewhat those of C.
rohindifolia, and were it not for the exserted style and the smaller
corolla it may be referred to the var. Alaskana of that species.

Idaho : Priest Lake, 1900, D. T. MacDoiigal, 66.

Castilleja linearis sp. nov.

A rather slender perennial, with a rootstock. Stem simple,
3-4 dm. high, finely villous pubescent : leaves narrowly linear,
about 5 cm. long and 2 mm. wide, pubescent, entire or the upper
with a pair of narrowly linear lobes : bracts broader, deeply cleft
into three linear lobes, tipped with brick-red : calyx villous, 2-2.5
cm. long, cleft almost equally deeply in front and behind, more than
halfway down : lateral lobes linear, 8-10 mm. long: corolla 2.5-
3 cm. long, greenish yellow : galea about i 2 mm. long, with red mar-
gins : lower lip dark green, about 3 mm. long, 3-lobed.

This species resembles in habit most the subarctic C. pallida,
but has the corolla of C. iiiincata. It grows in meadows at an alti-
tude of about 2500 m.

Colorado: Near West Spanish Peak, 1900, Rydberg & Vree-
land, j6i^.

Rydberg : Studies on the Rocky Mountain Flora 26

Castilleja trinervis sp. nov.

A tall perennial with rootstock. Stem more or less villous,
especially the upper portion, solitary, simple or branched, 3-6 dm.
high : leaves dark green, finely puberulent, entire, acute, usually
3-ribbed ; the lower linear, 5-8 cm. long and about .5 cm. wide ;
the upper lanceolate and often i 5 mm. wide : lower bracts green,
similar but shorter and broader and usually 3-5-lobed : the upper
with almost crimson tips : calyx densely white villous, about 3 cm.
long, equally cleft in front and behind, each division laterally cleft
into two oblong lobes, 5-8 mm. long, tipped with the same color
as the bracts : corolla about 4 cm. long, slightly curved, green, but
the galea with almost crimson margins : galea about i 5 mm. long :
lower lip 4-5 mm. long, dark green, with three narrow lobes : style
about 5 mm. longer than the corolla.

This species is nearest related to C. rhexifolia and C. confitsa,
but is characterized by the copious white villous pubescence of
the upper part of the stem and the calyx. It grows in open woods
at an altitude of 2700-3000 m.

Colorado: Headwarters of Sangre de Christo Creek, 1900,
Rydberg & Vreelaiid, ^620 (type); Gray-Back Mining Camps, 3621.

Castilleja luteovirens sp. nov.

A simple perennial with rootstock, often turning black in dry-
ing. Stem 3-4 dm. high, leafy, slightly pubescent when young,
soon glabrate, except the upper portion which is slightly villous :
leaves lanceolate to almost linear, 3-4 cm. long, 3-9 mm. wide,
finely puberulent, 3-ribbed, acute, entire, or rarely the upper 3-
lobed : lower bracts ovate, obtuse, entire, tipped with light green-
ish yellow or greenish white ; the upper ones 3-toothed at the
apex and greenish yellow throughout: calyx villous, 15-17 mm.
long, almost equally cleft in front and behind, laterally cleft about
3 mm.; lobes lanceolate : corolla 22-24 mm. long, greenish ; galea
6-"/ mm. long with yellow margins ; lower lip 2.5-3 "^"i- long,
bluntly 3-lobed.

This species has been referred to C. scptcjitrionalis and C. pallida,
but differs from both in the broader, less acuminate leaves and
broader, more entire yellowish bracts. Neither of the two species
mentioned is found in the southern Rockies. Mostly all the
material that has been determined as either belongs to the present
species. This grows in meadows at an altitude of 1200-2700 m.

Colorado: Sangre de Christo Creek, 1900, Rydberg & Vree-

27 RvDBERG : Studies on the Rocrv Mountain Flora

land, j6i6 (type) and 5(5/;; Wahatoya Creek, 3618 ; Middle Park,
1 861, 6^. C. Parry, 241; Chicken Creek, \d>g^, Baker, Earlc &
Tracy, JJ4 : near Denver, 1869, B. H. Smith; Ford of Chama,
1859, Neiuberry, in Macomb's Expedition ; Seven Lakes, Pikes
Peak, 1894, Ernst A. Bcssey.

Wyoming: Big-Horn Mountains, 1899, F. Tiuccdy, 2J40;
Laramie Plain, 1884, C. S. Sheldon, 80.

Utah : Salt Lake City, 1879, M. E. Jones, loji.

Mimulus gratioloides sp. nov.

A low, branched annual of more or less reddish color, gener-
ally less than i dm. high, somewhat viscid puberulent especially
above. Leaves oblong-lanceolate, sessile, about i cm. long, sinu-
ate-dentate : pedicels slender, in fruit i 5-20 mm. long ; calyx 7-
8 mm. long, cylindraceous in fruit : lobes subequal, short, broadly
ovate, acute, ciliate on the margins ; corolla yellow, about i cm.
long and 3 mm. broad, only slightly bilabiate : throat beardless.

This species is nearest related to M. riibellus and the specimens
from Colorado referred to that species may belong here. M.
gratioloides differs, however, in the smaller yellow corolla and the
acute calyx-lobes. It grows in exposed places among rocks and
gravel at an altitude of about 2300 m.

Colorado : Butte, 5 miles southwest of La Veta, 1900, Ryd-
berg &" Vr eel and, j66o.

Pedicularis lunata sp. nov.

A perennial, perfectly glabrous up to the inflorescense, with a
rather stout, but snnple rootstock : stem about 4 dm. high, slightly
striate and purplish : leaves alternate, dark green, 5-12 cm. long,
pinnately divided to near the midrib ; segments linear or linear-
oblong, crenate : spike 15-20 cm. long, rather lax; bracts pecti-
nately divided with prolonged endlobes, the lower often equal-
ing the flowers in length, slightly arachnoid villous : calyx about
9 mm. long ; its teeth broadly lanceolate, about 3 mm. long : cor-
olla pinkish, over 2 cm. long ; its tube about twice as long as the
calyx, strongly curved ; galea strongly arcuate, produced into a
rather long beak and almost crescent-shaped ; lower lip almost
meeting the beak of the upper, very broad, indistinctly 3-lobed
with large rounded lateral lobes.

The very broad lower lip, and long-beaked galea suggest
somewhat P. contorta and P. ctenophora, but the corolla-tube is

Rydberg : Studies on the Rocky Mountain Flora 28

much longer and the beak not spirally twisted. The species may
therefore be placed with P. Caiihyi, P. Parryi and P. Hallii, but the
corolla and beak are much more arcuate. It is a handsome spe-
cies growing at an altitude of nearly 2800 m.

Wyoming : Big-Horn Mountains, 1899, F. Tweedy, 2J17 (type).

Pentstemon erosus sp. nov.

A glabrous cespitose perennial, 2-4 dm. high. Basal leaves
oblanceolate, short-petioled, 3-5 cm. long, entire, mostly acute :
stem-leaves opposite, sessile, oblong or lanceolate, entire, acute or
the uppermost acuminate : flowers in a dense interrupted spikelike
inflorescence : calyx-lobes broadly ovate, almost cuspidate-acumi-
nate, tinged with dark purple and white and with an erose-dentate
margin : corolla purple with very dark limb : lower lip broad with
3 rounded reflexed lobes, bearded on the inside : upper lip with 2
erect narrower lobes : sterile stamens narrowly clavate, short-
bearded.

This species is nearest related to P. procerus, but easily dis-
tinguished by the erose sepals and more reflexed lower lip. It
grows at an altitude of 2000-2700 m.

Colorado: Indian Creek Pass, 1900, F. K. Vreeland, 615
(type); South Park, 1^7 1, John [F^^/^r (Wheeler Expedition), 2pj ;
Chicken Creek, West La Plata Mountains, 1898, Baker, Earle &
Tracy, 6j8.

Pentstemon Wilcoxii sp. nov.

An almost glabrous perennial with a cespitose caudex. Stems
3-6 dm. high, glabrous or slightly puberulent above, simple :
leaves opposite, glabrous and somewhat glaucous, dentate with
small sharp callous teeth, acute : the basal ones petioled, lanceo-
late : the lower stem-leaves sessile, oblanceolate or oblong, 3-5 cm.
long ; the upper lanceolate and slight!)' clasping : inflorescence
an elongated interrupted thyrse : branches 1-2 cm. long, fastigiate-
cymose : calyx glabrous, about 4 mm. long, cleft }^ its length :
lobes lanceolate, acute, slightly margined below and there often
sinuately toothed : corolla straight, purple, glabrous on the out-
side, about 1 5 mm. long ; the lower lip longer than the upper,
slightly bearded within : sterile stamen club-shaped, with a short
dense brown beard.

This species is probably nearest related to P. Jiuinilis ; but
easily distinguished by the tall stems, the sharper toothed leaves,
and the numerous purple flowers.

29 Rydberg : Studies ox the Rocky Mountain Flora

Montana: Kalispell, 1900, E. V. IVl/cox, jyo {type in United
States National Herbarium) and j68.

Polemonium delicatum sp. nov.

A small glandular perennial with slender horizontal rootstock.
Stem very slender, about i dm. high : leaves 3-8 cm. long ; leaf-
lets 5-1 1 pairs, very thin, ovate, ovate-lanceolate or oblong,
mostly acute, 3—10 mm. long: inflorescence usually branched;
branches 3— 4-flovvered ; pedicels slender, 5—15 mm. long; calyx
glandular, 4-5 mm. long: lobes lanceolate, acute: corolla blue,
open-campanulate, about 7 mm. high and 8 mm. broad : stamens
about equalling the corolla.

This is nearest related to P. parviflornm Nutt., but is still
smaller and characterized by its slender pedicels, small flowers and
acute calyx-lobes. It grows at an altitude of over 3000 m.

Colorado: West Spanish Peak, 1900, Rydberg &■ Vreeland,
5720 (type).

Colorado or New Mexico : Canadian ?, Dr. James.

Polemonium speciosum sp. nov.

A low viscid-villous plant, cespitose with a perennial rootstock.
Stem 1-2 dm. high with 2-3 leaves: basal leaves numerous, 8-15
cm. long with wing-margined rachis ; leaflets opposite, 7—9 pairs,
ovate, 6—12 mm. long, acute: stem-leaves similar but smaller : in-
florescence capitate : calyx viscid-villous, 15-20 mm. long, cleft to
about the middle : lobes lanceolate, acute : corolla pale blue, deeply
campanulate-funnelform. 2-2.5 cm. long: limb about 1.5 cm.
broad.

The form of the corolla, the dense capitate inflorescence and
the viscid pubescence place this species nearest P. viscosnm, and
P. covifertnin, but the leaflets are not verticillate, the corolla is
larger than that of the former and broader than in the latter.

Colorado: Mount Garfield, 1900, Fred. Clements.

Gilia Candida sp. nov.

A glandular perennial with short caudex. Stems often more
than one, ascending or erect, simple, 4—6 dm. high, leafy : leaves
rather fleshy, divided into linear-filiform, spinulose-tipped seg-
ments ; the upper gradually smaller : inflorescence a very narrow
panicle : branches usually very short and few-flowered : flowers
subsessile : calyx glandular-pubescent, about 8 mm. long ; its lobes
lanceolate, bristle-tipped : corolla white, or rarely pale pink, 2—3

Rydberg : Studies on the Rocky Mountain Flora 30

cm. lone, salverform : its lobes oval or obovate, obtuse : stamens
inserted unequally below the throat, included.

This species is nearest related to G. aggrcgata ; but differs in
the white corolla and its rounded obtuse lobes. It is also lower
and the upper leaves more reduced. It grows on dry hillsides at
an altitude of 2000-2700 m.

Colorado: Mesas near La Veta. 1900. F. K. Vrceland, 602
(type); Veta Pass, 1900, Rydberg & Vrceland, 3730; Calham,
1893, Dc A. Saunders.

Phacelia alba sp. nov.

A viscid-villous annual or biennial, 2-4 dm. high. Stem leafy,
strict and simple viscid-villous and glandular above : leaves twice
interruptedly pinnatifid, about i dm. long, glandular-puberulent,
hispid ciliolate on the margins and veins ; ultimate segments ovate
or oblong, 3-6 mm. long : inflorescence branched, dense, in flowers
almost capitate, but branches in fruit spiciform : flowers nearly
sessile, 2- ranked : calyx glandular, cleft to near the base ; sepals
broadly linear, obtuse, about 2 mm. long, about one third shorter
than the white corolla : the lobes of the latter rounded, crenate :
appendages fo, broadly ovate : stamens and style much exserted :
capsule ovoid, about 6 mm. long : seeds often solitary in each
cavity, finely pitted.

This species is nearest related to P. Nco-Mexicaiia and P. Popci,
resembling the latter most in habit, but having the viscid-pubes-
cence and the crenate corolla-lobes of the former. It differs from
both in the small white corolla and the long- exserted stamens. It
grows in mountain valleys at an altitude of 1800-3000 m.

Colorado: Sangre de Christo Creek, 1900, Rydberg & Vree-
land, 3755 (type); Valley of Upper Arkansas River, \^7l,Jolui

Wolfe (Wheeler Expedition), 99 ; Headwaters of Clear Creek,

1 86 1, C. C. Parry, 514.

New Mexico: Ruidoso Creek, White Mountains, 1895, E. O.

Wootoii.

Lappula calycosa sp. nov.

A hirsute annual, simple below, branched above with long
virgate branches. Stem hirsute, 3-4 dm. high : leaves oblong,
obtuse, 3-4 cm. long, 7-8 mm. wide ; those of the branches
smaller : pedicels short, in fruit 2-3 mm. long, generally 4 mm.
below the leaves ; corolla pale blue, about 1.5 mm. long, and i
mm. broad : calyx-lobes enlarged in fi it. foliaceous, 4-6 mm.

31 Rydberg : Studies on the Rockv Mountain Flora

long and reflexed : fruit about 4 mm. in diameter : nutlets sur-
rounded by a single row of subulate glochinate prickles, some-
what flattened at the base but not united into a wing-border : back
of the nutlets strongly muricate.

It is nearest related to L. occidentalis (Wats.) Greene, but dif-
fers in the oblong obtuse leaves, the enlarged and reflexed fruiting
calyx-lobes, and the virgate branches. It grows in deserted fields
at an altitude of about 2000 m.

Colorado: Walsenburg, igoo, Rydberg & Vreeland, fijif;.

Lappula cupulata (A. Gray)

Echinospcrinujii Redozi'skii var. cupulata A. Gray, Bot. Calif, i :
530. 1876.

I think that this should be regarded as distinct from L. Tcxana
(Scheeie) Britton, as the habit is quite different, L. cupulata is
diffuse, branching at the base, with elongated branches, while L.
Tcxana is simple at the base, and branched above with short
branches.

L. Tcxana is a southern plant, ranging from Texas to New
Mexico. The range of L. cupulata is from Nevada to Nebraska,
south to Colorado.

Cryptanthe minima sp. nov.

A dwarf hirsute annual, beginning to bloom when only i cm.
high. Stems several, erect, in the specimens seen only 1-4 cm.
high, long hirsute : leaves spatulate or oblanceolate obtuse, hirsute
on both sides, 5—1 5 mm. long and 2—3 mm. wide : flowers crowded,
2.5—3 iTim- long, subsessile : lobes of the calyx linear, hispid:
corolla white ; limb a little over i mm. broad ; nutlets whitish,
about I mm. long, dissimilar ; 3 ovoid, strongly muricate ; the
fourth somewhat larger, smooth ; the inside edge with a slender
groove, triangular-dilated at the base.

In size and general habit, this species is strikingly like C.
pusilla ; but the fruit is different ; the nutlets being dissimilar, one
of them smooth, and all with rounded lateral angles and more
rounded backs. The specimens of our collection are rather young
and only two full)' developed fruits were found. C. niinwia was
found on dry hillsides at an altitude of about 2200 m.

Colorado : Cucharas River, above La Veta, 1900, Rydberg &
Vre eland, 56 gy.

Ryduerg : Studies ox thk R(»ckv Mountain Flora 32

Mertensia lineariloba sp. nov.

A slender perennial with ijlabrous simple stem, 2—3 dm. high,
basal leaves 5-15 cm. long, long-petioled ; blades oblanceolate,
obtuse; stem-leaves subsessile, linear or linear-lanceolate, acute,
3-6 cm. long, 4-9 mm. wide, strigose above, glabrous beneath :
flower clusters 3-4-flowered in the axils of the upper leaves ;
pedicels very slender, 2-6 mm. long, strigose : calyx divided to
near the base ; lobes linear, acute, 3-4 mm. long, almost equaling
the tube of the corolla, glabrous on the back, but hispid ciliatc on
the margins : corolla blue, 7-8 mm. long, tube nearly of the same
length as the throat and limb ; the latter 4-5 mm. long : stamens
almost equaling the corolla ; filaments dilated, and broader than
the anthers.

It is nearest related to JI. Iwcaris, but characterized by the
narrow, strongly ciliate calj-x-lobes and the filiform pedicels. It
grows in shaded situations at an altitude of 2500-2700 m.

Colorado: West Indian Creek, Trichera Range, 1900, Ryd-
berg &- Vreelatid, j;6gi (type) ; near Empire, 1885, //. N. Patterson,

115-

Mertensia ovata sp. nov.

A low cespitose, somewhat fleshy perennial. Stems 1-1.5
dm. high, glabrous: leaves 2-5 cm. long, 1-1.5 cm. wide, mi-
nutely strigose above, glabrous beneath ; the lower obovate and
short-petioled ; the upper broadly ovate and sessile : flower-cluster
dense ; pedicels very short : calyx cleft to near the base : sepals
lanceolate, ciliate on the margin, about 4 mm. long and one third
shorter than the corolla-tube: corolla 10-12 mm. Ions:; its tube
nearly one half longer than the throat and limb : the latter about
7 mm. broad : stamens nearly equaling the corolla ; filaments
dilated and broader than the anthers.

This species is probably nearest related to M. lanceolata ; but
differs in the stunted habit and the broad leaves. It grows among
rocks, at an altitude of 2800-3500 m. •

Colorado: West Spanish Peak, igoo, RvdbtTg & Vreeland^
j6po (type) and jSpoa.

Mertensia obtusiloba sp. nov.

A low cespitose perennial. Stems ascending, glabrous, 1—2
dm. high : lower leaves spatulate or oblanceolate, obtuse, tapering
into a winged petiole, dark green, glabrous beneath, minutely
strigulose above, glabrate in age, 3-5 cm. long ; the upper broadly
lanceolate, sessile : flower-clusters several from the axils of the

33 Rydberg : Studies on the Rocky Mountain Flora

upper leaves : pedicels very short, often sparingly strigulose : calyx
divided to near the base ; sepals oblong, obtuse, ciliate on the
margins, 2-3 mm. long, scarcely more than half as long as the
tube of the corolla : corolla dark blue, about 7 mm. long : tube
nearly equaling the throat and limb : stamens short, included in
the tube, filaments very short, not broader than the anthers.

This species is nearest related to M. Tiucedyi, but differs in
the shorter, obtuse calyx-lobes, the broader and thinner leaves
and the stem which is not depressed or prostrate. It grows at an
altitude of 2000-3500 m.

Colorado: Pikes Peak, 1900, Fred. Clenioits (type); Garden
of the Gods and Pikes Peak, 1894, Ei'-nst A. Bessey ; Argentine
Pass, 1878, Marcus E. Joiics, j^.

Mertensia membranacea sp. nov.

A tall erect perennial with a rather thick tap-root. Stem gla-
brous or sparingly hirsute above, 6-8 dm. high : leaves all petioled,
or the upper sessile ; blades ovate, acute or more often short acu-
minate, 4-8 cm. long, 1.5-4 cm. wide, very thin, hispid-stringulose
on both sides : flower-clusters terminal and in the axils of the upper
leaves, branched and many-flowered : pedicels 5-10 mm. long, his-
pidulous: calyx-lobes 2-3 mm. long, hispidulous, lanceolate, acute,
one-third or one-fourth as long as the tube of the corolla : corolla
about I cm. long ; the pale blue or almost white tube longer than
the dark blue limb and throat : limb about 4 mm. broad : stamens
much shorter than the limb ; filaments dilated and broader than
the anthers : nutlets strongly rugose and spotted.

This species is related to M. panicidata. Mr. Bessey and my-
self collected it in 1897, but as the specimens were rather poor,
they were referred doubtfully to that species. As more and bet-
ter specimens have now been received, it has been possible to draw
a description. It differs from M. paniadata, in the thinner leaves
which almost always show an acumination, in the short calyx-lobes
which scarcely enlarge in fruit and the smaller and numerous
flowers. It grows in moist places at an altitude of 300-2000 m.

Idaho : Priest River, 1900, D. T. MacDougal, j (type) ; Cedar
Mountain, 1892, Sandberg, MacDougal & Heller, ^20.

Montana: Electric Peak, \Sgj, Rydberg & Bessey, ^864.

Mentzelia ctenophora sp. nov.

A diffuse cespitose scabrous perennial. Stems 3-6 dm. long,
branched, in age straw-colored : lower leaves linear or linear-Ian-

RvDHERG : Studies o\ the Rocky Mountain Flora 34

ceolate in outline, tapering at both ends, often 2 dm. long : the
upper lanceolate with a broad base : all pectinately laciniate with
linear-oblong divergent obtuse lobes ; hypanthium cylindrical,
15-18 mm. long; sepals ovate-lanceolate, acute, 4-5 mm. long:
petals obovate, light yellow, about 8 mm. long : seeds irregularly
angled, but not winged, finely muricate.

This species is perhaps nearest related to M. albicaulis, but
differs in the large size of the plant, the diffuse habit, and the larger
flowers. It was collected on railroad banks and in loose barren
soil on hillsides, at an altitude of 1 800-2 lOD m.

Colorado : On Cucharas River, below La Veta, 1900, Rydbcrg
& Vrecland, sydg (type) ; near Walsenburg, S7^^-

Impatiens aurella sp. nov.

A slender glabrous annual, about 6 dm. high. Petioles 1.5-3
cm. long : leaf-blades ovate or oval, thin, bright green, a little paler
beneath, 4-5 cm. long, coarsely toothed, acute ; teeth and apex
finely mucronate : peduncles ascending, mostly 2-flowered : bracts
minute, linear, about 2 mm. long : flowers orange, not mottled :
sack conical, about i cm. long and 6 mm. broad at the base ; its
' spur recurved, about 8 mm. long.

This is nearest related to /. biflora, but differs in the much

smaller flowers, which are scarcely more than half the size of that

species and without any spots, the comparatively longer spur and

less pale leaves. It grows in swamps at an altitude of about 600 m.

Idaho : Priest River, 1900, D. T. MacDougal, 20.

Geranium nervosum sp. nov.

A tall perennial with thick root and short caudex. Stems 4-8
dm. high, minutely retrorsely strigose. in age more glabrate : basal
leaves with petioles 2-4 dm. long ; blades reniform in outline. 8-
12 cm. in diameter, finely strigose on both sides, divided to near
the base into 5-7 obovate-cuneate or oblanceolate-cuneate divisions
which are again 2-3 -cleft and coarsely toothed : stem-leaves none,
except those subtending the inflorescence ; these sessile, 3-5 -cleft ;
branches of the cymes and calyx vei-y densely glandular pubes-
cent : sepals oval, 8-9 mm. long, terminated by a filiform tip, i-
2 mm. long: petals broadly obovate, 15-18 mm. long, pale violet
or almost white, with dark purple streaks : carpels as well as their
column densely glandular ; the latter nearly 2 cm. long : style be-
yond the column about 5 mm. : seeds glabrous, minutely reticulate.

This species is somewhat intermediate between G. viscosissiunnn

35 RvDBERG : Studies on the Rocky Mountain Flora

and G. RicJiardsoiiii, having the general habit, the leaves and pu-
bescence of the stem of the latter, but the densely glandular pu-
bescence of the inflorescence and the calyx and carpels of the
former. The color of the flower is most like G. RicJiardsonii, but
scarcely pure white and with much more prominent veins. It
grows at an altitude of 1800-2700 m.

Wyoming : Pish Creek, Teton Forest Reserve, 1897, F. Tiveedy,

494- (type).

Colorado: Continental Divide, Routt County, 1894, C. S.

CfandalL

Lupinus candicans sp. no v.

A low cespitose perennial, densely white-silky throughout.
Stems ascending, 1.5-2 dm. high, 3-4-leaved, often branched:
stipules linear subulate, 5-8 mm. long : petiole 3-8 cm. long : leaf-
lets about 7, densely white-silky and shining, 1.5-2.5 cm. long,
oblanceolate, acute, mucronate, mostly conduplicate : raceme rather
dense, 3-8 cm. long on a peduncle 2-5 cm. long : bracts small
and early deciduous : calyx densely silky-villous, only slightly
saccate on the upper side ; lower lobe about 4 mm. long : banner
dark blue with a light brown spot, about 7 mm. long, very broad,
and with the sides strongly reflexed ; wings dark blue, as well as
the banner glabrous, about 9 mm. long and about equaling the
keel : the latter whitish, tipped with dark purple : pod densely
white-silky, 3— 4-seeded.

This species has the white pubescence of L. scricens and L.
Hcllerae ; but is in every way a much smaller and more cespitose
plant : the flowers are much smaller and of a darker and more in-
tense blue.

Montana : (locality not given), 1900, E. V. Wilcox, ^j i (type

in U. S. Nat. Herb.) ; Boulder, i2j and I2g, in part ; Big Timber,

j<?5 ; Highwood Mountains, 42; Columbia Falls, 1897, R. S.

Williajiis.

Lupinus cyaneus sp. nov.

A stout and tall perennial with rather simple caudex. Stem
4-9 dm. high, densely villous, but not white, very leafy and in age
somewhat branched : stipules subulate, over i cm. long: petioles
5-10 cm. long : leaflets 7-1 1, oblanceolate, 3-9 cm. long, glabrous
above, almost velvety beneath, acute : raceme long and dense.
1.5-2 dm. long: bracts rather persistent, often over i cm. long:
flowers very numerous, 2-4 m each verticil, very short-pediceled :
calyx white-velvety, somewhat saccate above : lower lobe about

Rydberg : Studies on ihe Rocky Mountain Flora 36

8 mm. long : banner slightly hairy on the back, light blue, with
a light brownish spot at the center, about 8 mm. long : wings light
blue, about i cm. long, equaling the keel, which is whitish with
purple tip: pod densely silky-villous, about 25 mm. long and 8
mm. wide, 4-5 -seeded.

This species has the general habit and the long dense raceme
of L. IcucopJiyllus, but is much greener and the flowers are much
smaller and lighter in color. No. 435, cited below, is referred
here doubtfully. It is more silvery and more branched and had
apparently almost white flowers.

Montana: (locality not given), 1900, E. \\ Wilcox, ^46 (type
in U. S. Nat. Herb.) ; Coal Spur, 435 (?) and 7^9 (the latter in
fruit); Gallatin Valley, 1896,/ H. Flodman, 617.

Astragalus sulphurescens sp. nov.

A light green cespitose perennial. Stems ascending, about 4
dm. high, angled, glabrous, somewhat branched : stipules ovate to
lanceolate, membranaceous, free from the petioles : leaflets 13-19,
elliptic, obtuse or acutish, mucronulate, 12-30 mm. long, with a
few scattered strigose hairs : spike dense and elongated : flowers
ascending : bracts lanceolate-subulate, about 8 mm. long : calyx
white-strigose with scattered black hairs : tube about 5 mm. long :
lobes almost filiform, fully 5 mm. long : corolla light yellow :
banner narrow, 15-18 mm. long, much exceeding the wings and
keel : wings very narrow, only about 1.5 mm. wide : pod 2-celled,
with dorsal suture deeply inflexed, deeply obcordate in section,
about I cm. long, 3-4 mm. wide, strigose, with black hairs.

This species is closely related to A. adsiirgens, but differs in the
yellow, ascending flowers, narrow petals, more scant pubescence,
the long slender calyx lobes and the black hairy pod. It grows
at an altitude of 1600-3000 m.

Colorado: Georgetown, 1895, P. A. Rydberg (type); along
Platte River, Denver, 1878, M. E. Jones, 8ji.

Aragallus villosus sp. nov.

Densely and intricately cespitose perennial. Leaves basal,
numerous, 5-10 cm. long : leaflets 25-31, rather crowded but not
verticillate, 1-1.5 cm. long, lanceolate, very acute, densely but
somewhat loosely silky : scape about 1.5 dm. long, loosely silky,
almost hirsute, with spreading hairs : spike dense, 4-5 cm. long :
bracts linear or narrowly linear-lanceolate, 8-IO mm. long : calyx

37 Rydberg : Studies on the Rocky Mountain Flora

white-silky villous ; lobes very short, about 2 mm. long : corolla
ochroleucous without any purple, 12-15 r""^. long: keel very
short and round, abruptly tipped with a small tip : pod thin, per-
fectly 2-celled, white-silky, about 1.5 cm. long, more than twice as
long as the calyx, tipped with a slender beak which is abruptly
hooked at the apex.

This belongs to the A. campcstris group and is characterized
by the coarse spreading pubescence of the scape and the ochro-
leucous flowers without any trace of purple.

Montana : Craig, 1900, E. V. Wilcox, jy8 (type in U. S. Nat.

Herb.).

Trifolium lilacinum sp. nov.

A densely cespitose dwarf subscapose perennial with very deep
root. The short branches of the caudex covered with the scarious
stipules and remains of old leaves : leaves 3-foliolate, bright green ;
petioles 2-6 cm. long, strigose : leaflets elliptic or lanceolate-ob-
long, acute at each end, 1-2.5 cm. long, strigose : peduncle 5-12
cm. long, bracts minute, less than i mm. long, 3-toothed : flowers
reflexed in fruit : calyx densely strigose ; tube 3 mm. long ; teeth
subulate-filiform, 4-6 mm. long : corolla pale rose-purple or lilac,
in age turning light brownish, about i cm. long ; banner straight
and obtuse.

This species is related to T. dasyp/iyliui/i ; differing in the
minute 3-toothed bracts and their shorter and comparatively
broader leaflets. It is still nearer related to T. acuiniiiatiis Greene ;
from which it is distinguished by the smaller flowers and the not
acuminate banner. It is an alpine plant growing among rocks at
an altitude of about 3000 m.

Colorado: West Spanish Peak, 1900, Rydberg &■ Vreeland,
j-pso (type), J PS I and spS--

Lathyrus leucanthus sp. nov.

A glabrous or slightly pubescent perennial with a very slender
rootstock. Stem about 3 dm. high, angled : stipules very narrow,
semi-sagittate : leaflets 2-4 pairs, elliptic, veiny, glaucous, acute,
mucronate, 1-3 cm. long, 3-10 mm. wide: tendrils of the lower
leaves mere tips ; of the upper elongated and sometimes 3-di-
vided : racemes short, in the axils of the middle leaves, 3-4 cm.
long, 2-4-flowered : calyx glabrous, cleft to about the middle :
lobes lanceolate, acuminate : corolla white, about i 5 mm. long :
banner broad.

RvDBERG : Studies on the Rocky Mountain Flora 38

This is nearest related to L. Arizoniciis, but differs in the
broader lea\'es and banner. It grows at an altitude of 2400—
3000 m.

Colorado : Ojo, 1900, Rydbcrg & Ireelaiid, 6020 (type) ;
Pass Creek, 6021 ; West Indian Creek, 6022; mountain near
Veta Pass, 6o2j; ; Veta Pass, 1890, Mr. and Mrs. G. H. Hicks,
/p ,• Ridgway, 1895, F. Tweedy, 2j;g.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 13.

FURTHER STUDIES ON THE POTENTILLEAE.

BY P. A. RYDBERG.

NEW YORK
1901

[Reprinted from the Bulletin of the Toeeey Botanical Club, 28 : m-183, 27 Mar., 1901.1

Further Studies on the Potentilleae

By p. a. Rydberg

1. WEST AMERICAN SPECIES

Potentilla horrida sp. nov.

A stout perennial with very thick woody root and a short cau-
dex covered with the thick coriacious brown remains of old
leaves: stems ascending, 1.5-4 dm. iiigh, stout, few-leaved, hir-
sute-villous : petioles of the basal leaves 5-10 cm. long, stout,
densely hirsute-villous, broadened at the base and surrounding the
stem : leaves digitate; leaflets usually 5, linear-oblong or oblong-
lanceolate, obtuse, crenulate, 3-5 cm. long, 7-12 mm. wide, densely
pubescent on both sides, almost velvety : pubescence at first white,
later on brown, consisting of tomentum and longer silky hairs: stem-
leaves similar or trifoliolate, much smaller ; the upper subsessile ;
the stipulates lanceolate to linear-subulate, 1-3 cm. long, adnate to
the petioles : cyme rather congested : hypanthium and calyx
densely pilose : bractlets lanceolate, acute, about half as long as
the ovate-lanceolate sepals w^hich are about 5 mm. long : petals
yellow, broadly obovate, about 7 mm. long.

A species oiihe gracilis group, but resembling most P. Hacma-
tochrus in habit and pubescence. It differs, however in the coarser
pubescence, the 5-foliolate, instead of 7-foliolate leaves and the
yellow petals. The type specimens were found at an altitude of
2250 m.

Mexico, State of Chihuahua : Sierra Madre, near Colonia
Garcia, 1899, Toivuscud &- Barber, 16.

Potentilla brunnescens sp. nov.

A tall strict perennial with short scaly rootstock : stem 4-5
dm. high, pubescence, especially above, tinged with brown or pur-
plish : basal leaves digitately 5-7-foliolate ; petioles often 2 dm.
long, villous with spreading hairs ; leaflets i\-'j cm. long, broadly
oblanceolate in outline, brownish strigose on both sides, paler be-
neath, pectinately divided to near the midrib ; segments linear, ob-
tuse : stem-leaves similar but short-petioled or subsessile, smaller:
cyme many-flowered, corymbiform : hypanthium and calyx vil-
lous-strigose : bractlets linear, obtuse, about half as long as the

173

174 Rydberg : Further Studies on the Potentilleae

ovate-lanceolate, acute sepals : petals yellow, 5-6 mm. long, ob-
ovate, about one fourth longer than the sepals.

The species is a member of the gracilis group and nearest re-
lated to P. flabellifonnis, from which it differs in the light brown
or yellowish pubescence and the lack of tomentum on the lower
surface of the leaves. The type was collected at an altitude of
nearly 2000 m.

Wyoming : Spread Creek, in the Teton Forest Reserve, 1897,

i*. liveedy, 212.

Potentilla Townsendii sp. nov.

A slender perennial with a deep root and a short caudex,
covered by the brown remains of old leaves : stem ascending, 3-5
mm. high, sparingly silky : petioles of the digitate basal leaves 7-10
mm. long, slightly hairy; stipules adnate to the petioles; the
free portion- linear-subulate ; leaflets 5-7, oblanceolate to almost
linear, serrate, acute, glabrous above, sparingly hairy beneath,
principally on the veins : stem-leaves 2-3, similar, but with
shorter petioles : stipule large, 3-4 cm. long, lanceolate or ovate-
lanceolate : cyme rather open : hypanthium and calyx silky-
strigose : bractlets linear-lanceolate, equalling the lanceolate
sepals : petals yellow, broadly obcordate, about i cm. long and
almost twice as long as the sepals.

This species is perhaps nearest related to P. heptaphylla, but
easily distinguished by the narrow leaflets and the larger flowers.
It was collected at an altitude of about 2250 m.

Mexico, State of Chihuahua : Sierra Madre, near Colonia
Garcia, July 17, 1899, Toivnsend & Barber, 10 j.

Potentilla filipes sp. nov.

A tall perennial with several stems from the short caudex.
Stem 3-5 dm. high, slender, silky-hirsute, simple up to the in-
florescence : few-leaved : basal leaves mostly 7-foliolate, digitate,
with hirsute petioles, 5-15 cm. long ; leaflets obovate to oblanceo-
late, obtuse, coarsely crenate, 2-6 cm. long, green and sparingly
strigose above, white-tomentose beneath ; stem-leaves similar, but
smaller, usually 5-foliolate, short-petioled : stipules ovate-lanceo-
late, entire : cyxne open : pedicels very slender, as well as the
hypanthium and calyx silky hirsute and more or less viscid :
bractlets linear-lanceolate, a little shorter than the lanceolate, acute
sepals : petals yellow or somewhat orange, broadly obcordate or

Rydberg : Further Studies on the Potentilleae 175

obovate and emarginate, 6-8 mm. long, exceeding the sepals :
stamens about 20.

This species is nearest related to P. pulcherrima Lehm., and I
had referred all the specimens cited below, except the type, to that
species. It differs from P. ptilcherriina in the very slender pedicels,
the total lack of tomentum on the hypanthium and the calyx and
the evident viscidity of these organs and the pedicels. It grows in
mountain meadows and on hillsides at an altitude of 2400-3200 m.

Colorado: Wahatoya Cafion, Spanish Peaks, 1900, Rydbcro
& Vrecland, 6ojg (type); West Mancos Canon, 1898, Baker,
Earle & Tracy, ^06 ; La Plata Mountains, 182.

Montana: Bridger Mountains, 1896, /. H. Fiodman, ^61.

Potentilla permollis sp. nov.

A stout hoary perennial. Stems simple, about 6 dm. high,
densely and softly pubescent with long whitish hairs : basal and
lower stem-leaves usually 7-foliolate, digitate ; petioles about i dm.,
long, pubescent : leaflets obovate to oblanceolate, obtuse, coarsely
cleft nearly half way to the midrib, with lanceolate lobes, densely
soft-pubescent on both sides, greenish above, white and somewhat
tomentose beneath: upper stem-leaves 3-5-fonolate, similar, smaller,
short-petioled : stipules of the stem-leaves large, adnate to the peti-
ole, ovate, often over 3 cm. long : cyme rather open : hypanthium
and calyx densely white-pubescent : bractlets linear-lanceolate,
about equalling the lanceolate sepals : petals yellow, 6-7 mm.
long, obcordate, exceeding the sepals by about one fourth : sta-
mens about 20.

This species is nearest related to P. BlascJikeana ; but differs
principally in the denser and white pubescence and the smaller
flowers. It grows in meadows.

Washington: Endicot, Whitman Co., 1898, A. D. E. Elmer,

1830.

Potentilla obovatifolia sp. nov.

A low perennial with a thick tap-root : stems ascending or de-
pressed, 2-3-leaved, 1.5-2 dm. high, sparingly villous : basal
leaves mostly digitately 7-foliolate with petioles 3-10 cm. long :
leaflets broadly obovate, obtuse, coarsely crenate-serrate, green
and sparingly hairy above, white tomentose beneath, . 5-3 cm. long :
stem leaves mostly 3-foliolate, small, short-petioled with broadly
lanceolate stipules : cymes open but few-flowered : hypanthium
and calyx densely silky : bractlets and sepals lanceolate, sub-

176 Rydberg : Further Studies on the Potentilleae

equal : petals obcordate, yellow, about 8 mm. long, nearly twice
as long as the sepals.

This species is a member of the concinna group and nearest
related to P. couciiuiifornm ; but is larger ; the leaflets are greener
and more glabrous above and the bractlets are longer. By the
size it approaches the gracilis group.

Mexico : Pringle, 68 go.

Potentilla Hallii sp. nov.

A more or less villous-hirsute ascending or decumbent peren-
nial with a short rootstock or caudex : stem about 3 dm. long,
sparingly villous-hirsute, terete and light green : leaves mostly
digitately 5-foliolate or the upper 3-foliolate or simple : basal ones
with petioles 3-5 cm. long : leaflets obovate, 1—3 cm. long,
coarsely serrate-crenate, green and slightly hairy above : paler,
hirsute and sparingly tomentulose beneath : upper stipules ovate,
entire, 12-15 mm. long: cyme rather open: hypanthium and
calyx hirsute, more or less tinged with purple : bractlets ovate-
lanceolate about one third shorter than the ovate acute sepals :
petals yellow, about 5 mm. long, scarcely exceeding the sepals :
stamens 20.

This species is nearest related to P. fastigiata, but is greener
and with a more open cyme. As to the pubescence of the leaves,
it resembles P. viridescens ; but is in every respect smaller and
the stem is not erect. On account of its small size, it could
just as well be referred to the maadata group as to the graciles.
The type grew at an altitude of about 1600 m.

California : Pine Ridge, Fresno County, 1900, Hall &
CJiandler, 182.

Potentilla propinqua

Potentilla diffusa A. Gray, Mem. Am. Acad. 1849 : 41. 1849.
Not Willd.

P. Hippiaiia, var. diffusa Lehm. Ind. Sem. Hort. Bot. Hamb.
1849 : 8.

P.Hippiana, \-a.x . propinqua Rydb. Bull. Torr. Club, 24: 3. 1897.

The original specimens were rather depauperate and the most
prominent characters which separate it from P. Hippiana were not
noticed by me when I prepared my Monograph of the North
American Potentilleae. The plant is much greener than P. Hip-
piana ; the upper surface of the leaves, as well as the pedicels.

Rydberg : Further Studies on the Potentillfjve 177

hypanthium and the calyx, is only slightly silky, not at all tomen-
tose ; the cyme in well-developed specimens is more open and
flat -topped ; and the upper segments of the leaves are decurrent
and sometimes confluent, which is never the case in P. Hippiaim.
By the latter characters it approaches P. ainbigeiis Greene. From
this it differs in the appressed pubescence, the smaller flowers and
the less coarsely toothed segments. Baker's no. 390, as repre-
sented in the herbarium of the New York Botanical Garden, was
named P. anibigcns by Professor Greene, but belongs unquestion-
ably to this species.

Horkelia Chandler! sp. nov.

A low silky perennial with very thick and woody root and
short cespitose caudex with erect branches densely covered by the
remains of old leaves. Stems scapiform, slender, silky villous,
7-16 cm. high : basal leaves numerous, 4-5 cm. long, densely
silky, terete, worm-like from the numerous small crowded seg-
ments : flowers about 4 mm. in diameter, rather many in sub-
capitate dense clusters, hypanthium cup-shaped, as well as the
calyx densely villous : bractlets linear, about two thirds as long as
the triangular-ovate sepals: petals yellow, linear, 1.5 mm. long,
shorter than the sepals : stamens mostly 5 : filaments subulate-
filiform.

This species closely resembles H. Miiirii in the leaves and the
thick root and caudex : but differs in the taller scape, the more
numerous flowers, the longer bractlets, and the narrower linear
petals. The type grew at an altitude of about 3450 m.

California: Mt. Goddard, 1900, Hall & CJiaiidler, yoo.

Drymocallis gracilis sp. nov.

A tall and slender, glandular pubescent perennial with a more
or less cespitose caudex. Stem about 6 dm. high, glandular-
pilose throughout, branched above : basal and lower stem-leaves
pinnately 7— 9-foliolate, 1—3 dm. long : leaflets obovate to flabelli-
form, 1—5 cm. long, coarsely incised, except at the bases, sparingly
pubescent on both surfaces : upper stem-leaves 3— 5-foliolate,
subsessile : pedicels slender, .5-4 cm. long, ver>' glandular:
hypanthium and calyx glandular : bractlets linear, about half as
long as the lanceolate acuminate sepals ; these in fruit often 8 mm.
long : petals white, small, broadly obovate, scarcely exceeding the
sepals : stamens, 20—25 : style slightly fusiform.

This species resembles most D. glandulosa incisa in habit and

178 Rydberg : Further Studies on the Potentilleae

leaf-form ; but differs in the white petals, more acuminate sepals
and the denser glandular pubescence. The type was collected at
an altitude of about 1600 m.

California: Pine Ridge, Fresno County, 1900, Hall &
Chandler, ij8.

II. SOME SPECIES FROM GREENLAND

Some time ago Mr. Morten Pedersen, Assistant at the Botan-
ical Garden of Copenhagen, requested me to determine a collec-
tion of Potcntillae collected by him in Greenland. He also kindly
sent me several other specimens from the herbarium of the Botan-
ical Museum of such forms that I had not seen at all or of which
I had had only imperfect material. With the help of this material
I have been able to settle several important points and consider-
ably modify the disposition in my monograph.

Potentilla maculata Pourr. Act. Toloss. 3: 316. 1788

The following specimens additional to those given in my mono-
graph,* belong to this species :

Greenland: KinguaTasiusok(6i°45'), 1889; Scoresby Sund,
189 1 ; Danmarks, Oe. 1892 ; Jameson's Land, 1891, N. Harts;
Godthaabs Fjord, 1883,/ A. D. Jensen ; Alangua, 1885, 6". Han-
sen ; Umanaks Fjord, 1885, P. Eberlin ; (locality not given), Raben,
I ; Scoresby Sund, 1891, H. Hartz ; \ Kvan Valley, behind
Ujaragsugsuk, 1898, Morten Pedersen, 70 j ; Ekalunguit Itivnerit
^2g ; both on Disco.

Potentilla maculata var. firma Lange, Consp. Fl. Groenl.
235 is scarcely that of Lehmann,| although it agrees perfectly
with Lehmann'S description. The latter was in reality based upon
P. alpestris a firnia Koch. I have no means to find what this
really is ; but Lehmann cites P. alpestris c riibens Hegetschw. and
P. rubens Vill. as synonyms. These names represent a coarser
plant from Austria, Switzerland and Italy with longer and more
acute teeth to the leaves. I think it is a good species distinct from

* Mem. Dep. Bot. Columbia Univ. 2 : 59-

t This sheet is labeled Potentilla 7nacnlata Pourr. szx. gelida C. A. Mey. , as it
appears on the label from the fact that a few of the basal leaves are ternate, a condition
not uncommon in P. maculata. It has nothing to do with P. gelida C. A. Meyer.

\ Rev. Pot. 1 20.

Rydberg : Further Studies on the Potentilleae 179

P. inacuhita. The Greenland plant cited by Lange agrees fully
with the typical P. viacnlata except that the leaflets are broader,
more round and overlapping each other by the margins. I
scarcely think that it deserves a varietal name. If it does it must
bear some other name than finna, as this is preoccupied. The
specimens cited by Lange and the only one seen is the following:
Greenland : Holsteinsborg, 1884, Eug. Warming & TJi. Holm.

Potentilla Langeana sp. nov.

P. maadata ^ hirta Lange, Consp. Fl. Groenl. 6. 1880. Not
P. hirta L.

A cespitose perennial. Stems erect or ascending, 2-3 dm.
high, sparingly hirsute with erect branches : basal leaves digitately
5-foliolate : stipules large, adnate, lanceolate, 1-2 cm. long :
petioles 3-8 cm. long, sparingly hirsute : leaflets cuneate-obovate,
more or less densely silky-hirsute on both sides, 1-3 cm. long,
coarsely toothed above the middle w^ith oblong ovate teeth, the
cuneate base entire : lower stem leaves similar but smaller and
short -petioled : upper stem-leaves ternate or simple and subsessile :
stipules ovate, acutish, about i cm. long : cyme 3-7-flowered ;
hypanthium silky hirsute : bractlets oblong to lanceolate, acute or
obtuse, about one fourth shorter than the lanceolate sepals : petals
broadly obcordate, 7-8 mm. long : stamens about 20.

The species differs from P. maciilata not only in the characters
given by Lange, the several-flowered cymes and the long pu-
bescence of the leaves ; but also in the more acute teeth, the longer
and more acutish bractlets and narrower sepals. In P. maadata
both the teeth of the leaves and the bractlets are rounded at the
apex and the latter are only one half or two thirds as long as the
sepals. P. Langeana is in reality nearer related to P. verna than
to P. maadata but distinguished from that by the pubescence. In
Conspectus Florae Groenlandicae several more specimens are cited
belonging to P. maadata ^ hirta Lange. The following are the
only ones seen by me :

Greenland: Amarahk Fjord, 183 i,/. rW//(type) ; Kangerd-
luarsuk Fjord, 1884, Warming & Hohii ; Natsilik, 5. Hanson;
Ekalunguit Itivnerit on Disco, 1898, Morten Pedersen, ^ig ;
Ekigtok in Disco Fjord, 168^ (a taller and more glabrate form*) ;

■^ This was labeled /". w«rr«/rt/'<z var. /j)/r^;w?Va (Ramond) Lehm. , which is quite
a different plant, having thick leaves with short teeth directed forward.

180 Rydberg : Further Studies on the Potentilleae

Kuanersuit on Disco, 2770 ; Engelskmandens Havn, Disco, iSjc)
(a slender depauperate form*).

Potentilla emarginata Pursh, Fl. Am. Sept. 353. 18 14

Undeveloped or depauperate specimens of this species resemble
often very closely P. nana and I am now inclined to regard the
two species as forms of one. In Alaska and other arctic regions
of the American Continent they seem distinct enough but the plants
from Greenland, Spitzbergen and Novaja Semlja are often so con-
fusing that it is hard to tell to which to refer them.

The specimens collected by Mr. Pedersen are as follows, those
approaching P. nana are marked by (?).

Greenland: Skarvefjord, 1898, Pedersen, 2jg ; Ingigsok,
2i8a^.nA 2i8b ; Mudderbugten, <)66 (?) ; Ekalunguit Itivnerit,
1414. ; Plateau behind Ujaragsugsak, 3ogg ; all on Disco.

Potentilla nana Willd.; Schlecht. Mag. Ges. Naturf. Fr. Berlin.

7 : 296. 181 3

Among the many species of P. emarginata sent from Copen-
hagen, there are two sheets, however, that represent the typical P.
nana with its round teeth and rounded obtuse sepals and bractlets,
viz.:

Greenland: Manatsook, 1883, Dr. Berlin; Upernaviarsur,
1887, Ryder, 7.^

Potentilla nivea L. Sp. PI. 499. 1753

Of the typical P. nivea the collection sent contains many sheets,
the following was the only one collected by Mr. Pedersen :

Greenland: Rocks behind Kuanersuit on Disco, 1898, J/!
Pedersen, 2J06.

*0n the label is written ^^ Potentilla f?iaculata Pourr. var. debilis Lehm. in var.
hirtumV.gQ. transiens." Lehmann cites under the var. debilis, P. incisa Desf. as a
synonym. The latter is figured in Nestler's Monograph as a large plant with nar-
rowly cuneate leaflets deeply incised at the apex, and quite unlike the specimens cited
above. The latter I cannot distinguish from the typical P. Langeana except in size.

I The last was determined by Lange as P. tiivea prostrala, which must be a mis-
take as it is unlike the other specimen collected by Vahl and does not agree with the
description.

Rydberg : Further Studies on the Potentilleae 181

PoTENTiLLA NiVEA Altaica (Bunge) Rydb. Mem. Dep. Bot. Co-
lumbia Uni. 2: 86. 1898

Poteutilla nivca var. arcnosa Lange Consp. Fl, Gr. 236. Not
Turcz.

The specimens collected by Warming and Holm were deter-
mined by Lange as P. nivea aroiosa Turcz. but that variety is
described as having glomerate flowers, which is not the case in
Warming and Holm's specimens. These do not differ from the
common form of P. nivca Altaica except that the flowers are some-
what smaller. In fact they are as like the figure in Ledeb. Icon.
Fl. Ross. 4 : 329 as I have seen. If Lange's determination in
this case were correct the varieties Altaica and arenosa must be
united. I think, however, that the latter is quite different and re-
stricted to Asia. The following specimens belong here and are
not recorded in Conspectus Florae Groenlandicae :

Greenland : Amaralik Fjord, 183 1,/. Vahl ; Scoresby Sound,
1892, N. Harts ; Kakatsiak, 1885, 5. Hanson; Christianshaab,
1884, Warming & Hohn ; rocks behind Kuanersuit on Disco,
1898, M. Pedersen 2702-j, 2707-g (?) ; * Christianshaab, 1833,
/. Vahl {H) A

Potentilla nivea pallidior Sw. Sum. Veg. Scand. 19. 1874
Poteutilla nivea snbviridis 'L&deh. Y\. ^oss. 2: 57. 1844.

The following specimens belong here :

Greenland: Scoresby Sound, 1892, N. Harts; Godhavn,
1 87 1, Th. Fries.

Potentilla subquinata (Lange)

Potentilla nivea pentaphylla Lehm. Nov. Stirp. Pug. 9 : 69.
1 85 1. Not P. pentapJiylla Richt. 181 5 ; P. nivea subquinata
Lange, Consp. Fl. Groenl. 9. 1880 ; P. nivea guingiiefolia Rydb.

* AH of Mr. Pedersen' s specimens are sterile and without fully developed leaves.
They are remarkable for their long branching caudices and may belong to the variety of
the next species described below or perhaps to an undescribed species. They cannot
be determined satisfactorily.

■f- This was determined by Lange as var. trostrata but it does not agree with Rott-
boell's description especially as to the glomerate heads, sinuate-dentate leaves, revolute
margins and purple veins. I refer this doubtfully here as it differs from all other speci-
mens seen in the narrow leaflets.

] 82 RvDBERG : Further Studies on the Potentilleae

Bull. Torr. Club, 23: 302. 1896; P. quinquefolia Rydb. Mem.
Dep. Bot. Columbia Univ. 2: 76. 1898.

When I changed P. nivca pentapJiylla Lehm. to P. nivea quin-
quefolia, I did not know that it was identical with Lange's var-
S7ihq2iinata. From his description, especially from the words
" eleganter pinnatifidis " I was lead to the supposition that it
represented the quinate form of var. Altaica. Consequently I
referred Lange's var. subqidnata to that variety. As Lange's name
is older than my own, it should be adopted. I am well satisfied
with the change, as my own as a specific name is a misnomer. The
following specimens belong here.

Greenland : Disco Fjord, 1894, P. Soeimsen, joi ; Akingdlek,
1890, N. Hariz, i ; Kingua Orpiksuit, 2; Atanekerdluk and
Kuanersuit on Disco, 1871, Tli. Fries; Kingigtok, 1890; N.
Hartz ; rocks behind Kuanersuit on Disco, 1898, M. Pedcrsen,
2'ji8, 2yi'j, 2yij, 2yi^a, 2"/ 12 ; Disco Fjord, ;^(^o6 ; Skansen,
Disco, p.?7 ; Kuganguak Valley, 2j/j and 4g6 ; Vajat-shore, Un-
artuarsuk, //j.

Spitzbergen ; Kap Thorsen, 1896, E. Joergeiisen.

Potentilla subquinata Pedersenii var. nov.

Caudex with elongated branches covered by the remains of
old leaves ; stems low, less than i dm. high, 1-3-flowered, as well
as the petioles covered with long white hairs: leaflets small, 1-
1.5 cm. long, broadly obovate :. white silky on both sides and to-
mentose beneath, in age more glabrate above, bractlets very nar-
row, linear or linear-lanceolate.

Greenland: Vaigat Assuk, Disco, 1898, M. Pedersen, 4J0
(type) and io6ia. Rocks behind Kuanersuit, 2^11, 2^14, 2ji6,
and 2'jio ; Ingigsak, ^jj ; Rocks by Mudderbugten, g6j ; Mun-
garut, g24 ; all on Disco.

Potentilla Vahliana Lehm. Mon. Pot. 29 and 172. 1820

Fine specimens of this species were also in Pedersen's collec-
tion.

Greenland : Vajgat, Assuk on Disco, M. Pedersen 1061b.
Lyngmarken by Godhaven, 80; Rocks behind Kuanersuit on
Disco, 2'ji^b; Roede Elv, Disco, j^^ib.

Rydberg : Further Studies on the Potentilleae 183

POTENTILLA PULCHELLA R. Br. ill Ross' Voy. 1 42

Greenland: Kugsinarsuk, Disco, 1898, M. Pedersen, 4^4;
Vaigat-shore, Assuk, ./j/ and p.// ; Kutdlisat, jgSa ; Jameson's
Land, 1891, ^V. Hartz.

PoTENTiLLA PULCHELLA ELATiOR Langc Consp. Fl. Grocnl.

4. 1880

Greenland: Kardlunguak, 1890, iV. Hart:;; Atanekerdluk,
1 87 1, Th. M. Fries ; Vaigat Shore by Assuk, 1898, M. Pedersen,
462.

Argentina Egedii (Wormskj) Rydb. Mem. Dep. Bot. Colum-
bia Uni. 2 : 158. 1898
Greenland : Disco Fjord by Kuanersuit, 1898, M. Pedersen,
1448 ; Ungorsivik by Maligiak, 4j8.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 14

STUDIES ON THE ROCKY MOUiNTAIiN

FLORA— V.

BY P. A. RYDBERG

NEW YORK
1901

[Reprinted from the Bulletin of the Torrey Botanical Club, 28 : 266-28i. 21 May, igoi.l

Studies on the Rocky Mountain Flora.— V,

By p. a. Ryuberg

Sporobolus aiistatus sp. nov.

Vilfa depmiperata \-&.x. filiforviis Wats. King's Exp. 5 : 376, in
part (as to the awned form). 1871. Not Thurb.

Perennial with short branching rootstock. Culm usually only
4-6 cm. high : internodes short, usually covered by the open strongly
striate sheath ; ligules lanceolate, acute, about 1.5 mm. long : leaf-
blades 8-14 mm. long, and about i mm. wide, striate and puberu-
lent on the upper surface : panicle few-flowered, very narrow, with
short erect branches ; empty glumes almost equal, or the inner
slightly longer, 1-1.25 mm. long, less than half as long as the
flowerinsf gflume, ovate, acutish or obtuse and somewhat erose at
the apex : flowering glume about 2.5 mm. long, strongly veined,
long-strigose on the veins and tipped with an awn .5-1 mm. long ;
palet almost equal to the flowering glume (awn excepted) in
length, acuminate, but not awned, strigose.

This species is closely related to 5. filiformis (Thurb.) Rydb.
differing in the lower habit, less exserted panicle, firmer empty
glumes and the presence of a distinct awn. It grows in wet places
at an altitude of 2000-2500 m.

Wyoming: Big Horn Mountains, Sheridan Co., 1899, F.
Tweedy, 2ig6 (type) ; Spread Creek, 1897, 2^.

Utah : Bear River Canon, 1 869, 5. Watson, 1281.

Poa platyphylla Nash & Rydb.
Poa trivialis var. occidentalis Vasey, Desc. Cat. Grasses 85.
1885. '^ot Poa flexuosa vdiT. occidentalis YdiSey. 1878.

Poa occidentalis Vasey, Cont. U. S. Nat. Herb, i : 275. 1893.

Carex ebenea sp. nov.

Perennial by a cespitose rootstock. Stem 2-4 dm. high,
strongly striate ; sheath with a conspicuous membranaceous ligu-
lar portion, ligule proper rounded, about 2 mm. long ; leaf-blades
flat, 1-2 dm. long, 3-6 mm. broad, strongly nerved : spikelets
about I cm. long in a dense globular or rounded- conical head :

266

267 Rydberg: Studies on the Rocky Mountain Flora

shining, brownish black with a Hghter midrib, lanceolate, acute,
3-4 mm. long : perigynia lanceolate, tapering gradually into
a long beak, with the beak about 5 mm. long, dark brown ;
staminate flowers at the base and mixed with the pistillate :. upper
portion of the wings and the beak scabrous on the margins ;
teeth at the apex of the beak very short, subulate : styles 2 : achene
oblong, lenticular, nearly 2 mm. long and fully i mm. broad.

This species is nearest related to C. f estiva and has been labeled
C. f estiva Haydeniana, but it is not the same as the original of
that variety, which has broadly ovate perigynia. C. ebenea differs
also from all forms of C. f estiva in the form of the perigynia and
in the dark glossy color of the bracts and perigynia.

Colorado: Pikes Peak, 1900, F. Clements (type); Windy
Point, 1900; Mt. Harvard, 1896, 7; Grecian Bend, 1896, Bot-
tomless Pit, and Saddle, Clements ; between Cheyenne Mountain
and Seven Lakes, 1896,^5". A. Bessey ; I ronton, 1899, C. C. Curtis ;
Telluride, 1894, F. Tzveedy, 1^4; Chambers Lake and Cameron
Pass, 1896, C F. Baker; Clear Creek Canon, 1878, M. E. Jones,
26 J ; Silver Plume, 1895, Rydberg, 2460; Pagosa Peak, 1899, C.
F. Baker, 2jj.

Streptopus curvipes Vail sp. nov.

Simple, glabrous, except the margins of the leaves and the
peduncles. Stems 1-3 dm. high, from a slender rootstock cov-
ered with few fibrous rootlets : leaves sessile, ov^al or oblong-
lanceolate, 3-8 cm. long, acuminate at the apex, rounded and
slightly clasping at the base, 3-5 -nerved, the margins finely gland-
ular-ciliate : flowers 3-5, solitary: peduncles not geniculate, 5-15
mm. long, glandular-pubescent : flowers pale purple or rose-
colored : perianth-segments lanceolate, 5-7 mm. long, minutely
glandular-pubescent on the inner surface : anthers 2-beaked ;
beaks slender, about half the length of the anther : style 3-cleft,
the spreading branches stigmatic along the inner side : berry glo-
bose, when mature brick red, 7-9 mm. in diameter : seeds clavate,
3-4 mm. long, with many cross-striate ridges.

Differs from Streptopus roseus in its simple habit and the non-
geniculate and much shorter peduncles. The beaks of the anthers
seem to be a trifle longer than in 5. roseus. It is closely related to
S. brevipes Baker, but is larger and coarser in every respect. The
flowers of the latter species are not known.

British Columbia : Asulkan Pass, alt. 4400 ft., June to July,

Rydberg: Studies on the Rocky Mountain Flora 268

1897, Mrs. Zoe W. Palmer (type) ; Glacier, alt. 4122 ft., June to
July, 1897, Mrs. Zoe W. Palmer ; Glacier, June, 1897, Mr. &
Mrs. Cornelius Van Brunt.

Alaska : Juneau, along the mountain side, July 24, 1891, Miss
Grace E. Cooky ; Yes Bay, July 10, 1895, Thomas Hozvell, 1664
(all in Herb. N. Y. Botanical Garden or Columbia University) ;
Ferd. Bislwff.

Oregon : E. Hall, j2i.

Washington: Skamania Co., Aug., 1886, Suksdorf ; Mt.
Adams, 3-400 ft., Suksdorf, 44; June, 1879,/. Hozvell {\.\\q last
five specimens in Herb. Gray).

Vagnera brachypetala sp. nov.

A tall stout perennial, 5-8 dm. high. Stem striate, puberu-
lent, especially the upper portion : leaves subsessile or short-
petioled, oval or ovate, 8-18 cm. long, 4-7 cm. wide, often short-
acuminate and twisted at the apex, with 5-7 stronger nerves and
numerous weaker ones: panicle on a peduncle 4-5 cm. long,
rather dense, 3- 10 cm. long and 2-5 cm. broad : petals and sepals
oblong, 1-1.5 mm. long, scarcely half as long as the broadly di-
lated, lanceolate-subulate petaloid filaments : style about .5 mm.
long : berry dark purple, about 6 mm. in diameter.

This species is closely related to V. racemosa and V. amplcxi-
catdis, perhaps most nearly to the former, but is easily distinguished
by the short petals and the purple fruit. In V. racemosa the
petals and sepals are almost as long as the filaments and the fruit
is 7-8 mm. in diameter and red with purple spots. V. brachypetala
grows on hillsides up to an altitude of 2500 m.

British Columbia: Glacier, at the "Loup," 1897, Mr. &
Mrs. Cornelius Van Brunt (in fruit, type) ; Deer Park, Columbia
River, 1890, John Macoun ; Victoria, Vancouver Island, 1893,
John Macoun, 5gg8.

Washington: Seattle, 1891, C. V. Piper, ig8.

Idaho: Lake Waha, 1892, Sandberg, MacDougal & Heller,
228.

Vagnera leptopetala sp. nov.

A slender glabrous perennial, 2-3 dm. high. Rootstock very
slender for the genus, white, only about 2 mm. in diameter: stem
erect, strict or the upper portion somewhat zigzag, striate and
pale : leaves oblong-lanceolate, 6-8 cm. long, 1-2 cm. wide, light

269 Rydberg: Studies ox the Rocky Mountain Flora

green, sessile, long-acute, with 3-5 stronger nerves and 12-14
weaker ones : raceme terminal, simple, 3-6-flowered : pedicels
5-8 mm. long : petals and sepals linear, acute, about 4 mm. long
and less than i mm. wide, very thin, white : fruit not seen.

This species is nearly related to V. stcllata and V. liliacea, but
differs in the narrow petals and sepals, the slender rootstock and
the lighter green foliage. It grows in rich soil in caiions at an
altitude of 2000-2100 m.

Colorado : Headwarters of Sangre de Christo Creek, 1 900,
Rydberg & Vreeland, 6^^r (type); Dark Cafion, Pikes Peak,
1900, Fred Clements.

Limnorchis purpurascens sp. nov.

A rather stout plant, 3-5 dm. high, with fleshy-fibrous roots.
Leaves ovate to lanceolate, acute, 6—10 cm. long, 1.5—3 cm. wide,
dark green : bracts lanceolate, the lower exceeding the flowers :
spike rather dense : flowers 10-12 mm. long : lateral sepals green,
oblong-linear, or linear, obtuse, 4-5 mm. long ; the upper sepal
tinged with purple, broadly ovate, erect, obtuse : petals slightly
shorter, erect, purple, lanceolate, oblique : lip broadly linear-lan-
ceolate, about 5 mm. long, purplish, scarcely at all dilated at the
base, the edges almost straight : spur scarcely more than half as
long as the lip, much thickened and saccate.

This species belongs to the L. Jiyperborea group, and is perhaps
nearest related to that species. It differs, however, in the purple
petals and lip and the shorter and more saccate spur. The spur
has almost the same form as that of L. stricta ; but from that
species it differs in the dense spike and the broader lip. L. pur-
purascens grows in damp woods at an altitude of 2700-3000 m.

Colorado: Iron Mountain, 1900, Rydberg & Vreeland, 6^1 /f.
(type); Manitou, i goo, Fj-ed Clements, ijz ; Georgetown, 1878,
M. E.Jones, J14.

Piperia gen. nov.

Montolivaea Rydb. Mem. X. Y. Bot. Gard. i : 106. 1900.
Not Reichenb.

Professor C. V. Piper of the Agricultural Experiment Station
at Pullman, Washington, has called my attention to the fact that
the genus Montolivaea was based not on PlatantJicra elegans Lindley,
or Habenaria elegans Bolander ; but on Mo7itolivaea elegans ^e.\ch.-

Rydberg: Studies on the Rocky Mountain Flora 270

enb., or Habcnaria clcgaiis Jackson, of a much later date. The
genus that I shortly characterized under the name of Montolivaci
in the Memoir cited above, is therefore without a name. As I
find that no genus has at yet been dedicated to Professor Piper, I
take the pleasure in naming this for him. The genus contains at
least three species, viz.

Piperia elegans (Lindl.)
PlatantJicra elegans Lindi. Gen. & Sp. Orch. 285. 1835.
Hahenaria elegajis Boland.; Wats. Bot. Calif 2: 133. 1876.
Moiitolivaea elegans Rydb. Mem. N. Y. Bot. Gard. i : 106.
1900. Not Reichenb.

Piperia Unalaschensis (Spreng.)

Spirantlies Unalascheitsis Spreng. Syst. 3: 708. 1826.

Habenaria Unalaschensis Wats. Proc. Am. Acad. 12 : 277.
1876.

Montolivaea UnalascJiensis Rydb. Mem. N. Y. Bot. Gard. i :
107. 1900.

Piperia elongata sp. nov.

A slender strict plant, 4-7 dm. high. Corm ellipsoid, about 2
cm. long and i cm. in diameter : leaves 2, near the base ; blades
lanceolate or oblanceolate, acute, 8-15 cm. long, 1-2 cm. wide:
spike elongated and lax, 2-3 dm. long : flowers about i cm. long :
sepals green, about 5 mm. long, the upper lanceolate, the other
two linear obtuse : petals lanceolate, oblique, equalling the sepals :
lip entire, broadly lanceolate : spur slender, scarcely clavate, almost
cylindrical, arcuate, 10-12 mm. long, about .7 mm. thick.

This is somewhat intermediate between the two preceding.
It has almost the same flowers as the first and the general habit of
the second. P. elegans, with which it has generally been con-
fused, has a stouter, more fleshy stem, thicker bracts and a short
crowded spike. P. Unalaschensis has the same habit as the present
species, but the spur is very short and saccate. P. elongata grows
in open woods, up to an altitude of 700 m.

Idaho : Priest Lake, near lower end, 1900, D. T. MacDongal,
168 (type); Priest River Valley, ij^.

Washington : W. Klickitat County, 1885, W. N. Snksdorf ;
Seattle, 1889, C. V. Piper, loSi.

271 Rydberg: Studies on the Rocky Mountain Flora

Corallorrhiza Vreelandii sp. nov.

Scape rather stout, 2-4 dm. high, dark purplish brown.
Sheaths, especially the lowest one, much inflated, purplish brown
with dark purple striations : flowers 6-15, about 15 mm. long, in
fruit drooping : sepals oblong, 7-8 mm. long, brown with 3 purple
stripes : petals similar but slightly broader and with 4 stripes :
lip ovate, entire, with a small gibbosity at the base : spur none.

This is nearest related to C. striata, from which it differs
in the smaller flowers and narrower sepals and petals. The type
was found growing among alders and willows at the margin of a
small lake, at an altitude of about 2700 m. It was first dis-
covered by Mr. F. K. Vreeland, my enthusiastic associate and
pleasant companion in field work last summer.

Colorado : Veta Mountain, 1900, Rydberg & Vreeland, 64.1S.

Salix Wyomingensis sp. nov.

A shrub i m. high or less. Bark of the main trunk gray ;
that of the branches dark-brown or grayish, shining, some-
what flaky ; young branches appressed-villous ; buds brown,
shining, pubescent when young : leaves small, rather crowded ;
petioles 2-4 mm. long ; blade oval or obovate, acute, 1-3 cm.
long, entire or exceedingly minutely callous-denticulate, upper
surface green and shining, slightly pubescent when young, glabrate
in age ; lower surface pale bluish green, appressed silky-villous :
aments borne on very short leafy branches, about 3 cm. long,
8-10 mm. in diameter: bracts brown, oblong or ovate, obtuse:
pedicels scarcely i mm. long : ovary villous, but green, when fully
developed only about 3 mm. long: style .5 mm. long: stigmas
short, slightly 2-cleft.

This species is nearest related to 5. glaucops Anderson, but
differs in the small oval or obovate leaves and the much smaller
and greener ovaries. It grows at an altitude of 2100-2700 m.

Wyoming : Eastern Slope of Big Horn Mountains, headwaters
of Clear Creek and Crazy Woman River, 1900, F. Tiveedy, 3434.
(type) and 34.33.

Salix stenophylla sp. nov.

A shrub 2—5 m. high. Bark of older stems light brownish
green, smooth ; that of the slender strict branches brown : young
shoots silvery pubescent : leaves narrowly linear, 'i^—'j cm. long,
about 3 mm. wide, acute, entire or minutely and distantly dentic-
ulate, permanently finely silky strigose, but not white : pistillate

Rvdberg: Studies on the Rocky Mountain Flora 272

aments 3-4 cm. long at the ends of short leafy branches : bracts
yellow, oblanceolate, deciduous, acute, 2-2.5 ^^- long- rather
sparingly silky-villous : pistil green, about 4 mm. long, glabrous-
except the swollen portion at the end just below the red stigma:
stipe about i mm. long : style none : stigma minutely 4-lobed :
fruit glabrous, narrowly ovoid-conic, about 6 mm. long, and 1.5
mm. in diameter : staminate aments 2-3 cm. long, 5 mm. in diam-
eter : bracts like those of the pistillate aments, but more obtuse
and more villous : stamens 2, about twice as long as the bracts :
filaments slightly hairy, especially below.

This species is a member of the longifolia group and nearest
related to .S". exigjia Nutt., at least as understood by Professor
Rowlee ; but differs in the narrower leaves, the distinct stipe and
the peculiar swelling just below the stigma. It grows on river
banks up to an altitude of 2500 m.

Colorado: Cuchara River, below La Vcta, 1900, Rydberg &
Vreeland, 6jp2 (staminate flowers) and 6jpj (pistillate, type) ;
Mancos, 1898, Baker, Earlc & Tracy, 12 j (staminate) and 102
(pistillate in fruit).

New Mexico: 185 1-2, Wright, 1874 and iSyj ; 1847, Feitd-
Icr, 813.

Arizona: San Francisco Mountains, 1881, Rusby, jyi.

Salix padifolia sp, nov.

A shrub 1-7 m. high, with light brown smooth bark. Young
twigs strict, glabrous and shining, yellow, or often brown or purple :
stipules ovate or rounded, glandular-dentate : leaves with petioles
5 8 mm. long ; blade oval or broadly elliptic, crenate, short-acute
or obtusish, rounded at the base, 3-5 cm. long, 1.5-2 cm. wide,
when young sparingly covered with silky hairs, but soon glabrate,
dark green above, paler beneath : pistillate aments 3-4 cm. long,
densely flowered, appearing usually before the leaves, borne on
very short branches and subtended by 1-4 small leaves : bracts
obovate, fuscous, covered on the outside with white wool : pistils
nearly sessile, glabrous : style about 1.5 mm. long : stigmas 2,
nearly i mm. long, 2-cleft : capsules ovate-conic, about 6 mm.
long: staminate aments almost sessile, 2-3 cm. long, 10-12 mm.
in diameter : bracts as those of the pistillate aments : stamens 2;
filaments glabrous.

This is a species of the cordata group and nearest related to
5. Mackenziana ; but differs in the shorter oval leaves, the shorter
and thicker aments and the exceedingly short stipes. It has gen-

'273 Rvdberg: Studies ox the Rocky Mountain Flora

erally been referred to 6". cordata ; but has much thicker and
shorter ainents and the leaves are quite different, thin, very short
and broad, never acuminate and never sharply serrate. It grows
along streams at an altitude of 2000—3000 m.

Colorado : Tributaries of Turkey Creek, 1900, Rydbcrg &

Vreelaiid, 6j8g (fruit, type); Ojo, 6j86 (staminate) and 6j8y

( pistillate ) ; Pass Creek, 6j88 (pistillate ) ; Los Finos, 1899, Baker,

2"]! ; Bob Creek, iSgS, Baker, Earle & Tracy, lys > Silverton,

1895, F. Tu'ccdy, 268 (?).

Wyoming : Golden Gate, 1899, Aven & Elias Nelson, jj4Q.

Montana : Ten miles east of Monida, 1899, Aveti & Elias
Nelson, 342^.

Salix flava sp. no v.

A shrub or small tree, 4—7 m. high, with grayish yellow rough
bark. Branches short and divergent, light yellow, smooth and
shining : stipules rounded, entire : leaves with petioles 2-6 mm.
long ; blades lanceolate, short -acuminate or acute, entire or indis-
tinctly crenulate, rather firm, yellowish green, glabrous on both
sides, 3—7 cm. long: pistillate aments 2-3 cm. long, almost ses-
sile, subtended by 1-2 leaves or naked : bracts very short, obo-
vate, fuscous, densely long-woolly : pistil stipitate, glabrous ; stipe
in fruit often 2 mm. long: style about .5 mm. long: stigmas ob-
long, subentire : capsule ovate, about 6 mm. long : staminate
aments 2-3 cm. long, sessile : bracts as in the pistillate aments :
stamens 2 ; filaments glabrous.

This is also a member of the cordata group and nearest re-
lated to S. lutca Nutt., differing in the less acuminate and suben-
tire leaves, which are not paler beneath, and the longer stipes. It
grows along streams at an altitude of i 500-2000 m.

Wyoming : Green River, 1895, Rydbcrg (type) ; Spread Creek,
1897, F. Tiveedy, J02.

Montana: Boulder River, 1888, F. Tu'ccdy, 6j.

Idaho: Beaver Caiion, jSg^, Rydbcrg.

Nevada : Unionville, 1868, 5. IVatson, logy.

Utah : Wahsatch Mountains, 1 869, 5. Watson, iog6, in part.

Colorado : South Park, 1873,/. Wolfe ; Cucharas Valley, near
LaVeta, \ goo, Rydbcrg &■ Vrecland,6j'/'/ ; Manitou, 1878, J/ E.
Jones, JO.

Blitum hastatum sp. nov.

A slender annual, 2-4 dm. high. Leaves very thin ; blades

Rydberg: Studies on the Rocky Mountain Flora 274

3—7 cm. long, ovate or ovate-lanceolate in outline, but often hastately
lobed at the base, which is broadly cuneate and dccurrent on the
slender petiole ; the upper ones smaller and generally not hastate :
lower petioles 2—4 cm. long ; flowers in small glomerules in the
upper axils and on a slender interrupted terminal spike : seeds
brownish black, smooth and shining, about i mm. long.

This species differs from B. capitatum in the hastate or entire,

never toothed, thin leaves and in the inflorescence which is more

inclined to be naked above, more lax and with smaller glomerules.

It grows at an altitude of 2000—3000 m.

Wyoming: Buffalo, 1900,/^. Trcir^^', j^p f (type).

Utah: Alta, Wahsatch Mountains, 1879, M. E. Jones,

1181.

Nevada : Above Thousand Springs Valley, 1868, 5. Watson,

977-

Alsine Curtisii sp. nov.

Slender, erect, branched, 2—6 dm. high with a slender horizon-
tal rootstock. Stem sharply 4-angled, glabrous, except at the in-
florescence : leaves linear or lance-linear strongly i -nerved, sessile,
rounded and half clasping at the base, long-attenuate at the apex,
3-6 cm. long, 3—5 mm. wide, those of the inflorescence smaller :
branches of the many-flowered cymes divergent, very viscid-pubes-
cent : sepals about 3 mm. long, thick, ovate-oblong, obtuse or
acutish with a narrow white margin : petals about 7 mm. long,
cuneate with a broad and deep sinus at the apex : seeds very few,
dark brown, almost 2 mm. in diameter.

This species is nearest related to A. Jamesii and has been con-
fused with it ; the differences are mostly in the leaves and sepals.
In A. Curtisii the former are scarcely half as wide as those of A.
Jamesii, and the sepals of the former are rather thick while in the
latter almost membranous. A. Curtisii grows at an altitude of 1800
to 3000 m.

Wyoming : Headwaters of Cliff Creek, 1900, C C Curtis
(type).

Utah : Wahsatch Mountains, 1 869, S. Watson, i^g (in part) ;
Beaver Valley, 1877, E. Palmer, ji^i; Alta, Wahsatch Moun-
tains, 1879, 3f. E. Jones, 1160.

Nevada : East Humboldt Mountains, 1868, 5. Watson, ijg (in
part).

Arizona: Mogallon Mountains, 1881, //. H. Rushy.

275 Rydberg: Studies on the Rocky Mountain Flora

Arenaria confusa sp. nov.

A slender diffuse plant with cespitose, perennial, but not ligneous
base. Stems slender, branched, 4-6 dm. long, finely puberulent :
leaves opposite, linear-lanceolate, pointed, with a strong midrib,
puberulent, 1—2 cm. long : pedicels in fruit divergent, about i cm.
long, slightly bent under the calyx : sepals linear-lanceolate, acumi-
nate, scarious-margined, not tuberculate-punctate, about 3 mm.
long : petals about three fourths as long as the sepals : capsule
broadly ovoid, about 4 mm. long.

This species is related to A. lanuginosa and A. saxosa, and some-
what intermediate between the two. In habit it resembles most
the former, but does not have the subverticillate or fascicled leaves ;
the sepals are narrowly lanceolate instead of ovate and not tuber-
culate-punctate, the stem is not retrorsely pubescent as in that
species and the petals are larger. In all specimens of A. lanuginosa
seen by me the petals are either lacking or not more than half as
long as the sepals. A. saxosa is subligneous at the base, has low
stems, short leaves, mostly less than i cm. long and sepals even
in flower 4-5 mm. long. Dr. B. L. Robinson, in the Synoptical
Flora, remarks under A. alsinoides [A. lanuginosa): "A more
western form, represented from New Mexico by Fendler's 58 and
62 and Wright's 864, has slightly firmer stems, more numerous
subpaniculate flowers, and leaves less narrowed at the base. In
all these respects it shows a transition to the following." I have
not seen the numbers of Fendler's collection cited ; but Wright's
864 as represented in Columbia University Herbarium belongs to
A. confiisa. On the sheet of Rusby's 38, cited below, found in
the Columbia University herbarium. Dr. Robinson has written in
pencil : " Apparently this is merely a lax form of A. saxosa Gray.
The earliest leaves are crowded, the upper internodes much
elongated, and later flowers are always smaller. (B. L. R.)"
Wilcox's plant cited below was also determined by Dr. Robin-
son as A. saxosa. Both of these agree perfectly, however, with
Wright's specimens ; but neither with our material of A. lanuginosa
from the Southern States and Mexico, nor with the type of A.
saxosa. A. confusa grows in canons at an altitude of 2000-3500 m.

New Mexico: White Mountains, 1897, E. 0. Woofon, 2pj
(type); 185 1, C. Wright, 864; Burrow Mountains, 1880, H. H.
Rusby, j8.

Rydberg : Studies on the Rocky Mountain Flora 27G

Arizona: Ft. Huachuca, 1892, T. E. Wilcox ; Rincou Moun-
tains, 1891, Neally, iig ; Flagstaff, 1894,/. W. Tourney.

Colorado: La Plata Mountains, 1896, F. Tweedy, 4.26;
Wahatoya Creek, 1900, Rydberg & Vreeland, 6274; Pagosa
Peak, 1899, C. F. Baker, jog.

Delphinium ramosum sp. nov,

A tall perennial, often 2 m. high. Stem glabrous and shining
up to the inflorescence, often tinged purplish or bluish : petioles
1-1.5 dm. long; leaf-blade glabrous above, finely puberulent be-
neath, divided near the base into 5-7 segments, which are 4-7 cm.
long, oblanceolate or obovate, cuneate in outline, unequally 3-cleft
and these segments again cleft into oblong or lanceolate lobes 3-6
mm. broad : inflorescence with slender branches, these sparingly
and finely puberulent : bracts linear-subulate, 5-10 mm. long :
pedicels slender, 1-3 cm. long: bracdets filiform, 2-5 mm. long,
inserted 2-5 mm. below the calyx: sepals dark blue, striately
nerved, slightly brownish at the base, about i cm. long, ovate ; the
upper and lower ones acute ; the lateral ones obtuse, all puberulent :
spur about i cm. long, straight, or slightly curved at the apex :
upper petals dirty-white, tinged with brown and blue, about 8 mm.
long : spur slender, almost i cm. long : lateral petals dark blue,
bearded, with a slender claw, somewhat corniculate, bent at right
angles and only slightly 2-lobed at the apex: follicles 3, 12-15
mm. long, somewhat spreading and with divaricate beaks.

This species has without doubt been included in D. scopidorum,
but is evidently distinct from the type of that species. It is taller,
with more branched inflorescence, glabrous stem and the flowers
are usually tinged with brown at the base. The type of D. scopu-
loriim has narrow, very acute and divergent lobes of the leaves
and the calyx is purely dark blue. D. ramosum grows at an alti-
tude of 2000-3000 m.

Colorado : North Cheyenne Caiion, 1896, E. A. Bcssey{ty^€) ;
also in Williams Caiion, Artists Glen and other places near Pikes
Peak; "Colorado," I'i^c), Alice Eastivood.

Delphinium robustum sp. nov.

A tall and stout perennial, often 2 m. high or more. Stem
leafy, striate, puberulent throughout, often i cm. in diameter at
the base; petioles 1-2 dm. long, puberulent; leaf-blade finely and

277 Rydberg: Studies on the Rocky Mountain Flora

sparingly puberulent on both sides, divided to the base into 5-7
segments, which are 6—12 cm. long and twice cleft into linear lobes
3-5 mm. broad : inflorescence branched, densely many-flowered :
pedicels 1-3 cm. long, rather stout, usually equalled or surpassed
by the linear bracts : bractlets filiform, 5-10 mm. long, inserted
close under the calyx : sepals dark blue, paler at the base, ellipti-
cal, all except the upper one obtuse, 12-15 ^n"^- lo"g ; spur 12
mm. long, slightly curved : upper petals narrow, light brownish,
striate: lateral petals with slender claws bent at right angles,
bearded only within, deeply 2 -cleft at the apex.

This species in perhaps closest related to D. cncidlatiivi A.
Nelson but differs in the longer and narrower leaf-segments, the
more branched inflorescence, the stouter habit, the longer bracts
and bractlets and narrower upper petals. It grows at an altitude
of about 2500 m.

Colorado : Wahatoya Creek, below the Spanish Peaks, 1900,
Rydberg & Vreeland, 621 y (type); Colorado Springs, 1895, E. A.
Bessey ; Ruton, 1840, Albert.

Erysimum alpestre (Cockerell)

Erysijimrn aspenini f. alpestre Cockerell, Bull. Torr. Club, 18 :
168. 1891.

A tall strigose perennial with a deep tap-root ; stem strict, sim-
ple, 3-6 dm. high, striate : basal leaves linear or narrowly linear-
oblanceolate, 5-10 cm. long, 2-7 mm. wide, subentire or sinuately
denticulate, grayish strigose, gradually tapering below with a short
petiole : stem leaves mostly narrowly linear, the upper sessile :
raceme at first short and corymbiform, in fruit much elongated :
pedicels short, in fruit scarcely i cm. long, ascending: calyx 10-
1 2 mm. long, yellow : two of the sepals strongly saccate at the
base : petals with slender claws : blades broadly obovate-cuneate,
slightly emarginate, about 8 mm. long and broad, varying from
orange or brown to rose-purple or the older pale yellow : pods
four-angled, slender, erect, J-^ cm. long and 1.5 mm. in diameter :
style about 2 m.m. long, stout.

In general habit, this species closely resembles E. aspenun.
Mr. Cockerell, who seems to have been the first to observe this in-
teresting plant of the Sangre de Christo region, took it to be a form
of E. asperiini with peculiarly colored petals, found only at high
altitudes. I was first of the same opinion until I saw the fruit
in the type number and still better developed in Mr. Bessey's

Rydberg: Studies ox the Rockv Mountain Flora 278

specimens. This resembles more that oi E. asperriimini {Clieiran-
tlius aspcrrimmii Greene) than that of E. asperum. The present
species is also less densely and more finely strigose than either
of them. E. alpcstn grows at an altitude of 2500 m. or more.

Colorado: Headwaters of Pass Creek, i goo, Rydberg & Vrcc-
land, 6200 (type); mountain near Veta Pass, Srgg ; Veta Moun-
tain, 6igj ; tributary of Turkey Creek, 6ig8 ; East Indian
Creek, 6ig6 ; Ojo, 6202; Bald Mountain, near Pikes Peak,
1896, E. A. Bcsscy.

Draba aureiformis sp. no v.

A slender grayish stellate plant with perennial tap-root : stem
strict, 2-4 dm. high, simple or branched above, with slender erect
branches : basal leaves spatulate, 2-3 cm. long, acute, entire :
stem-leaves closely sessile, oblong-lanceolate, acute, entire or
sinuately denticulate : petals sulphur yellow, 3-4 mm. long,
broadly spatulate : pedicels ascending : pod erect, linear-oblong-
lanceolate, 10-15 mm. long, about 3 mm. wide, slightly if at all
twisted, stellate : style about i mm. long.

This species is nearest related to D. aiirca, but characterized
by the small light yellow petals, the slender style, the less dense
pubescence, and slender stem. It grows in dry soil at an altitude
of 2700-3 300 m.

Colorado : Headwaters of Pass Creek, 1900, Rydberg & Vree-
land, 61 j J (type); Middle Park, 1861, C. C. Parry, loj ; Gray-
mount, 1885, G. W. Letternian, 28.

Physaria vitulifera sp. nov.

A densely tufted finely stellate-pubescent perennial with deep
tap-root. Basal leaves numerous, fiddle-shaped, the larger 4—5
cm. long ; terminal lobe nearly orbicular to broadly obovate, sub-
entire, obtuse ; upper sinuses rounded: lateral lobes 1—2 pairs,
much smaller : stems ascending, or decumbent, 1—2 dm. high :
stem-leaves 1—2 cm. long, obovate or oblanceolate, entire : ped-
icels short, usually curved in fruit : sepals about 4 mm. long, ob-
long, acute : petals clawed, 8-9 mm. long : fruit obtuse at the
base, deeply divided above ; cells much inflated, round-obovate,
divergent, about 5 mm. in diameter : style about 5 mm. long.

This species, as well as the two following, differs from P. didy-
mocarpa in the smaller fruit, which is divided only above, not at

279 Rydberg: Studies on the Rocky Mountain Flora

the base. In this respect they resemble P. Ncwberryi ; but the
fruit is much smaller and not flattened on the sides. P. vitiilifera
differs from the following in the form of the leaves and the curved
pedicels. It grows in dry places at an altitude of about 2500 m.
Colorado : Idaho Springs, 1895, P. A. Rydberg {\.y^€)\ Middle
Park, 1 86 1, C. C. Parry, loi.

Physaria floribunda sp. no v.

Densely tufted perennial with thick tap-root. Flowering stems
very numerous, ascending or almost erect, 1-2.5 elm. high: basal
leaves less crowded and more erect than in the preceding, oblan-
ceolate or spatulate, sinuately toothed, acute, 5-10 cm. long :
stem-leaves oblanceolate, entire, about 3 cm. long: sepals about
5 mm. long, linear-lanceolate : petals bright yellow, 8-9 mm.
long, oblanceolate with broad claws : fruit obtuse or slightly cordate
at the base, deeply divided above : cells much inflated, about 8
mm. in diameter, almost globular : style 6 mm. long.

In the size and form of the fruit, this species is intermediate
between the preceding and P. didymocarpa ; but it is taller than
either. Sometimes the leaves are deeper sinuate, and then become
slightly fiddleform ; but the terminal lobe is sinuately toothed
and acutish. From P. didymocarpa it differs in the smaller fruit,
which is less cordate at the base, and the longer, more acute
leaves. It grows in loose sandy soil in mountain valleys at an
altitude of 2000-2700 m.

Colorado: Sangre de Christo Creek, 1900, Rydberg &
Vreelajid, 61 j^ (type) and 6ij6 ; Hills about Golden, 1892, Cran-
dall, 5J,

Physaria acutifolia sp. nov.

A small tufted perennial with a deep rather slender tap-root.
Basal leaves numerous, 1.5-2.5 cm. long, oblanceolate or obovate,
acute, entire or slightly wavy, very finely stellate : stem-leaves
rather few, oblanceolate : flowering stems 4-6 cm. long, ascending
or depressed : fruit small, obtuse or slightly cordate at the base,
deeply divided above : cells inflated, almost spherical, 4-6 mm. in
diameter : style about 5 mm. long.

This species differs from P. didymocarpa in the smaller size, in
the acute leaves, and the smaller fruit, which is more deeply divided
above and less so below. In general habit it resembles more P.
Geyeri, from which it differs in the turgid more spherical cells. It
grows in the mountains, reaching an altitude of 3000 m.

Rydberg : Studies on the Rocky Mountain Flora 280

Colorado : Grand Junction, 1893, Alice Eastzvood (type).
Wyoming: Mount Leidy, 1897, F. Tweedy, ^gi .

Cardamine cardiophylla sp. nov.

Stem stout, 3—10 dm. high, densely hirsute with short white
hairs, almost pilose : leaves rounded-cordate or the lower reniform,
simple, coarsely sinuately toothed : blades 3—4 cm. broad, more
or less pubescent : petioles more or less densely pilose, those of
the basal leaves 5—8 cm. long, those of the upper stem-leaves
very short : sepals also pubescent, about 3 mm. long : petals white,
broadly spatulate, about 8 mm. long : pedicels in fruit spreading,
1.5—2 cm. long: pods 2.5—3.5 cm. long, ascending, 1.5 mm. wide.

This species is nearest related to C. cordifolia, and differs
mainly in the thicker leaves and the dense pubescence. It reaches
an altitude of 3300 m.

Colorado : Tennessee Pass, Lake County, 1900, Geo. E.
Osterhont, 2i'/8 (type); 1893, De Alton Saunders ; Camp beyond
Bent's Fort, 1845, Fremont, ^2g.

Thlaspi Coloradense sp. nov.

Perennial ; rootstock or caudex cespitose, the branches rosu-
liferous at the apex. Basal leaves broadly spatulate, 1-2 cm. long,
fleshy, 4—7 mm. wide, sinuately crenate or subentire : flowering
stems 2-8 cm. long, leafy : stem-leaves oblong or obovate, 3-10
mm. long, obtuse : inflorescence short and dense, even in fruit
seldom over 2 cm. long : sepals rounded-ovate, obtuse, with a
white margin, about 2 mm. long : petals large, obovate-spatulate,
5-6 mm. long : silicle obovate, about 6 mm. long, only slightly
emarginate at the apex : wing-margins very narrow ; carina on the
sides rather prominent : seeds red, about i mm. long.

This species has been included by American authors in T. alpestre,

which is not found in America. The European species has very

small flowers and a different pod which is obcordate with a narrow

and deep sinus at the apex. Professor Greene has determined T.

Coloradense as T. Fendleri Gray. The latter was described from

flowering specimens and might be the present species ; at least

it is much like it. Specimens in fruit from the same region as the

type of T. Fendleri were collected later and were referred by

Dr. Gray to T. Fendleri ; in these the fruit is larger, decidedly

inverted triangular in outline, broader margined at the corners and

almost truncate at the apex.

281 Rydberg: Studies on the Rocky Mountain Flora

T. Coloradense groves at an altitude of 2500 m. or more, among
rocks.

Colorado : Bald Mountain near Pikes Peak, 1896, E. A.
Bessey (type) ; South Cheyenne Canon, Colorado Springs, 1900,
Rydberg & Vrccland, 6126 ; West Spanish Peak 61 2j and 6128 ;
Pikes Peak, 1894, E. A. Bessey ; Pikes Peak and Baldy, 1896,
F. Clements ; Pikes Peak and Tennessee Pass, 1893, De Alton
Saunders ; Laramie County, 1893, C. S. Crandall, 60 ; Pikes Peak,
1 89 1, Dr. E. Penard, 34. ; Arapahoe, 55 ; Los Pinos, 1899, C. F.
Baker ; Cheyenne Mountain and Seven Lakes, 1896, £". A. Bessey.

Thlaspi purpurascens sp. nov.

Perennial with a tap-root crowned by a very short caudex and
a rosette of leaves ; flowering stems generally several, less than
I dm. high : basal leaves oval or broadly spatulate, 2-3 cm. long,
petioled, usually more or less sinuate-dentate : stem-leaves ovate,
obtuse, with a truncate base, closely sessile: sepals 2.5-3 mm.
long, oblong-oval, obtuse, purplish with broad white margins :
petals broadly spatulate, about 6 mm. long : siHcle triangular-
obovate, distinctly winged above and with a very broad and shal-
low sinus at the apex, 7-8 mm. long.

This species is closely related to the preceding and inter-
mediate forms are not lacking. The principal difference is in the
fruit, which in T. purpurascens is comparatively broader, distinctly
winged and with a broad and open sinus at the top, almost trun-
cate and less keeled on the sides. The sepals in all specimens
seen are purplish and broader, the stem leaves are larger, more
ovate and usually with a truncate base, and the branches of the
caudex are very short.

Arizona : 1876, E. Palmer, ^ji (type) ; San Francisco Moun-
tains, 1 88 1, H. H. Rushy, 28; 1887,/?/-. E. A. Mearns, 34 ;
Flagstaff, 1898, i^. T. MacDo?/gal, 228.

Colorado: Headwaters of Sangre de Christo Creek, 1900,
Rydberg & Vreeland, 6123 ; Iron Mountain, 6124.

Sophia glandulifera sp. nov.

Biennial. Stem simple below, branched above, about 6 dm.
high, slightly grayish pubescent with branched hairs, decidedly
glandular-viscid above ; branches spreading, with upwardly curved
ends : leaves pubescent and viscid, bipinnately divided ; segments

Rvdberg: Studies on the Rocky Mountain Flora 282

linear-lanceolate or of the upper leaves filiform, acute or attenuate :
flowers light yellow : petals scarcely exceeding the sepals : pedicels
very short in flower ; in fruit 4-6 mm. long, ascending or almost
erect : pods 8-10 mm. long, erect, more or less curved, especially
those of the branches, scarcely exceeding 7 mm. in width, more or
less constricted between the seeds ; beak slender, short, about .5
mm. long : seeds uniserial, red, a little over i mm. long.

This species has the short pedicels and erect pods of 5. Hart-
ivegiana, but the pod of the latter is more slender, and often
curved, the whole plant is greener and conspicuously glandular,
and the se2:ments of the leaves are more slender. The latter char-
acter and the short ascending or erect pedicels distinguish it from
^. incisa Engelm. It grows at an altitude of 1000-1500 m.

Wyoming : Rolling plains between Sheridan and Buffalo, 1900,
F. Tivecdy, 35g2.

Sedum frigidum sp. nov.

Perennial with a fleshy rootstock, dioecious. Stems usually
less than i dm. high, light green : leaves flat, 1-1.5 cm. long and
5-7 mm. wide, sessile, obovate or oblong-obovate, often dentate
above the middle, or entire, acute : inflorescence dense, usually dark
purple : flowers 4— 5-merous, usually 5-merous : sepals of the
staminate flowers lanceolate, acute, 1.5-2 mm. long, dark purple or
rarely greenish : petals oblanceolate or oblong, acute, about 3 mm.
long, dark purple or very rarely greenish tinged with purple : fila-
ments filiform, purple, about one third longer than the petals, the
pistillate similar but with somewhat shorter and more obtuse petals :
follicles 3-5 mm. long, oblong, with a very short beak about .5
mm. long, divergent or at last recurved.

This species has gone under the name of ^. roseuvi (L.) Scop.,
but is quite unlike the northern European plant, which must be re-
garded as the type of RJiodiola rosea L. This has a very short,
almost tuberous rootstock, more oblanceolate leaves about 3 cm.
long, usually with very sharp dentations, usually yellow petals,
longer filaments almost twice as long as the petals and follicle^, 6-
8 mm. long. This form is also found in the mountains of southern
Europe ; but there seems to be another European plant ; this is de-
scribed and figured under the name RJiodiola rosea in the " Flora von
Deutschland" published by Schlechtendal, Langethal and Schenk.
It resembles more the Rocky Mountain plant in the purple,
flowers and short stamens, but it has more inversely deltoid leaves.

283 Rydberg: Studies on the Rocky Mountain Flora

the fertile flowers have very minute petals or none and the follicles
have long beaks. In all systematic botanies of Scandinavia
and Russia, S. rosciiui is described as having yellow petals, which
is never the case in the Rocky Mountain and Alaskan plants.
In this respect plants from eastern North America agree with the
European.

5. frigiduui is an alpine-arctic plant growing in Colorado at an
altitude of 3000 m. or more. In Montana it is found at an alti-
tude of about 2700 m. and in Alaska at low altitudes. It grows
among rocks, associating with several species of Saxifraga and
Adoxa Moschatellina.

Montana: Old Hollow Top, 1897, Ryciberg & Bessey, 4248
(type); Long Baldy, 1896, Flodman, jij ; Haystack Peak, 1899,
P. Koch.

Colorado: West Spanish Peak, 1900, Rydberg &■ Vreeland,
61 1 2 (9); 61 14 i^$) and 6115 (with greenish, merely purple-
tinged flowers) ; Grayback Mining Camp, 61 16 ; Pikes Peak,
1900, F. Clements ; 1821, Dr. James ; Arapahoe Peak, 1891, Dr.
E. Penard ; Pikes Peak, 1894, E. A. Bessey ; 1893, De Alton
Sainiders.

Nevada: Mineral King, Sierra, Nevada, 1891, Coville &
Funsioti, ij2p.

Idaho: Packsaddle Peak, 1892, Sandberg, MacDougal &

Heller, 860.

Washington: Mt. Paddo, 1886, VV. H. Snksdorf, 8sg.

Alaska: Muniak Island, \'^(^\, Jas. M. Macoim, 48; Shu-
magin, 1 871-1872, J/. W. Harrington; King Island, 1897, E.
A. McIlJiejiny, 8j.

Sedum polygamum sp. nov.
Perennial with a thick fleshy rootstock. Stem 1-3 dm. high :
leaves obovate or oblanceolate, acute, sessile, flat, entire or minutely-
denticulate, 1.5-2.5 cm. long: inflorescence dark purple, dense :
flowers dioecio-polygamous : sepals of the staminate ones lanceo-
late, 2 mm. long, acute : petals oblanceolate, dark purple, about 3
mm. long : filaments about half longer than the petals, purple,
broader than in the preceding species, abruptly acuminate above :
fertile flowers with more lanceolate petals and usually with sta-
mens which however have shorter filaments scarcely exceeding
the petals : follicles 6-8 mm. long with an ascending or spreading
beak, about i mm. long.

Rydberg: Studies on the Rocky Mountain Flora 284

This species is closely related to the preceding, but is stouter,
the fertile flowers usually with stamens, the follicles larger and
with less divaricate beaks. It has also been included in 5. roscum^
but is really nearer related to 5. atropiirpiirewn of eastern Asia,
which however has larger leaves. S. polyganiuvi grows at an alti-
tude of 3000 m. or more.

Colorado : West Spanish Peak, /poo, Rydberg & Vrecland,
61 1 3 (type); Silverton, 1895, F. Tti'ccdy 124; Mt. Hesperus,
I $98, Baker, Earlc & Tracy, 481 ; Basin Creek, La Plata Moun-
tains, 4S0 ; Mt. Lincoln, 1873, /. M. Coulter; Chambers Lake,
1894, C. S. Craiidail.

New Mexico: White Mountains, 1897, E. 0. Wooton, 677.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 15

STUDIES ON THE ROCKY MOUNTAIN

FLORA.-VI

BY P. A. RYDBERG

NEW YOEK
1901

[Reprinted from Bulletin Toeeky Botanical Club, 28 : m-5l3. 30 Sept. 1901.]

Studies on the Rocky Mountain Flora.— VI

By p. a. Rydberg

Salix padophylla n. n.

5. padifolia Rydb. Bull. Torr. Club, 28: 272. 1901. Not
Anderson.

In the May number of the Bulletin appeared a new species of
willow under the name 5. padifolia. The name should have been
S. padophylla, as S. padifolia is preoccupied by one of Anderson's
species. Besides, padiis is originally a Greek word, -aoo^.

Trifolium stenolobum sp. no\'.

A densely cespitose and scapose perennial. Branches of the
caudex covered by the large stipules which are 2 cm. long ; petioles
5-10 cm. long, finely strigose ; leaflets 3, narrowly lanceolate or
oblanceolate, 1.5-4 cm. long, finely strigose, very acute or acumi-
nate : scape 1-1.5 dm. long : heads many-flowered ; bracts minute ;
flowers reflexed in fruit : calyx-tube silky-strigose, 4-5 mm. long ;
teeth almost filiform, 7-9 mm. long: banner about 18 mm, long^

obtuse, mucronate.

This is related to T. dasypliyllum but is easily distinguished
by the minute bracts, the long slender calyx-teeth and the larger
corolla. It grows at an altitude of 3600 ni,

Colorado : La Plata Mountain, 1896, F. Tweedy, ^57 (type in
U. S. Nat. Herb.).

Trifolium subcaulescens A. Gray, Ives, Rep. Colo. Riv. Bot.

10. i860

Trifolium iicmor ale (jx&txvQ, Pittonia, 4 : 136. 1900.

Dr. Watson referred this species to T. gymnocarpuvi, to which
it is nearest related, but it differs in the larger size and the many-
flowered head. The specimen of C. F. Baker's no. 446 in the
herbarium of the New York Botanical Garden matches perfectly
Newberry's specimen in the Columbia University herbarium. The
former is the type number of T. neinorale Greene, and the latter
that of T. siibcaidescens A. Gray. Unless the type sheets repre-

499

500 Rydberg : Studies on the Rocky Mountain Flora

sent different species from those represented in our herbaria, Prof.
Greene's name has to pass into synonymy.

Trifolium bracteolatum

Trifolium lilaciniwi Rydb. Bull. Torn Club, 28: 37. 1901.
Not Greene. 1896.

Vicia producta sp. no v.

A very slender, sparingly hairy cespitose perennial. Stems
decumbent or ascending, 2-3 dm. long, much branched and leafy,
striate and somewhat angled : stipules narrow, semi-hastate, entire,
3-5 mm. long; leaflets 3-5 pairs, oblong to linear, 5-15 mm.
long, obtuse, mucronate ; tendrils 3-cleft : peduncles 2-4 cm.
long, usually 2-flowered, produced beyond the upper flower : calyx-
tube about 2 mm. long, strigose, teeth about i mm. long, lance-
olate-subulate : corolla about 8 mm. long, yellowish-white, tipped
and tinged with purple.

This species is probably closest related to V. lutmilis H. B. K.,
but is characterized by having the peduncle produced beyond the
two flowers and by different calyx-lobes. The type was growing
among rocks on the south side of a butte, at an altitude of about
2400 m.

Colorado : Butte, 5 miles southwest of La Veta, 1900, Ryd-
berg & Vre eland, 6006.

Primula Americana sp. nov.

Primula farinosa A. Gray, Syn. Fl. 2^: 58, in part. 1878.
Not L. 1753.

Perennial with a short rootstock and fleshy-fibrous roots.
Basal leaves oblong or oblong-oblanceolate or spatulate, 2-8 cm.
long, obtuse, gradually contracted at the base, usually with short
winged petioles, sinuate dentate above the middle or subentire,
thin, more or less mealy, especially on the lower surface : scape
1-2 dm. high, mealy when young: bracts 6-10 mm. long, linear-
lanceolate, usually acute : pedicels in flower little if any exceeding
the bracts, in fruit sometimes 2 cm. long, erect from the beginning :
calyx more or less mealy, 6-8 mm. long, lobes oblong-obtuse :
corolla lilac ; tube 8-9 mm. long, only slightly exceeding the
calyx ; lobes of the corolla obcordate, 2-3 mm. long.

This species has been confused with P. farinosa, but I think it
distinct. All specimens from the Rocky Mountain region under
that name differ from the European and especially the Scandi-

Rydberg : Studies on the Rocky Mountain Flora 501

navian P. farinosa in the longer bracts, in the longer calyx and
in the short corolla lobes. The specimens of northeastern America
seem to be like the European plant. In the latter the bracts arc-
only 4-6 mm. long, acuminate, the calyx less than three fourths
the length of the tube of the corolla and the lobes of the latter
4-5 mm. long. /'. Americana ^gxows, in swamps and wet meadows
of the Rocky Mountain region at an altitude of 1200-2500 m.

Montana: Deer Lodge, 1895, P. A. Rydberg, 2746 (type).

Wyoming: Little Laramie River, \d>C}6, Avcn Xclson, ig6i ;
Hams Fork and La Barge, 1900, C. C. Curtis.

Alberta: Devil's Head Lake, Banff, 1899, \V. C. McCalla,

2.f.22.

Colorado: North Park, 1896, G:o. E. OstcrJiout.

Cuscuta gracilis sp. nov.

Stem filiform, about .25 mm. in diameter. Flowers in dense
globular clusters : calyx gamoscpalous but cleft to near the
base, lobes ovate : corolla urceolate, less than 2 mm. high, lobes
ovate, widely spreading, acute, delicate, about i mm. long ; scales
ovate, crenate, not divided, almost half as long as the corolla-tube :
filaments subulate, about twice as long as the anthers : styles dis-
tinct, equal, about as long as and somewhat thicker than the red
filiform curved stigmas : capsule about 2 mm. high, acute-globose,
circumscissile near the base : seeds about i mm. long.

This species is nearest related to C. epili/unn, which, however,
has shorter and broader corolla-lobes, shorter filaments, scarcely
longer than the anthers and short emarginate and crenate scales.
C. gracilis grows parasitic on species of Erigeron, CJirysotliamnns
and Solamim.

Wyoming: Rolling plains between Sheridan and Buffalo, 1900,
F. Tweedy, 32^2 (type) ; Laramie, 1894, Aven Nelson, iijg.

Cuscuta megalocarpa sp. nov.

Stem stout, usually over i mm. in diameter. Flowers in dense
globular clusters : pedicels very short, at most 2 mm. long :
calyx gamosepalous, lobes rounded, scarcely i mm. long : corolla
about 3 mm. high and broad, lobes broadly triangular, acutish,
about I mm. long, with incurved tip ; fringed scales attached near
the bottom, equalling about half the corolla-tube, deeply 2-lobed,
and fringed only in the open sinuses : stamens about as long as the
lobes of the corolla, filaments subulate, about twice as long as the

502 Rydberg : Studies on the Rocky Mountain Flora

rounded anther : styles distinct, short, stigmas capitate : capsule
5-6 mm. in diameter, acute-globose, about 4-seeded : seeds about
2.5 mm. long and 2 mm. broad, finely muricate.

A plant parasitic on willows and other shrubs, at an altitude of
about 2000 m.

Colorado : Cucharas Creek, near La Veta, 1900, F. K.
Vre eland, 6jo.

Wyoming: Dayton, Sheridan Co., 1899, F. Tioccdy, 22j8.

Monarda comata sp. nov.

Perennial with a horizontal, slender rootstock. Stem obtusely
4-angled, 4-6 dm. high, simple, sparingly silky-villous, especially
below the nodes and on the upper portion : lower leaves with more
or less hairy petioles, which are about 1 cm. long, the upper sub-
sessile ; blades ovate or lance-ovate, more or less cordate at the
base, acute or acuminate, serrate with small teeth which are di-
rected forward, 4-6 cm. long, sparingly silky-strigose on both
sides: bracts green or tinged with purplish, ovate, 1-2 cm. long:
calyx about i cm. long, minutely puberulent, with a ring of hairs
at the base of the teeth, which are subulate, .7 mm. long: corolla
red-purple, almost wine-color, densely villose-puberulent, the ex-
serted portion about 2 cm. long.

This is nearest related to M. stricta Wooton, but differs in the
long pubescence of the stem, leaves and bracts, shorter calyx-teeth
and darker flowers. It grows in meadows among bushes, at ah
altitude of 20QO-2500 m.

Colorado : Wahatoya Creek, 1900, Rydberg & Vreela^nd, jdjj
(type); Ruxton, i goo, Fred Clements.

Castilleja Wyomingensis sp. nov.

A more or less cespitose perennial with a short woody caudex.
Stems 3-4 dm. high, striate, sparingly puberulent and the upper
portion viscid-pubescent : leaves linear, 4-5 cm. long, 4-7 mm.
wide, minutely puberulent, more or less distinctly 3-nerved, entire,
or the upper 3-lobed : bracts lanceolate to ovate, puberulent and
ciliate on the margin and veins tinged with sulphur-yellow, 3-
lobed at the apex or the lower entire, middle lobe oblong and
rounded at the apex, the lateral ones lanceolate or linear, acute :
calyx villose, nearly 2 cm. long and almost equalling the corolla,
sulphur-yellow, cleft about half way down, slightly deeper on
the lower than on the upper side, lateral cleft 3-4 mm. deep :
corolla greenish, tinged and bordered with sulphur-yellow, galea
about 6 mm. long, lip about 2 mm.: capsule ovoid, 12-13 mm. long.

RvDBERG : Studies on the Rocky Mountain Flora 503

The species is nearest related to C. sulp/iuna, from which it
differs in the narrow leaves and in being more puberulent. It
grows in northern Wyoming at an altitude of 2200-2700 m.

Wyoming: Big Horn Mountains, Sheridan Co., 1899, F.
Tweed}', 2j;^i (t\'pe) and 2J42 ; Headwaters of Clear Creek and
Crazy Woman River, 1900, Tzveedy, J400.

Pentstemon suffrutescens sp. nov.

PcntstcDwn cacspitosns var. suffntiicosus A. Gray, Syn. Fl. 2' :
270. 1878. ^oX. P. suffrnticosiis V)o\x^. 1846.

Excellent specimens of what I take to be Gray's variety cited
above, which was described from imperfect material, have been
collected by Mr. Tweedy, and these show that the plant is nearer
related to P. pimiilus than to P. cacspitosns, but the calyx lobes
are broader, shorter and blunter than in that species, the corolla
slightly smaller and the leaves glabrous.

Colorado : Ridgway, 1895, F. Tweedy, ijo.

Sambucus microbotrys sp. nov.

A low shrub, 5-20 dm. high, glabrous throughout and with
pale green foliage. Leaflets ovate or rarely ovate-lanceolate, acute
or short-acuminate, 3-9 cm. long, mostly rounded and oblique at
the base, coarsely serrate : cyme thyrsoid-paniculate, small, about
as long as broad, about 3 cm. in diameter and of the same height ;
flowers whitish : fruit bright red, 4-5 mm. in diameter : seeds
finely punctate-rugose.

This species is nearest related to 5. pnbcns and perhaps all
specimens from the southern Rockies referred to that species
belong here. The main distinctions from 5. pubens are the total
absence of pubescence and the smaller inflorescence. It grows
at an altitude of 1500-2700 m.

Colorado : Bottomless Pit, and below Halfway House, Pike's
Peak, 1896, E. A. Bcsscy (type); Gray's Peak, 1895, P. A.
Rydberg ; East Indian Creek, 1 900, Rydberg & Besscy, ^^82 ;
Little Veta Mountain, JJc^J.

Chrysothamnus filifolius sp. nov.

A small shrub, about 3 dm. high, very bushy. Bark of the
stems and larger branches gray and somewhat fissured, that of
the young branches white and shining: leaves very narrow, linear-

504 Rydberg : Studies on the Rocky Mountain Flora

filiform, 1-2 cm. long, less than i mm. wide, dark green and
glabrous except the minutely scabrous-ciliolate margins : cyme
large, flat-topped : heads very' numerous, 5-6 mm. high, 2 mm.
in diameter, turbinate ; bracts in 5 vertical rows with usually 3 in
each row, lanceolate-acuminate, the outer with green tips, the
inner wholly chartaceous.

In habit and leaves, this species resembles C. stcuopJiylliis, but
the leaves are still narrower and in that species the bracts are
obtuse or merely acute.

Colorado: Granite, I'&gG, Fi-cd. Clements, jgo.

Chrysothamnus scoparius sp. nov.

Perennial with a woody caudex and deep tap-root. Stems 1-2
dm. hicfh, elabrous and shining, with a white bark : leaves nar-
rowly linear, light bluish-green, glabrous, 2-3 cm. long, a little
over I mm. wide, more or less twisted : cyme flat-topped : heads
numerous, 6-8 mm. high, 2-3 mm. wide ; bracts chartaceous,
lanceolate, acuminate.

This species is still more like C. stenopliylhis in habit, but the
bracts are diiTerent, being acuminate instead of obtuse or acute
and more keeled. The perennial portion is also much shorter and
the general color of the plant is much lighter.

Colorado: Mesas, La Veta, 1900, F. K. Vreeland, 6g8.

Aster crassulus sp. nov.

Aster midtiflorus A. Gray, PI. Wr. 2 : 75. In part. 1853.

Aster incano-pilosits Rydb. Mem. N. Y. Bot. Gard. i : 393.
In part. 1900.

Perennial with a horizontal, stoloniferous rootstock. Stems
erect or ascending or rarely decumbent, usually simple below,
branched above, 4-7 dm. high, terete, brownish, densely hirsute
with short divaricate hairs : stem-leaves linear, or oblong-linear, 3-
6 cm. long, 2-3 mm. wide, rather firm, hirsute with short spreading
hairs, sessile and slightly clasping, in age usually reflexed, acute
with a short spinulose tip ; those of the branches smaller ; those
of the branchlets only 3-5 mm. long and merging into the bracts
of the involucres : heads numerous, borne at the end of leafy
branchlets which are 1-2 cm. long ; involucre turbinate, 5-8 mm.
high, 8-10 mm. broad ; bracts in 3-4 series, the outer successively
shorter, broadly spatulate, minutely spinulose-mucronate, very
thick, hirsute especially on the margin, chartaceous at the base
with a broadly oval herbaceous tip : rays numerous, white, 5-7
mm. long and fully i mm. wide : achenes strigose.

Rydberg : Studies on the Rocky Mountain Flora 505

This is a member of the A. uiultiflonis group and has been
confused with A. commntahis (T. & G.) Gray or A. incano-pilosiis
Sheldon. In A. commiitatiis, the stem is strigose-pubescent, the
branchlets more elongated and the outer bracts longer than in A-
crassidiis and often equalling or surpassing the inner bracts. It
may be the same as described by Lindley in Hooker's Flora and
DeCandoUe's Prodromus as " A. ramnlosus (i, incano-pilosus " ;
but here it is to be noticed that the word " incano-pilosus " is
printed in the same type as all the descriptions and not the special
type used for names. Hence "incano-pilosus" was a descriptive
term for the A. ramnlosus ,5, and not the name. If. however,
it should be taken as a name, which Mr. Sheldon did, it is a
nometi muiuni, as no further description is given, and then should
be disregarded. As Sheldon's Aster incano-pilosus, which is amply
distinct, has as a synonym, the older A. comniutatus (T. & G.)
Gray it must be dropped and A. conunutatus substituted.

A. crassulus grows on dry plains and table-land at an altitude
of 1 200 m.

Colorado : Mesas, La Veta, 1900, F. K. Vrceland, 6goa (type)
and 6c)0\ Denver, 1871, Dr. Geo. Smith, Sj ; Ridgway, 1894, F.
Tweedy, J 06 ; Pagosa Spring, 1899, C. F. Baker, Sj-/.

New Mexico : 185 i, (f. Wright, 1155; Rio Dolores, AVw^mj.

California: Valley, near San FiHpe, 1858, S.Hayes.

Wyoming: Bear Lodge Mountains, 1898, L. W. Carter;
Moorcroft, 1897, Z. W. Carter; Dayton, 1899, F. Tweedy, 2031.

North Dakota: Minot, 1891, Wright; Custer, 1892,7?/^/-

bcrg, 773 .

Montana: Park Co., 1887, F. Tweedy, 339 ; Colgate, Sand-
berg, MacDougal %i Heller, 1021 ; Montana Valley, Madison Co.,
1899, Aven & Elias Nelson, 683c).

Aster exiguus (Fernald)
Aster ciliatus Muhl. ; WiUd. Sp. PI. 3: 2027. 1804. Not
Walt. 1788.

Aster hebecladus K. Gray, Syn. Fl. 2' : 185, under ^. nmltiflorus.

Not DC.

Aster multijlorus var. exiguus Fernald, Rhodora, i : 187. 1899.
I think that Aster ciliatus Muhl. is distinct from A. nmltiflorus ;

506 Rydberg : Studies on the Rocky Mountain Flora

but unfortunately it is antedated by A. ci7ia/?is Walt. In A. iiuilti-
flonis the pubescence is mostly appressed and the bracts are nar-
row-oblanceolate and rather thin. In A. cxigmis the pubescence is
much denser and divaricate and the bracts are thick, broadly spatu-
late or obovate. In fact A. exigiiiis differs from the preceding only in
the smaller, more crowded heads and the shorter rays ; the former
are less than 5 mm. high and the rays are only 5—6 mm. long. The
New Mexican form which Gray regarded as A. liebecladiis DC. and
for which Dr. Greene has revived the name belongs here, but is
evidently not A. liebccladus DC. for this is described as having
glabrous stem and linear bracts.

A. exiguus grows on prairies and plains along roads, etc., and
westward is much more common than A. mnltiflorus. It ranges
from Vermont and Pennsylvania to Washington and south to
Texas and Arizona.

Machaeranthera rubricaulis sp. nov.

Biennial or perhaps perennial. Stem erect or nearly so, simple
below, branched above, 4-6 dm. high, terete, striate, usually tinged
with red or purple, finely strigose-puberulent, not viscid : leaves
linear or linear-oblanceolate, sessile, acute, entire or sparingly
" dentate, scabrous-ciliolate on the margins ; the largest 8 cm. long,
and 8 mm. wide : heads corymbose-cymose, numerous : involucre
10-12 mm. high, 12-15 mm. in diameter : bracts numerous, linear-
lanceolate with attenuate, almost terete squarrose tips, viscid-
puberulent : rays numerous, dark blue, about 1 2 mm. long and i
mm. wide : pappus sordid : achenes flat, sparingly and minutely
strigose.

The species grows in dry soil at an altitude of 2000— 2700 m.

Colorado : Mesas, La Veta, 1900, F. K. Vrceland, 681 (type) ;

Tel lu ride, 1894, F. Tzucedy, jo^ and j 05.

Eiigeron laetevirens sp. nov.

Perennial with a woody tap-root. Stems several, 1.5-2 dm.
high, slender, erect or ascending, striate, silky strigose : basal
leaves very narrowly linear-oblanceolate, acute, 5- 10 cm. long,
1—3 mm. wade, sparingly strigose ; stem-leaves narrowly linear,
2—6 cm. long, scarcely over i mm. wide : heads solitary : invo-
lucre 12—15 nim. in diameter, grayish villous-hirsute : bracts
in 2—3 series, subequal, linear-lanceolate, long-acuminate : rays
numerous, blue or purple, y-^ mm. long, 1-1.5 mm. wide.

Rydberg : Studies on the Rocky Mountain Flora 507

This species is nearest related to E. Montainiisis, from which it
differs in the narrower, blue or purple rays and longer and less
pubescent leaves. It grows at an altitude of 2000-2700 m.

Wyoming : Little Goose Creek, 1899, F. T-wecdy, 200 j; (type) ;
Headwaters of Clear Creek and Crazy Woman River, 1900, ,^0/0.

Carduus Centaureae sp. nov.

(?) Cnicus carlinoides var. Amcricanus A. Gray, Proc. Am.
Acad. 10: 48. 1874. Not Cirsiuin acaidc var. Amcricanus
A. Gray. 1863.

(?) Cnicus Amcricanus A. Gray, Proc. Am. Acad. 19 : 56.

Biennial or perennial. Stem rather slender, 6-10 cm. high,
striate, often tinged with red, slightl)' floccose when young : basal
leaves about 2 dm. long, thin-petioled, deeply pinnatifid to near
the midrib, above light green, sparingly floccose when young,
glabrate in age, beneath more or less permanently grayish-to-
mentose ; lobes lanceolate or triangular, 2-4 cm. long, more or
less toothed and tipped with weak spines, 1-3 mm. long ; lower
stem-leaves similar but petiole more winged, dilated at the base
and short decurrent ; the upper sessile and clasping with an
auricled base ; the uppermost lanceolate and undivided : heads
2-2.5 cm. high and 1.5-2 cm. in diameter : outer bracts narrowly
lanceolate, yellowish, often with darker center, fimbriate on the
margin and tipped with a flat weak spine 1-2 mm. long : the
inner with dilated deltoid scarious fimbriate tips which are merely
acuminate : flowers ochroleucous.

This species may be the Cnicus Amcricanus A. Gray or Carduus
Americamis Greene, but the name Amcricanus is preoccupied. The
species is, however, not nearest related to C. Parryi, where Dr.
Gray placed it ; but to C. leioccphalus and C. canovirens, from
which it differs in the less spinose, more dilated and fimbriate
bracts. A seedling of apparently this species was collected by
Cowen and it has long oblanceolate 3-4 dm. long, undivided
leaves. It grows at an altitude of 2200-2600 m.

Colorado : Laramie County, 1895,/. H. CozvcniX-y^^) ; 4-mile

Hill, Routt County, 1896, C. F. Baker; Penn's Gulch, 1865,

Letterman, yS.

Carduus erosus sp. nov.

Biennial. Stem stout, about 7 dm. high, angled and striate,
often red, loosely floccose when young : lower leaves about 2 dm.
long, rather thin, green and slightly floccose above when young :

608 Rydberg: Studies on the Rocky Mountain Flora

grayish-tomentose beneath, deeply pinnatifid to about one fourth
from the midrib : lobes about 2 cm. long, lanceolate, few toothed or
lobed and tipped with moderately strong spines 4-10 mm. long :
heads hemispherical, 2.5-3 cm. high and 2.5-3.5 cm. broad;
outer bracts ovate, slightly floccose when young, entire, with a
narrow dorsal line, tipped with weak spines 1-2 mm. long, the
inner with dilated deltoid erose scarious acuminate tips : flowers
ochroleucous.

A species closely related to the preceding, differing in the
larger heads, longer spines, broader bracts and with erose instead
of fimbriate tips. It grows in moist meadows at an altitude of
about 2200 m.

Colorado: Durango, 1896, Frank Tzvccdy, 317 {ty^Q in U.

S. Nat. Herb.).

Carduus Americanus (A. Gray).
Cirsmm aciile var. Americanus A. Gray, Proc. Acad. Phila.

1863 : 68.

Cm'cus Drummondii acaulcscciis A. Gray, Proc. Am. Acad. 10 :

40, in part.

Dr. Gray's Cirsinm acanle var. Anicricamivi is amply distinct
from C. Drummondii not only by the acaulescent habit, but also
by the narrower bracts and their long stout spines which are 1-2
cm. long while in C. Drummondii they are only a few mm. in
length and very weak. It is an alpine species growing at an alti-
tude of 2800-3500 m. To this belong the following specimens.

Colorado : Silver Plume, 1895, P. A. Rydberg ; Como, South
Park, 1895, C. S. Crandall ; South Park, 1884, Letterman.

Carduus acaulescens (A. Gray)
Cnicus Drummondii var. acaulescens A. Gray, Proc. Am. Acad.

10 : 40, in part.

Gray's variety acaulescens was based on two previously pub-
lished things, viz., Cirsinm acaule var. Americanus A. Gray, given
above, and Cirsinm Driimmondii, " acaulescent form " D. C. Eaton
in King's Report. As the first already has a valid name, I here
adopt the name acaulescens for the other part on which the variety
was founded. C. acaidesccns resembles closely C Americanus in
habit, but the spines of the outer and middle bracts are shorter and

Rydberg : Studies on the Rockv Mountain Flora 509

weaker, seldom over 5 mm. long, and the inner are acuminate,
wholly unarmed. The true C. Driiininondii, even in its subacaul-
escent state, has very broadly ovate outer bracts and the spine is
so fine that it is better called a bristle. The inner bracts have a
long acuminate very crisp tip. In C. acan/csccns it is scarcely
crisped. The following specimens belong to C. acmi/csccits :

Utah : Bear River Valley, 1869, 5. Watson, 6po.

Wyoming : Woods Creek, 1897, A^l-// Nelson, j-^Sj.

Colorado: Grizzly Creek, 1896, C. F. Baker.

Carduus oreophilus sp. no v.
Cirsinm Dnuninondii, D. C. Eaton, King's Rep. 5: 195. 1871.

Not T. and G.

Biennial stem 4-8 dm. high stout, angled and striate, slightly
arachnoid-hairy when young, leafy : basal leaves about 2 dm.
long, thin, short-petioled, green and glabrate above, more or less
grayish-tomentose beneath, pinnately lobed a little more than
half way to the midrib ; lobes triangular, usually 2-3 -lobed, with
rather slender spines 3-10 mm. long: heads hemispherical, 2-2.5
cm. high, 2.5 cm. broad: outer and middle bracts narrowly lan-
ceolate, slightly arachnoid, tipped with flat spines 3-6 mm. long,
the inner with a long attenuate soft tip : flowers red.

This species has been included in C. Druvimondii but is taller
with smaller head, narrower bracts, the outer with longer spines,
and the tips of the inner scarcely crisp. It grows at an altitude of
I 500-3000 m.

Colorado : Georgetown, 1895, P. A. Rydberg (type) ; Pagosa
Springs, 1899, C. F. Baker, 644.

Nevada: Run Valley, 1868, J5. Watson, 68g.

Carduus griseus sp. nov.

Biennial. Stem stout, sparingly floccose when young, angled
and striate : basal leaves oblanceolate in outline, thin, glabrate
above, grayish-tomentose beneath, deeply pinnately divided ; seg-
ments 3-4 cm. long, deeply 2-3 -lobed and toothed, tipped and
margined with rather slender spines 3-5 mm. long ; the upper
similar but less deeply divided, sessile and half clasping by the
dilated rounded shortly decurrent base : heads about 3 cm. high
and broad : bracts subequal, without dorsal glandular ridge, the
outer narrowly lanceolate, with a long-attenuate tip gradually
changing into a flat spine, and somewhat laciniate or spinulose-

510 Rydberg : Studies on the Rocky Mountain Flora

dentate on the margin ; the inner bracts less rigid, their tips not
spinose but long-attenuate, scarious and somewhat crisp : flowers
ochroleucous.

This species is probably nearest related to C. Iciocepliahis (D. C.
Eaton) Heller but differs in the less divided, less spinose leaves and
the bracts which are lacking the long yellow spines on the margins.
C. Iciocepliabis has red flowers. Cardials griscus grows at an alti-
tude of 3400 m.

Colorado: Telluride, i<S94, F. Tzuccdj, j2i (type in U. S.
Nat. Herb.).

Carduus pulcherrimus sp. nov.

Biennial. Stem about 4 dm. high, very leafy, grayish-tomentose,
especially when young, angled : lower leaves 1-1.5 dm. long, ob-
lanceolate in outline, short-petioled, pinnately divided, bright green
and glabrate above, densely white-tomentose beneath ; segments
lanceolate or triangular, lobed and toothed with slender spines 3—5
mm. long ; the upper leaves similar, sessile and half clasping : heads
2.5—3 cm. high, 2.15—4 cm. broad : outer bracts lanceolate with a
narrow glandular ridge, slightly floccose, tipped with yellow spines
about 5—7 mm long ; the innermost narrowly lanceolate and with
long-attenuate tips : flowers rose-colored.

In habit this species resembles most vS. nndulatits and vS.
odiroccntnis, but has smaller heads, narrower bracts and narrower
glandular ridge. It is perhaps intermediate between those species
and C. canovircns, from which latter it is easily distinguished by
the dense tomentum on the lower surface of the leaves. It grows
in open woods, etc., at an altitude of about 2500 m.

Wyoming : Headwaters of Clear Creek and Crazy Woman
River, 1900, F. Tiucedy, 30-f.S (type); Medicine Bow, 1898,
Elias Nelson, 43 g6.

Carduus oblanceolatus sp. nov.

Apparently perennial. Stem rather slender, 3—6 dm. high, al-
most terete, densely white-tomentose : lower leaves about i dm.
long, oblanceolate, thick, short-petioled, merely spinulose-dentate
or rarely with a few triangular lobes, loosely floccose above,
densely white-tomentose beneath, spines slender, 2-5 mm. long ;
upper leaves lanceolate, sessile and half clasping : heads 1—3, cam-
panulate, 3-3.5 cm. high, 1.5-2.5 cm. broad: bracts slightly floc-
cose at first, with a narrow glandular dorsal ridge, the outer ovate-
lanceolate, the middle lanceolate, with a slender divergent spine^

Rydberg : Studies ox the Rockv Mountain Flora 511

3-5 mm, long, the innermost long-attenuate, unarmed : flowers
rose-colored.

This species is nearest related to C. Flodmanii, differing in the
more simple habit, the smaller, thicker, not pinnatifid leaves and
shorter and weaker spines. From C. a/fissinins it differs in the
denser tomentum, the simple habit, the narrower and fewer bracts
and their narrower dorsal ridges.

Colorado: Twin Lakes, 1896, Fred. Clements, j8j (type).

Agoseris arachnoidea sp. nov.

Perennial with a tap-root, more or less floccose throughout.
Leaves 1.5-2 dm. long, more or less runcinate-lobed, oblanceolate
in outline, long-attenuate at the apex, densely floccose when young,
more glabrate in age : scape 1.5-4 dm. high, densely woolly above,
less so below: head 2-2.5 cm. high, 1.5-2 cm. broad: bracts in
about 3 series, linear-lanceolate, long-attenuate, the inner one-third
longer than the outer : flowers at least in age rose-purple : beak
of the achenes fully as long as the body.

Perhaps nearest related to A. aurantiaca but differing in the
larger heads, the dense pubescence and the lighter flowers. It
grows in meadows at an altitude of 2000-2800 m.

Colorado: Gray-Back Mining Camp, 1900, Rydberg &
Vreeland, S553 (type) ; La Veta, 1896, Fred. Clements, 167.

Agoseris pubescens sp. nov.

Perennial with a tap-root and short branching caudex. Leaves
narrowly oblanceolate, about i dm. long, .5-1 cm. wide, acuminate,
slightly glaucous but even in age villous-pubescent : scape 1.5-2
dm. high, sparingly villous: involucre about 2 cm. high and 1.5
cm. wide, villous and somewhat viscid : bracts with a dark
median line and sometimes tinged with purple, the outer ones
ovate-lanceolate, about 2^ the length of the linear-lanceolate
long-acuminate inner ones : flowers lemon-yellow, the outer stri-
ate or tinged with purple : beaks of the achenes short and striate
throughout.

This species is nearest related to A. glauca and A. scorzonerae-
folia but differs in being very pubescent. It grows in moist
meadows at an altitude of 2000-2500 m.

Wyoming: Big Horn Mountains, Sheridan County, 1899, F.
Tiveedy, 20 J j (type) ; Buffalo Fork, i8gy,j2S; Amethyst Creek,
1899, Az'en & hlias Nelson, ^ydg (in part).

512 Rydberg : Studies on the Rocky Mountain Flora

Taraxacum Alaskanum sp. nov.

A low delicate plant with perennial root : leaves linear-oblan-
ceolate in outline, 3-5 cm. long, deeply runcinate-pinnatifid with
triangular retrorse lobes : scape 4-5 cm. high : involucre cam-
panuate, 10-12 mm. high, 6-8 mm. broad: bracts fuscous, not
corniculate ; the inner linear-lanceolate, long-acuminate ; the outer
scarcely half as long, lanceolate, spreading or somewhat reflexed :
achenes brownish, spinulose-muricate above, fusiform, 4 mm. long.

This in size and habit most resembles T. scopidomm, but is
easily distinguished by its triangular acute lobes. It was found
" on steep side bank facing the ocean."

Alaska : Pt. Barrow, 1 898, McIUicnny.

Crepis glaucella sp. nov.

Perennial with a slender tap-root. Stem slender, about 3 dm.
high, glabrous and shining : basal leaves glabrous and shining,
somewhat glaucous, thin, 5-10 cm. long; blades oblanceolate,
acute at the apex, tapering downward into a more or less winged
petiole, remotely sinuate-dentate with divaricate or retrorse short
teeth or entire ; stem-leaves 1-2, much reduced, 1-2 cm. long,
linear or nearly so : heads 1-4, on slender branches : involucre
turbinate, about i cm. high and 6-8 mm. wide, sparingly hirsute ;
bracts linear-lanceolate, acuminate : achenes brown, lo-striate, gla-
brous : pappus white.

This species is nearest related to C. glaiica but differs in the
smaller, narrower, longer-petioled leaves and hirsute involucre.

Wyoming: Pacific Creek, 1897, F. Tweedy, 6oj.

HiERACiUM MACRANTHUM Nutt. Trans. Am. Phil. Soc. II. 7 : 446.

1840

Hicj'aciiini iniibellatwii Hook. Fl. Bor. Am. I : 300, in part ;
Rydb. Mem. N. Y. Bot. Garden i : 463, mainly. Not L.

The plant of the Rocky Mountain region which has been re-
ferred to H. iimbcllatwn and sometimes to H. Canadeiuc is quite
distinct from both. The leaves are usually broader than in the
first but narrower than in the second and the heads are usually
fewer than in either. The best character by which to distinguish
it is, however, the pubescence. The leaves are densely puberu-
lent and decidedly scabrous-ciliolate on the margins. The upper
portion of the stem and the branches of the inflorescence are also
scabrous puberulent.

RvDRERG : Studies ox the R(ickv Mountain Flora 513

//. iiiacranthuin ranges from Wyoming and Washington north-
ward to subarctic America. It apparently also extends eastward
to the upper Mississippi River (Nicollet's specimens seem to be-
long here) and therefore may be Schweinitz' //. scabcrrimniii,
which has been referred to H. iiinbcllatuiii and //. Canadcnsc. I
have not seen Schweinitz' original description and can not venture
an opinion. Nuttall's name belongs to the plant without an)'

doubt.

Hieracium Columbianum sp. nov.

Perennial. Stem about 6 dm. high, terete, more or less tinged
with purple, more or less white- or yellowish-hirsute below, gla-
brous or puberulent above : lower leaves oblanceolate, 8— lo cm.
long, light green and somewhat glaucous beneath, usually more
or less silky-hirsute, sinuately dentate, acute ; the middle leaves
lanceolate or ovate-lanceolate with rounded or obtuse base, sessile,
more glabrate ; the uppermost much reduced and bract-like : in-
florescence corymbose-C)'mose, small and contracted : heads 2—6,
12—15 nim. high : bracts lanceolate, unequal and more or less im-
bricated, dark, puberulent when young, glabrous in age : pappus
very light brownish.

This species is nearly related to H. Canadeiisc and H. iivibcl-
latiini, but differs from both in the long hairs of the lower part of
the plant. The leaves are thin as in H. Canadcnsc but narrower
and the heads are fewer. It grows in low ground at an altitude
of about 600 m.

Idaho : Priest River Valley, 1900, D. T. MacDoiigal, log
(type); cafions near Farmington Landing, \Z<^2, Sandberg, Mac-
Doug al & Heller, jjj.

CONTRIBUTIONS FROM THE NEAV YORK BOTANICAL

GARDEN— No. 16

VANISHING WILD FLOWERS

BY ELIZABETH G. BRITTON

NEW YORK
1901

[Reprinted fiom Torreya, Vol. I, No. 8; So-93, August, 1901. J

VANISHING WILD FLOWERS

IjY ElIZAKEIm' G. 15RITTON

A number of articles on this topic have been pubhshed this
year. They have awakened the interest of many readers, caused
much comment and discussion, and prompted investigation as to
the reasons for this calamity, which, if it does actually come to
pass, is as much to be deplored as the extermination of the buf-
falo, the seal or the beaver. As in the case of mammals and
birds, ereed and thoughtlessness combine to do the harm, and
fashion and selfishness are the motives.

The Neiv York Tribune of May 5th had the following article :
" Now that spring is really here, the picnicking parties are in-
vading the woods north of the Harlem, and have begun the an-
nual systematic destruction of a large proportion of all wild
flowers within reach. The authorities of the Botanical Gardens
are on the lookout for them, and within their own precincts will
guard the blossoms as thoroughly as possible under a well
planned system ; but the rest of the Bronx will be at their
mercy, and that means death to many a poor little plant. It is
not that these ruthless explorers fail to appreciate the beauty of
flowers — they "just love them," in all probability. The trouble
arises from their ignorance of the extent of the damage they do,
and from an utter inability to comprehend that a flower or any-
thing in the vegetable world has rights which the lord of crea-
tion himself is bound to respect. Thanks to the picnickers and
alleged botanists, the arbutus, loveliest of spring blossoms, has
been almost exterminated in the Bronx region. Its delicate pink
and white used once upon a time to hide under the leaves all
through the northern woods in that part of the suburbs ; now it
may be found only in spots where it commands less enthusiastic
admiration. The mountain laurel has shared a similar fate."

86

"The most curious feature of this destructive energy is that
the plants and flowers so carelessly torn from their homes prob-
ably give little or no satisfaction to those who take them away
with them. Is there, for instance, a more unsatisfactory flower
to make attractive in a vase than the arbutus ? It pines for its
screen of leaves ; the loveliness that seems so perfect when half
hidden becomes quite inadequate when pulled out into the glare
of the day and put down in a city room. The same is true of
most other delicate wood plants. They depend upon the charm
of their surroundings." In Tlic House Bamtiful, July, 1901.

" Is there a flower lover who has gone out into the country with
a party of young people unaccustomed to find themselves sur-
rounded with green who has not noted with something very like
hopeless rage the immediate rush on every growing thing in the
neighborhood, its instant uprooting and subsequent careless toss-
ing aside ? Later in the day, before going home, when all the blos-
soms in the immediate neighborhood have been destroyed, there
is a search for fresh fields, and another spot is denuded. A few
dejected blossoms are all that is left when home is reached ;
nothing of any value remains out of all the lives butchered to
make an East Side holiday. The saddest part of it is, not that the
children do it, for that might be pardoned on the score of ignor-
ance, but that those in authority permit it without a remonstrance."

" The flower's right to existence nobody takes into account, or
the harm done to the children by allowing them to think that
they may destroy life as they choose."

And in this connection arises the question of public rights on
private property. I know that less than fifty miles from New
York, a man of wide and varied culture and sympathies, a mem-
ber of a variety of horticultural and agricultural societies, own-
ing a large tract of land away from any large town, has at-
tempted to plant the waste roadside places and private woodland
with wild and cultivated flowers, and repeatedly seen great bunches
of them carried off by people, walking or driving by, who did not
realize all the trouble and expense he had been to, in order to
beautify the roadsides for tlicm and for others who might come
later. Many a prized Azalea bush has been rifled before its owner
knew it, often thoughtlessly and without evil intention by those
who "just love them."

87

The destruction of wild flowers not only takes place while
they are in bloom but there is another cause which is even worse
than picking and that is fire. In early spring, March and April,
when the woods are dry and high winds prevail, a match thrown
carelessly among the dry leaves will start a fire which soon at-
tains serious proportions and is often difficult to conquer, so that
in a short time nothing remains but charred stems and scorched
earth from which weeds only can derive sustenance. Fires often
result, in the New York Botanical Garden, from the carelessness of
smokers, and they always start near the paths.

The same selfish disregard of consequences impels picnic
parties to come and scatter unsightly papers, boxes and broken
glass, in spite of the fact that according to the city ordinances
they are guilty of a misdemeanor and liable for each offence to
a fine of from one to five dollars.

These evils are due to thoughtlessness and selfishness ; now let
us see what fashion is doing. In the July number of the House
Beautiful occurs the following paragraph :

" The rarest flower in Europe, the edelweiss, is becoming
scarcer every year, and unless measures are taken to prevent in-
discriminate gathering it is likely to disappear altogether. The
edelweiss only grows 2,500 or 3,000 yards above the level of
the sea and under special climatic conditions. Unfortunately, the
edelweiss has become the ' fashionable ' flower in Germany since
the Emperor commenced wearing it."

The truth of the matter is that for commercial purposes, the
wild supply of edelweiss has long been insufficient and for many
years it has been impossible to gather it " indiscriminately," for it
is only to be found in the most inaccessible places. But it has
lone been cultivated for sale to tourists and makers of souvenirs.
In fact, the edelweiss is not difficult to grow in suitable localities,
and even in unsuitable ones for it, such as the New York Botanical
Garden, it has grown and blossomed for two years in succession,
in one of the Composite beds of the Herbaceous Grounds, next
to its North American allies, the everlastings and cudweeds.
The living plants were obtained from the Buffalo Botanical Gar-
den, where it has also been grown, but a succession of hot, dry
summers has killed it. In the shaded and moist rock-garden
yet to be built it may probably be made to live.

88

It would be interesting to learn how many boxes of arbutus
are annually mailed in the United States and how near extermina-
tion it is at the several stations where it was formerly abundant.
We know that at Lakewood there is little of it left, and we hope
that George Gould will protect it within the limits of his estate.
It is the only way that certain rare plants and birds have been
preserved in England, and we are rapidly finding such restrictions
necessary. At Natural Bridge all persons are forbidden picking
wild flowers. Various places have their fashionable favorites ; in
the Berkshires it is the fringed gentian, in Boston the Sabbatia,
at several places in Pennsylvania it is the Rhododendron, Kalmia
and Azalea, and New York may well claim first place as destroyer
of the Holly and Prinos berry. We may well ask, also, where
will the Christmas trees and greens come from in the future, if
they do not cultivate the balsams and spruces, and cease the reck-
less destruction of ground-pine and laurel. We are sending now
to the southern states for most of the holly and mistletoe and to
the states northeast of us for Christmas trees.

Before it became the fashion to use "Galaxy" for funeral
wreaths, Galax was very abundant in the southern Alleghanies,
but now that the leaves are picked by the crate-full, it is becom-
ing more expensive. It is to be hoped that they do not " kill the
goose that lays the golden t^z^

The custom of filling jardinieres with ferns has destroyed many
pretty nooks in Bronx Park and is the cause of endless trouble,
as the propensity to take them and ignore the signs, seems to be
a prevalent feminine failing. None of our native ferns are par-
ticularly suited to this purpose, however, and invariably need fre-
quent renewing, so that it would be easy to exterminate any one
species very soon, if the depredations were permitted and con-
tinued. In the heat of summer nothing is more beautiful and
restful than a fern bank ; but the sight will not be allowed to New-
Yorkers if energetic folk who "must have green things about"
have their way. Much care has been taken to transplant into
suitably prepared nooks and crevices of the Fern corner, the rarer
species and varieties of North American ferns and to surround
them with beds of mosses and rocks and shade. The Walking-
fern has been exceedingly difficult to establish. There are sev-
eral stations for this fern within a radius of fifty miles from New

89

York City, but the stations are kept secret by those members
of the " Torrey Botanical Club " who know them, for fear that it
will be exterminated. This is the experience of a New England
botanist who mourns about her losses in Rhodora for March.
"We find the 'Walking-leaf — to use Dr. Bigelow's English
name for the Caviptosorus — but a pretty habit that it has may be
the death of it. Growing amongst mosses and other low plants
that need but little depth of soil, and whose interlaced roots weave
the whole together, it frequently carpets the flat tops of rocks —
a beautiful sight which draws the attention even of idle picknick-
ers who, not realizing that they are destroying years of growth,
find it amusing to peel off these mats and then, without a pitying
thought, throw them on the ground to die. Thus they have laid
bare the rocks within sight of their walks ; but away from the
paths the interesting fern is still common."

In Connecticut the Hartford or Climbing fern was so nearly
exterminated for decorative purposes that a law was passed pro-
tecting it. We hope that the newly-awakened popular interest in
ferns will not lead to the same sad results, but rather cultivate the
love of these beautiful plants and help to protect them.

The Natural Science Committee of the Associate Alumnae of
the Normal College has been doing a good work, among the
children of the East side, in distributing "Nature Material,"
holding flower-shows in various places and making " loan-col-
lections " for the use of the teachers. We quote from the last
Annual Report which reaches 1,500 members.

During the past year the Natural Science Committee has done
all in its power to arouse greater interest in Nature Study, es-
pecially among the teachers and children of the public schools.
With this object in view, the following lines of work have been
carried on :

First, the distribution of "nature material" to the Alumnae
School Representatives. There have been five of these distri-
butions and the specimens have been as far as possible " accord-
ing to season," that the children might come into touch with
what was going on in the great world of nature, even though
obliged to spend their days amid brick and mortar.

October 12, 1900. Fruits and seeds of all kinds, as well as
some of the late flowers, as witch-hazel and fringed gentian.

90

December 7, 1900. Evergreens of many kinds, holly, ground-
pine, etc.

January 16, 1901. Birds' nests, cocoons, starfish, sea-urchins,

shells, minerals, etc.

February 28, 1901. Budding twigs in great variety, also
mosses, lichens, fungi and sea-weeds.

April 16, 1 90 1. Material for aquaria: frogspawn, tadpoles,
snails and aquatic plants. A number of maple seedlings in paper
pots were also distributed. In a number of instances barrels of
labeled specimens have been sent direct to various down-town
schools. "About twenty boxes and baskets were sent every
week to the ten vacation schools."

"Through the distributions we reach the teachers, but by means
of the flower shows we come into direct touch with the children.
Last May the experiment was tried of holding a flower show in
one of the down-town schools. It proved even more successful
than we had anticipated and we hope to make this a permanent
branch of our work, as it seems to be more far-reaching in its
effects than anything else we undertake. The flowers are en-
joyed, not only by the children of the school in which the exhi-
bition is held, but by the neighboring schools which are invited
to visit the show after school hours ; the mothers of the children
are asked to come, and if there chances to be a little leisure time
the children from the street are invited in."

It is evident from the above quotations that such wholesale
quantities as these must be judiciously selected, or there will be
no wild flowers left within easy transportation of the city. One
member is reported to have sent i 50 pitcher-plants from a bog at
Plymouth, Massachusetts! We question the wisdom of gathering
either fringed gentians, pitcher-plants or christmas-fern, in quan-
tities sufficient for distribution to seventy-five teachers, and we
hope the pupils of country schools may be guided to make a wise
selection, if they are encouraged to send flowers in quantities to
the city schools.

The flowers which may be picked in large quantities, without
likelihood of extermination are dandelions, violets, daisies, butter-
cups, black-eyed Susan, wild carrot, clovers, sweet melilot,
golden-rod, asters, and grasses innumerable. But the rarer,
dainty woodland wild flowers, that fade as soon as they are picked

91

such as spring-beauties, dogtooth-violets, bloodroot and hcpati-
cas, columbines, anemone, arbutus and pyxie, azalea and laurel,
wilk pinks, geraniums and roses and lilies and orchids, dogwood
and viburnums, are far better left to reproduce their kind and add
new loveliness in new places next year and for manj' years after.
Several times since our connection with the New York Botanical
Garden I have stopped children and teachers who were picking
flowers or breaking branches of trees, and have been told it was
for " nature study " or for " school " and when asked if they did
not know it was against the rules of all public parks to pick any-
thing, they almost invariably replied either that they had been in
the habit of picking in this place before it became a Park or a
Garden and did not see why they should not continue to do so ;
or they implied that the object for which they were to be used
justified the breach of law. The making of loan collections
for the teachers is an excellent plan, and the accumulation of
local floras at two or three different educational institutions also
will help them. For the children, there are the Museums of
National History and Botany and the Children's Museum in
Brooklyn, but we hope that besides these, we shall have for a
long time yet, places near the city, where the wild flowers may
be seen growing and that the children of the public schools may
not only learn to ''know them by name and enjoy them,'' but leave
them to continue their growth. The greatest destruction of all,
comes from the draining, clearing and cultivating of wild lands ;
and in the vicinity of large cities, by the continued extension of
their limits ; this, of course, is unavoidable.

The Metropolitan Park Commission of Boston has printed a
Flora of the parks within their jurisdiction, compiled by various
local botanists who volunteered their services, organizing and
cooperating for this purpose. It was published in 1896 and
special localities were given for a number of rare plants, among
them Pogonia verticillata, Habenaria fimbriata, Epigaea repcns,
Kalmia latifolia and Conopholis Americana and in the preface we
find the following references to them : " The public should be
exhorted, if they come across such plants as these, to preserve
them rigidly. The true botanist and lover of nature needs no
such exhortation."

It would be interesting to know, whether any injurious results

92

have followed its publication, or whether the Boston public has
reached such a high degree of culture both moral and aesthetic
that no one makes an exception even of himself?

• But the climax has been reached in an advertisement which
has been printed in Rhodora, the Journal of the New England
Botanical Club, since May. It announces that on the Bangor
and Aroostook Railroad, there is to be had the " best botanizing
in the Eastern States," and proceeds to give the names of stations
and lists of rare plants to the length of two whole pages. We
ask with amazement, where did they get the information ? What
botanist sold his birthright for a few railroad passes ? For-
tunately, many of the plants listed are so rare that only the dis-
criminating and trained botanical specialist will be sure to find
them, and the general public will hunt a long time and not
know them when they see them.

Mr. Redfield used to tell the story of Rafinesque that when he
first found Corenia Conradii, he threw himself down upon it and
stretching out his arms, said "all that I cover is mine." It is
not always the most enlightened who are the most unselfish.
There have been botanists, even in the Torrey Club, who up-
rooted plants wantonly and made no good use of them after they
were picked. But the custom of carrying "tin trunks" has
been largely superseded by presses, and only a few duplicates
are now made of each species.

The flora of Great Britain is, perhaps, the best known of any
in the world ; and there is more knowledge among the working
people of special and difficult branches of botany, probably than
in any other country, Germany not excepted. Dr. Braithwaite
told me that he had sold a great many copies of the British Moss-
fiora to the Manchester weavers.

But many of their rarest plants have been exterminated by
botanists, as shown by the following quotations taken from the
Journal of Botaiiy for July :

" The accuracy in general matters for which the Daily ATail has
long been conspicuous, extends to its botanical information. We
reproduce the most recent item in the hope that the publicity now
given to the methods of the ' professional botanist ' will cause him
to abstain from this nefarious means of adding to his income.

" Four of the daintiest of English wild plants are rapidly dis-

93

appeann<^ from this country, and one, at any rate, can rarely be
seen outside Kew gardens. This is the Cypripediuin Calccolus
commonly known as the ' lady's slipper.' It is really a wild
orchid, with a pretty yellow flower resembling in shape the article
which has given it its popular name. The other vanishing plants
are the Osnninda rcgalis, the Scolopendrium vulgai-e (hart's
tongue), and the Asplenmni viridc (green spleenwort), all of which
are ferns. Their disappearance is due to the depredations of the
tourist, especially of the cyclist, and the professional botanist,
who scours the woods and disposes of his ' finds ' for a few pence
in the streets of the nearest large town." — Daily Mail, June 26.

" There can, however, be little doubt that, apart from the
ravages of ' professional botanists ' and the destructive efforts of
various local bodies, who throughout the country are engaged
in destroying grassy roadsides and scarifying hedgebanks, to the
great advantage of the nettles, docks and other weeds which take
the place of the native vegetation, our British plants are threat-
ened with a new danger."

"I have before me the programme of the Essex Technical In-
struction Committee for Field Studies in Natural History. The
course for 1901 is intended to instruct teachers in the elements
of botany by means of rambles in search of wild flowers. One
leading feature is a vacation course of ten days in the New
Forest. The teachers are to be accompanied by local guides,
and their attention is particularly directed to the rarest species,
which are specially named, as well as the places in which they
are known to grow. To collect, dry and identify plants is the
chief aim of the leaders, who not only urge every teacher to
make his own collection, but suggest that duplicate plants will
prove useful for ' special fascicles.' It seems to me lamentable
that teachers should be advised to study natural history by
schedules, and to gather plants merely in order to name and dry
them. I imagine that they will be worse and not better for
working through so dry and barren a course. Nothing shows
the want of judgment of the promoters more clearly than that
untrained botanists should be seriously advised to pay particular
attention to the difficult and uncertain subspecies of the common
bramble. But all of us, whether we are concerned with the
teaching of botany or not, have an interest in the preservation of

94

»

our native plants. The Essex Committee is simply organizing a
raid upon plants which are already near to extinction. I hope
that they will fail to discover the rarities which they selfishly
covet; their enterprise is, I venture to say, an injury to natural
history and to education alike. It may not be too late to get
this programme cancelled, and I would beg those who care for
live natural history to use their influence in diverting the atten-
tion of the Essex collectors to some other pursuit where they
will do less harm."

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 17

THE TYLOSTOMACEAE OF NORTH AMERICA

BY V. S. WHITE

NEW YORK
1901

[Reprinted from the Bulletin of the Torrey Botanical Club, 28 : i^l-MH,, 27 Aug., 1901.]

The Tylostomaceae of North America

Bv V. S. White
(With Plates 31-40)

The members of this family arepuffball-like plants, which form
underground in the shape of rounded masses, appearing at first on
the mycelium as minute thickenings, and gradually reaching their
full development. The ball has a thick outer covering, and an
inner, and usually thinner one, commonly known as the peridium
proper. The upper and larger portion of the ball is composed of
sporogenous tissue, and there is a lower sterile portion which elon-
gates when conditions are most favorable, forcing the upper portion
up through the surface of the ground, and consequently rupturing
the outer coat which originally enveloped the whole mass. In
some genera this outer coat remains at the base as a distinct cup-
like volva, the upper portion then being carried up almost intact
and falling away or adhering to the peridium ; in other cases the
outer coat is ruptured irregularly, bearing very slight traces at the
base of the stem or even none at all, and in the genus Tylostoina
this coat adheres more or less to the peridium, nearly always leaving
some traces in the form of a collar at its base. The stem is com-
monly of a firm, almost woody texture, and the plants can be quite
satisfactorily preserved in a dry condition. They vary greatly as
to size and shape, but are all constructed on the same general plan,
having a more or less irregularly globose peridium, and a distinct
footstalk, which is usually, though not always, more than twice
the length of the peridium in mature forms. The methods of

[Issued 27 August.] 421

422 White : Tylostomaceae of North America

dehiscence are very different in the various genera, some having a
definite apical mouth, the peridium withering and collapsing as
the spores escape ; others rupture irregularly from above down-
wards, more as in the Sclerodermataceae. Battarrca has a regular
line of dehiscence where the upper portion of the somewhat hem-
ispheric peridium breaks away, leaving the lower portion at the
summit of the long stem.

In the peridial characters, Tylostoma is closely allied to the
Lycoperdaceae. The relation of this family to the Phallaceae
is one of analogy mainly, shown in the elongation of the stem
which forces the spore-bearing parts to a higher position for the
better scattering of the spores.

In his latest treatment of the family, Fischer* recognizes
four genera : Tylostoma with forty species, widely distributed ;
Qiicletia with a single species from France, failing to note that it
had also been reported from America ; Battarrea with eight spe-
cies ; and Sphaericeps with one species from Angola. The genus
Chlaviydopjis, described by Spegazzinif from Argentina, after
Fischer's first draft was prepared, was later too summarily assigned
as a synonym of TylostomaX for the differences are so marked that
it would be unfortunate to include these two diverse types under
one genus.

Within the past year several specimens of a Oilamydopus have
been found at Mesilla Park, New Mexico, and it was the original
purpose of this paper simply to describe and figure these. It soon
became necessary to study the members of the allied genus
Tylostoma, of which extensive suites of specimens were found in the
Ellis collection, mainly from the western half of the country, rep-
resenting some species hitherto undescribed, so that it was finally
concluded to prepare a revision of the entire family as represented
in North America. Finally some specimens collected by Mr. E.
Bethel, of Denver, Colorado, and sent by him to Mr. Ellis, proved
to belong to an undescribed genus. This large amount of ma-
terial, mostly forming a part of the Ellis collection at the New
York Botanical Garden has made the preparation of this paper

*Engler& Prantl, Die nat. Pilanzenfam. i'**: 342. I900.
I An. Mus. Nac. Buenos Aires, 6 : //. 4. f. 2, 3- 189. 1899.
X Engler & Prantl, Die nat. Pflanzen'^am. i'** : 357. 1900.

White : Tvlostomaceae of North America 423

possible. Mr. E. Bartholomew has kindly loaned material, for
the better description of one species, and Mr. E. S. Salmon has
kindly looked up some data at Kew, England.

Special thanks are due to Professor Charles H. Peck, of Al-
bany, for material and suggestions, and more especially to Pro-
fessor L. M. Underwood, of Columbia University, under whose
direction the work has been undertaken and whose private collec-
tion has furnished considerable additional material.

The genera of the family Tylostomaceae may be recognized
by the following synopsis :

Peridium opening by an apical mouth.

Peridium with a collar underneath, formed about the cylindrical stem ; volva in-
definite. I. Tylostoma
Peridium without a collar ; stem much enlarged where it joins the peridium ;
volva cup-like, flaring. 11. Chlamydopus
Peridium circumscissile.

Peridium hemispheric or nearly plane below, dehiscing at the margin of the

plane of the hemisphere. Ill- Battarrea

Peridium spherical, dehiscing at the equator. Sphaericeps (extra-limital).

Peridium opening irregularly.

Peridium readily separating from the stem ; capillitium free.

IV. QUELETIA

Peridium closely attached to the stalk ; capillitium embedded in a membranous
tissue. V. DiCTYOCEPHALOS

I. TYLOSTOMA Pers. R6mer, Neues Mag. Bot. i : S6. 1794
The first reference to a plant certainly belonging to Tylostoma,
is to be found as early as 1696, in Ray's Synopsis * under the
name of Fuiigics pidvendentus minimus. He describes it in a few
Latin words, " Pediculo longo, insidens. In agris circa Londinum,"
and adds in English, •' the least dusty mushroom, with a long foot-
stalk, collected by D. Tancred Robinson, M.D." The next men-
tion is made in 1700 by Tournefort t who also gives the first figure
of a Tylostoma ; this has the characteristic short tubular mouth of
T. pediinailatitm, a slender, smooth stem, and closely resembles
specimens of this plant found in our own country. Micheli % next
figured the other variety of T. pediinadatiim under the name, sqiia-
mosiim. This figure is not so easy to identify as that of Tournefort ;

*Syn. Meth. Stirp. Brit. 2d ed., l6. 1696.

flnst. rei Herb, i : 563. //. JJ/- / E, F. 1700.

J Nova Plantarum Genera, 218, nos. 10 and IT. //. gy. f. i. 1729-

424 White : Tylostomaceae of North America

none of the European specimens of this variety seen, have such a
rough stem, or a peridium of the exact shape as shown in the figure,
but it is evident notwithstanding, what it was meant to represent.
T. peduncidatum being the commonest European species, and
widely distributed tends to vary and extreme specimens are found
to differ so widely that were it not for the many intermediate
stages, the extremes might well be kept distinct, as indeed has
been done by several writers.

Linnaeus* refers to this same plant by its first binomial,
Lvcoperdon pedimcidatuui, and cites the above-mentioned descrip-
tions of Ray and Tournefort. His description is in the following
words : " Stipite longo, capitulo globoso glabro ; ore cylindrico,
integerrimo. Habitat in campestribus." In 1794 Persoon
established the genus Tidostoma.\ He described it — " Peridium
stipitatum, ore cylindrico cartilageneo " — and mentions two species
T. brumalc {L. pcdwiadatiim L.) and T. imbricatwn (Micheli Ly.
11); these two species have since been commonly referred to,
under the name of T. inammosjim (Mich.) Fries, but all these
specific names are antedated by pednnadaUnn of Linnaeus.

Since the time of Persoon, various species of Tylostonia have
been described, and De Toni ]: enumerates thirty -four species from
different parts of the world.

Our own species were studied in 1890 by Morgan, § but
evidently from a limited amount of material. In his paper
he enumerates five species, of which T. Diamviosiim and
T. fimbriahun are originally European, two were new native
species, T. vcrriicosiun and T. canipestre, and the fifth he
called T. Meymiamun Kl., but it is evident that the original
T. Meyenianuni was not a true Tylostonia but a poorly figured
species of the genus Chlamydopus, and as the material on which
Morean based his determination is not now available, we are uncer-
tain as to what his last species really is. T. pcdunculatuin was first
reported from this country in i8i8by Schweinitz|| under the name

*Sp. Plant. I184. 1753.

f Tylostoma from ruXoa, callous skin, and aTdfia, a mouth.
X Saccardo, Syll. Fung. 7 : 60 ; 9 : 268 ; II : 159 ; 14 : 258.
^Jour. Cin. Soc. Nat. Hist. 12 : 163; //. /6. f. 1-5. 1890.
II Syn. Fung. Car. 34. 1818.

White : Tylostomaceae of North America 425

of T. hrumale Pers. from North Carolina, and in 1834* he gives
two species, T. bruniale and T. sqiiaumlosiim,\ of which latter he
writes "not a variety of the preceding but closely resembling
Rinella of Rafinesque,:}; " but notwithstanding his assertion to the
contrary T. squamosum is now commonly regarded as a variety of
T. pedimculatum {brumalc). In 1837, Montagne§ described T.
cxaspcmhim from Cuba, collected by Ramon de la Sagra, the
well-known Cuban naturalist, and this species has since been re-
ported from India. In 1867 Curtis'!! mentions T. fimbriatum and
T. mamtnosum as " common in North Carolina," and six years
later Berkeley** reported T. fimbriatum, T. mammosum, and
T. Meyenianum, collected by Wright in New Mexico. As late
as 1 89 1, Masseeft published Berkeley's description of T. WrigJitii,
also from New Mexico. Other American species have been de-
scribed since, T. obcsum C. & E., %% T. punctatum Peck, §§ and T.
semisulcatum Peck. *^ Much of the western material found in the
Ellis collection had been referred to T. obesum, although it repre-
sents very diverse forms. T. punctatiim has been reported from
several localities, but of T. semisulcatum only the type specimens
are known, and these are inaccessible at Albany, if indeed they
are still in existence. Extensive collections have been made in
different parts of the United States in the past few years and we

* Trans. Amer. Phil. Soc. 4: 256. 1834.

t Since writing the above it has been possible to examine Schweinitz' original
specimens, which are carefully preserved in the Philadelphia Academy of Natural
Sciences. There is a question as to the specimen marked T. brtanale being T. pedtin-
culatnm, to which it is usually referred, owing to the color of the peridium, which is a
bright terra-cotta red, unlike anything seen before — but this may be due to the color of
the clay in which the plant grew or to other causes of unknown origin. The specimen
marked T. squamosum resembles greatly some of our specimens of T. fimbriatum in
general habit and spore characters, with the exception of the mouth which is entire, but
not tubular and prominent, like the mouth of T. pedtinadahtm — which fact may be
owing to the wasting of the cartilaginous tissues occasioned by time — so that it is hardly
possible to determine what this plant really was.

JSyst. F. Veg. 139.

§ Ann. Sci. Nat. II. 8 : 362. 1837.

T[Geol. and Nat. Hist. Survey North Carolina, 3 : no. 1867.

** Grevillea 2 : 49. 1873.

ft Grevillea 19 : 95. 1891.

XX Grevillea 6 : 82, //. 100. f. 24. 1878.

\l Bull. Torr. Club, 23 : 419. 1896. ^

ll^Bull. Torr. Club, 22 : 209. 1895.

426

White : Tylostomaceae of North America

find among them several additional species. The indications are
that we are only at the beginning of our knowledge of their distri-
bution.

The following table shows the distribution of the hitherto
known species of Tylostoina :

'J

2
o

(A

S

V ■

C(t

(Li

6

rt

ug

6

In

Tylostoma.

North Am

South Am

Europ

*5)

lis

<

1
tn

3

1"
8<:
'%

1

en

album.

+

Angolense.

+

+

Barlae.

+

Barbeyanura.

+

Berteroanum.

+

Bonianum.

+

Bossieri.

+

campestre.

+

caespitosum.

+

carneum.

+

Cesatii.

+

exasperatum.

+

+

fimbriatum.

+

+

+

•

Giovanellae.

4-

granulosum.

+

Jourdani.

+

laceratum.

+

leprosum.

+

Leveilleaiium.

+

mammosum.

+

+

+

+

+

+

maximum.

+

Mollerianum.

+

montanum.

+

obesum.

+

pulchellum.

+

pusillum.

+

'+

Patagonicum.

+

Schweinfurthi.

+

squamosum.

+

tortuosum.

+

volvulatum.

+

verrucosum.

+

Wightii .

+

Wrightii.

+

The species of Tylostoma are found principally in sandy regions,
in dry barren fields, rarely in woods or shaded places. As yet
very little is known of their habits, growth and development as no
little difficulty attends the study of these strange and interesting
plants in their early subterranean stages. It is hoped that stu-

White : Tylostomaceae of North America 427

dents in the districts where these plants are most common will
gather material in the earlier stages of growth — digging down
where mature forms are found above ground in the hope of find-
ing the young peridia.

As far as can be ascertained, only two papers have appeared
treating of the growth of these plants. Schroeter * has published
an account of the development of the spores and basidia. Pro-
fessor Bessey t has a short note on the growth of T. jiianimo-
sum in which he says, " The ball forms underground, and reaches
maturity there so far as the spores are concerned. Tylostoma
agrees with Lycoperdon in having the interior of the ball composed
of two portions, first a spore-bearing part which occupies most of
the interior, and second a sterile base composed of tissue which
does not produce spores. In Tylostoma a portion of this tissue of
the sterile base remains living until the spores ripen. Then this
tissue begins a rapid growth and a cylindrical stalk is produced
which forces the ball through the overlying earth and carries it up
several inches." Massee,| speaking of Tylostoma mammosnm says,
" resembling a Lycoperdon with an elongated stem, but readily dis-
tinguished by the groove between the apex of the stem and the
peridium, and by the threads of the capillitium being nodulose at
the base of the septa." And writing of the stem, "it is some-
times smooth, and at others broken with small irregular fibrillose
squamules which are sometimes arranged more or less in circles."

Synopsis of our Species

Mouth entire, short tubular.

Spores smooth, or occasionally with a few minute scattered warts.

1. T. albicans.
Spores uniformly and densely verrucose.

Peridium smooth ; capillitium not much swollen at the joints.

2. T. pedunctilatum .
Peridium warty ; capillitium swollen at the joints. 3. T. verrucosum.

Mouth lacerate-fimbriate, not tubular.
Spores smooth or nearly so.
Mouth plane.

Stem tapering toward the base ; peridium depressed.

4. T. gracile.

*Cohn's Beitr. Biol. Pflan. 2 : 65. 1877.
fAmer. Nat. 21: 665. 1887.
+ Ann. Bot. 4 : 85. 1S89.

428 White : Tvlostomaceae of North America

Stem not tapering ; peridium globose.

Capillitium freely septate ; stem short (1-1.5 cm.), collar close.

5. T. viinutiim.
Capillitium rarely or never septate ; stem longer (1.5-5 cm. ), collar
distant. 6. T. Kaiiscnse.

Mouth raised, convex.

Capillitium freely septate ; outer peridium long persistent.

7. T. poculatum.
Capillitium rarely or never septate ; outer peridium soon receding.

8. T. obestim.
Spores verrucose or granular.

Mouth plane. 9- ^- campestre.

Mouth raised, convex.

Spores furnished with long blunt spinules ; stem short (0.75 cm. ), rough

spiny. 10. T. exasperattuii.

Spores with coarse scattered tubercles ; stem longer (3 cm. ), even, nearly
smooth. II- T. tuberculatum.

Spores finely granulate-verrucose.

Capillitium sparingly septate, not much swollen at the joints.

Stem elongate (6 cm.), with copious persistent mycelial strands.

12. T. fibrillosutn .
Stem shorter (3.5 cm.), without mycelial strands.

13. T. subfuscum.
Capillitium freely septate, swollen at the joints.

Peridium smooth ; inouth fimbriate. 14- T. fitubriatuni.

Peridium pitted ; mouth lacerate. 15. T. punctatum.

I. Tylostoma albicans sp. nov.

Peridium depressed globose, 0.7-1 cm. high, 1-1.5 cm. in
diameter : outer peridium scaly, retreating and leaving a smooth
whitish surface to the inner membranaceous peridium : mouth short
tubular, entire, prominent : collar irregular, 3-5 mm. distant from
the stem : stem equal, slightly thickened at the base, the outer coat-
ing lacerate scaly, whitish like the peridium, 4-6 cm. long, 0.5 cm.
in diameter : capillitium whitish, hyaline, branched, septate, slightly
swollen at the joints, 4-6 // wide, free ends rounded : spores reddish
brown, irregularly globose, pedicled, smooth, or if rough only a-
vcry few of them so, 4-5 /^ in diameter. (PI. 31, f. A--?-)

Texas : E. D. Cope.

2. Tylostoma pedunculatum (L.) Schroeter ; Cohn, Beitr. Biol.

Pflan. 2 : 65. 1877
Lycoperdon pedunculatum L. Sp. PI. 1 1 84. 1753-
Tulostoma bnimalc Pers. Romer Neues Mag. Bot. i : 86.

1794.

Tulostoma mammosum Fr. Syst. 3: 42. 1829.

White: Tylostomaceae of North America 429

Lycoperdon II. Mich. Nov. Plant. Gen. 218. 1729.
Tiilostoma imbricatiim Pers. Romer Neues Mag. Bot. i : 86.

1794.

Lycoperdon sqiiamoswn Gmel. Syst. Nat. 2 : 1462. 1796.
Tidostoma sqiianiosiiui Pers. Syn. Meth. Fung. 139. 1801.
Tidostoma mainmosiim Fr. Syst. 3 : 42. 1829.
Tylostoma mammostivi var. squamosinn De Toni ; Saccardo,
Syll. Fung. 7: 61. 1888.

Peridium subglobose, 1-1.5 cm. high, 1-2 cm. in diameter,
the brown outer peridium soon retreating, leaving the inner peridium
smooth and membranaceous : mouth short tubular, entire, promi-
nent : collar inconspicuous : stem slender, lacerate scaly, or nearly
smooth, with a small mycelial bulb, stuffed with loose silky threads,
1-5 cm. long, 2.5 mm. in diameter: capillitium 4-7 // thick, sep-
tate, somewhat swollen at the joints, light colored, hyaline : spores
subglobose, 3-5 11 in diameter, minutely verrucose, some short
pedicled. (PI. 31, f. 1-3.)

Exsicc. E. & E. N. A. Fungi 2734 (as T. inainmosum). Ra-
venel. Fungi Am. Ex. 137 (as T. mammosiini).

New York, Undcrivood, Fischer; New Jersey, Ellis ; Penn-
sylvania ; Michigan, McBride ; Wisconsin, D. V. B ; North
Carolina, Curtis, Wood ; Kansas, Kellerman, 770; Indiana, Cook,
Undcrzvood ; Florida, Underwood; Nebraska, Webber ; Texas,
Harris; Colorado; Iowa; New Mexico, Wright, Fcndhr, Cock-
erell ; Mexico, C. L. Snnth.

3. Tylostoma verrucosum Morg. Jour. Cin. Soc. Nat. Hist. 12 :

163. //. 16. f. 2. 1890
Peridium depressed globose, 1-1.2 cm. high, 1-1.5 cm. in di-
ameter : outer peridium of scales and warts persistent : inner
peridium brownish, becoming smooth with age : mouth entire,
raised, prominent, short tubular : collar of delicate lacerate points
descending about the top of the stem : stem 4-5 cm. long, .5 cm.
in diameter, slender, the surface lacerate scaly, brown, hollow,
cylindrical and having a large basal bulb composed of the mycelial
strands and adherent earth : capillitium almost white, hyaline,
slender, septate, some of the joints rather swollen, ^-^ 1^ wide:
spores subglobose, granular, lightish cinnamon-colored, 4-7 //. in
diameter. (PI. 31, f. 8-10.)
Ohio : Morgan, C G. Lloyd.

Growing on the ground in rich soil in woods. The mycelial
bulb is usually larger than the peridium.

430 White : Tylostomaceae of North America

4. Tylostoma gracile sp. nov.

Peridium depressed globose, 1-1.5 cm. high, 1.5— 2cm. in di-
ameter : outer peridium retreating, leaving but slight traces on the
inner peridium which is thin, smooth, whitish and rather shiny :
mouth plane, lacerate : collar entire, restricted around the top of
the stem : stem slender, 3-5.5 cm. long, 6 mm. in diameter at the
top, 3 mm. in diameter at the base which is slightly enlarged,
somewhat sulcate, lacerate, whitish within and without, fibrillose
stuffed, becoming hollow : capillitium dark ferrugenous, 4-6 [/.
wide, ends rounded, and having swollen places mostly near the
ends, branched, rather thick-walled, many threads flattened, septa
very scarce : spores subglobose, 3-5 jjl in diameter, smooth, short
pediceled. (PI. 32, f 1-3.)

New Mexico : Las Cruces, ]Vootou.

5. Tylostoma minutum sp. nov.

Peridium globose, 0.7-1 cm. high, 0.7-1.2 cm. in diameter;
outer peridium scaly, retreating, leaving smooth, brownish surface
to the thin inner peridium, remnants of the outer peridium forming
a cup around the base of the inner peridium : mouth plane, fimbri-
ate, small ; collar entire, indistinct : stem 1.5—2 cm. long, 3 mm. in
- diameter at the top, 5 mm. in diameter at the base, slender,
slightly enlarged at the base, hollow, fibrillose stuffed : capillitium
yellowish, hyaline, sparingly branched, rather thin walled, ends
rounded, septa swollen at the joints, 3-5 n wide : spores irregularly
globose, nearly smooth, pediceled, thick-v/alled, 3-5 jj. in diameter.
(Pi. 31, f 11-13.)

Colorado, Bethel (type) ; Kansas, Kellcrnian.

6. Tylostoma Kansense Peck, sp. nov.

Peridium subglobose, 1-2 cm. broad, 1-1.5 cm. high, thick,
firm, glabrous, white, somewhat flattened at the base : mouth
slightly lacerate on the margin, plane, sometimes slightly promi-
nent : collar wide, membranaceous, 2.5 mm. distant from the stem :
stem equal or slightly narrowed toward the base, 1.5—7 c"^- lo"S>
6—8 mm. thick, hollow or stuffed with silky fibrils, somewhat
sulcate-striate, white within and without, slightly and abruptly
bulbous: capillitium sparingly branched, colored, 7— ii jJ- wide,
hyaline, septa not seen : spores subglobose, 4—5 ii in diameter,
brownish ferruginous. (PI. 32, f 7-9-)

KANS.A.S, BartJiolomeio. July. It has rarely two ostiola.

White: Tvlostomaceae of North America 431

7. Tylostoma poculatum sp. no v.

Peridium globose, somewhat depressed, 1-1.5 cm. high, 1-2 cm.
in diameter, smooth, fawn-colored, membranaceous : outer peridium
scaly, but more persistent than in most species, remaining in the
shape of a cup-like involucre round the base of the peridium,
mouth slightly raised, fimbriate, mostly large : collar entire, in-
conspicuous : stem 1—3 cm. long, ^—6 mm. in diameter, cylindri-
cal, firm, slightly bulbose, hollow or stuffed, often with consider-
able remnants of the outer peridium attached : capillitium lightish
yellow, sparingly branched, septate, swollen at joints, 4-7 /i wide,
rather thick walled : spores ferruginous, subglobose, smooth, or
irregularly ridged in the older specimens, owing to the shrinking of
the inner substance, short pedicelcd, 4-5 li in diameter. (PI. 34,
f. 4-6.)

Nebr.\ska : Lone Pine, Bates, 462 (type) ; Alabama : Tuske-
gee, Carver ; Colorado : Boulder, D. M. Andres. Plants grow-
ing singly or in groups of twos and threes.

8. Tylostoma obesum C. & E. Grevillea 6 : 82. pi. 100. f. 24.

1878

Peridium globose pyriform, 2.2 cm. high, 2 cm. in diameter,
smooth, leathery, whitish, with a kid-like finish : outer peridium
scaly, retreating, leaving a narrow ring around the lower part of
the peridium : collar entire, descending about the stem : mouth
raised, lacerate, rather large : stem 4 cm. long, hard, leathery,
somewhat sulcate, slightly tapering at the base, 8 mm. in diameter
at the top, 5 mm. in diameter at the base : capillitium dark
ferruginous, slender, free ends rounded, branched, septa none or
very scarce, 3—6 ix wide : spores globose, smooth, short pediceled,
3-6 p. in diameter. (PI. 32, f. 4-6.)

Colorado.

This description and the figure are based on the specimen of
this species which Mr. Ellis retained when he sent other specimens
to M. C. Cooke for a description. It will be seen that the dimen-
sions and shape here given differ considerably from the one fig-
ured in Grevillea. The type is at Kew (2715 Cke.), and through
the kindness of Mr. E. S. Salmon, it was possible to compare some
of the spores and capillitium which agree in all respects with the
original here, but the Kew specimens are apparently not so
mature.

432 White : Tyi-ostomaceae of North America

9. Tylostoma campestre Morg. Jour. Cin. Soc. Nat. Hist.
12 : 163. pi. 16. f. 4. 1890

Peridium subglobose, 1-1.5 cm. high, 1-2 cm. in diameter:
outer peridium brown, scaly, retreating and leaving an irregular
ring around the lower part of the smooth inner peridium : collar
irregular, descending about the top of the stem : mouth plane,
lacerate, in some specimens being a mere crack or slit : stem
rather thick, 3— ID cm. long, i — 1.5 cm. in diameter, darker than
the peridium, fibrillose stuffed, becoming hollow, outer brownish
coat lacerate, scaly, and having a small thickened mycelial bulb :
capillitium yellowish, hyaline, cylindrical, 4-8/^ wide, branched,
septate, somewhat swollen at the septa : spores subglobose, warty,
some short pedicled, 3-5 n wide. (PI. 33, f. 10-12.)

Exsicc : E. & E. N. A. Fungi, 3297 (as T. gramilosiun
Sw. ?), 3514.

Nebraska: Webber, Bates; California: Underwood, Mc-
Clatchie ; Colorado, Crandall. Growing singly and in groups
in sandy soil.

10. Tylostoma exasperatum Mont. Ann. Sci. Nat. II. 8 : 362.

1837

Peridium subglobose, 1.2 cm. high, 1.3 cm. in diameter: outer
peridium composed of long spinulose scales, somewhat reflexed,
which recede from the top downward, leaving the brown inner
peridium marked with a series of regular pits : mouth raised, fim-
briate lacerate, rather large : collar inconspicuous : stem equal,
slightly enlarged at the base, 5-7 mm. in diameter, 3.5 cm. long,
covered with reflexed lacerate scales, like those on the outer peri-
dium : dark brown: capillitium whitish-yellow, hyaline, 4-6 «
wide, branched, free ends rounded, sparingly septate, not much
swollen at the joints : spores globose, furnished with long blunt
spinules, 5-8/^ in diameter. (PI. -^i, f. 1-3.)

Cuba, Wriglit.

1 1 . Tylostoma tuberculatum sp. nov.

Peridium depressed globose, 1-1.2 cm. high, 1-1.8 cm. in di-
ameter : outer peridium scaly, retreating, leaving a wide band
around the base of the whitish, smooth, and rather thin, inner
peridium : mouth raised, fimbriate, roundish : collar close, incon-
spicuous : stem slender, whitish, hollow or stuffed, slightly en-
larged at the base, 2-3 cm. long, 3 mm. in diameter : capillitium
light yellow, hyaline, branched, thick-walled, variable as to width,
4-8 n wide, rather flattened, ends rounded, broad, sparingly sep-

White: Tylostomaceae of North America 433

tate, swollen at joints : spores subglobose, nearly smooth or with
occasional tuber-like warts, thick-walled, 3-5 // in diameter. (PI.

33, f- 7-9-)

British Columbia: JSIacoun (type); Colorado: Fort Collins,

Baker, 405.

12. Tylostoma fibrillosum sp. nov.

Peridium globose, 1-2 cm. high, 1-2.5 cm. in diameter : outer
peridium retreating leaving a smooth whitish surface to the thin,
membranaceous inner peridium, and leaving a thick portion round
the base of the peridium forming a ring : collar indistinct, close :
mouth raised, fimbriate, roundish : stem equal or slightly tapering
toward the base, sulcate, firm, white and slimy within and without,
somewhat hollow, fibrillose stuffed, and having a small thickened
bulb which in some specimens falls off leaving a flat, whitish sur-
face to the base of the stem, 5-7 cm. long, 5 mm. in diameter :
capillitium whitish, hyaline, 4-8 // wide, branched, some threads
rather thicked walled, sparingly septate, not much swollen at the
joinits, rather flattened : spores subglobose, minutely warted,
darker than the capillitium, some short pediceled, 3-5 fx in dia-
meter. (PI. 33, f. 4-6.)

Ontario : Dcarncss (type) ; Michigan, Hicks. Growing on sand
dunes.

The stem is much covered usually for upwards of 2 cm. with
the mycelial strands, and adhering sand. The specimens from
Michigan have rather more swollen septa.

13. Tylostoma subfuscum sp. nov.

Peridium globose, 0.8-1.3 cm. high, 1-1.5 cm. in diameter:
outer peridium scaly, retreating, leaving a ring around the base of
the smooth, brownish inner peridium : mouth raised, fimbriate :
collar inconspicuous, close : stem brown, somewhat lacerate scaly,
leathery, sulcate, with a small thickened bulb at the base, 2-3
cm. long, 3-5 mm. in diameter ; capillitium lightish yellow, hya-
line, branched, threads long and slender, 4-6// wide, free ends
rounded, occasionally septate, somewhat swollen at the joints :
spores subglobose, minutely warted, some short pediceled, 3-5 //
in diameter. (PI. 34, f. 10-12.)

Colorado, Bethel, 21.

14. Tylostoma fimbriatum Fries, Syst. Myc. 3 : 43. 1829

Peridium subglobose, 1-1.5 cm. high, 1-1.5 cm. in diameter:
outer peridium retreating, having a smooth, brownish surface to

434 White : Tylostomaceae of North America

the thin inner peridium : mouth raised, fimbriate, collar irregular,
inconspicuous, close : stem white within and fibrillose stuffed,
brownish and lacerate scaly outside, with a small mycelial bulb
at the base, 1.5-5 cm. long, 5 mm. in diameter: capillitium light-
colored, almo.st white, hyaline, freely septate, swollen at the joints,
4—8 fJi wide : spores subglobose, dark ferruginous, verrucose, some
short pediceled, '^-6(1 in diameter. (PI. 34, f 7-9.)

Exsicc. : Ravenel, Fungi Car. Ex. 5 : 80 ; Ravenel, Fungi Am.
Ex. 724.

New York, Peck ; South Carolina, Ravenel ; North Caro-
lina, Curtis, JVood ; Alabama, Earle ; Texas, Wright ; Wyom-
ing, Nelson ; New Mexico, Wriglit ; Colorado, Cockcrcll ; Kan-
sas, Kellerman.

I 5. Tylostoma punctatum Peck, Bull. Torr. Club, 23 : 419. 1896

Peridium subglobose, i — 1.3 cm. high, i — 1.3 cm. in diameter:
outer peridium scaly, retreating, leaving a ring around the base of
the peridium : inner peridium thin, whitish, covered with irregular
shallow pits : mouth raised, lacerate, irregular, rather large : collar
indistinct, close : stem equal, darker than the peridium, lacerate
scaly, with a small mycelial bulb, white within, hollow or stuffed,
2—3 cm. long, 3 mm. in diameter : capillitium whitish, hyaline,
branched, septate, much swollen at the joints, 4—8 p. wide : spores
subglobose, yellowish cinnamon-colored, minutely verrucose, some
short pediceled, 3—5 p. in diameter. (PI. 34, f. 1—3.)

Kansas, Bartholomew ; Nebraska, Webber; Washington,
D. C, Braeneile. Growing in sandy pasture land.

Species not seen

16. Tylostoma semisulcatu.m Peck, Bull. Torr. Club, 22 :

209. 1895

" Peridium subglobose, usually a little longer than broad, 6-8
lines broad, 9 lines long, glabrous above, ferruginous-tomentose
on the lower half: osteolum entire : stem equal, about 2 inches
long, even and glabrous or but slightly furfuraceous on the upper
part, the lower part longitudinally sulcate, whitish : spores ferru-
ginous, globose, .00016 to .0002 in. broad: threads of the capil-
litium not septate.

"■ Sandy soil, Nevada. Collected by C. W. Irish, communi-
cated by T. Taylor.

"This species is separated from T. inanimosum Fr. by its peri-

White : Tvlostomaceae of North America 435

dium, which is tomentose on the lower half and not depressed,
and by its stem which is distinctly furrowed on the lower half."

As this species was not seen the above original description is
quoted. The same is the case with the following description of
T. Wrightii, and the species which Morgan calls " T. Mtycniannm."
It has been thought best to include these descriptions in order
that our present knowledge of the group may be accessible in a
single paper.

17. Tylostoma Wrightii Berk. Grevillea, 19: 95. 1891

"Stem 6 cm. high, 4 mm. thick, hollow, equal, ochraceous, even,
glabrous ; peridium spherico-depressed, 2 cm. broad, minutely
umbonate, pale, ochraceous, glabrous, the wall of the umbo dis-
appears at maturit)' and forms a small circular stoma ; mass of
spores yellowish brown ; threads of capillitium h)-aline, thick-
walled, aseptate, equal, very long, branched, axis lunate, 5 fi in di-
ameter : spores glabrous, globose, pale yellow brown, minutely
warted, 5-6 />« in diameter.

" On the ground. Rio Grande, North * Mexico (Wright).
Type in Herb. Berk.

" Distinguished from Tiilostoma Meycnianuni in the entire mouth
and the hollow, even and not striated stem."
Tylostoma sp. (Described by Morgan, Jour. Cin. Soc. Nat.

Hist. 12 : 163. //. 16. f. J, as T. Mcyeuiamnn Kl., but it is

clearly distinct from that species, though it might possibly be

referred to T. obcsiun C. & E.)

" Peridium depressed, globose ; the cortex soon receding, leav-
ing a smooth whitish or yellowish surface to the submembrana-
ceous inner peridium, the apex plane with a lacerate mouth :
stipe long, thick, unequal, fusiform or tapering, nearly solid, sul-
cate : threads of the capillitium long, much thicker than the
spores, branched, hyaline : spores subglobose, even, pale brown,
4.5-5.5 mic. in diameter.

"Growing in sandy soil. New Mexico, Wright; Colorado,

Webber. Plant 2-4 inches in height, the peridium 3^'-i inch in

diameter, the stipe about j^ of an inch in thickness at the thickest

part. Specimens referred to T. Angolense W. & C. do not differ

otherwise than in having the stipe thickest at the apex and taper-

* Probably a misprint for New as Wright was not known to have collected across
the Rio Grande.

436

White : Tylostomaceae of North America

ing downward instead of fusiform. T. obcsiim ,C. & E. appears
to be founded on a specimen with the short thick stipe not fully
developed."

Morgan's description was made solely from Webber's Colo-
rado specimen which was returned to Mr. Webber who is now
unable to find it. The New Mexican reference is merely a quo-
tation from Berkeley who determined Wright's original specimens
under this name.

The following tabulation will give more compactly our pres-
ent knowledge of the distribution of Tylostoma in North America :

Tylostoma.

albicans.

campestre.

exasperatum.

fibril losum.

fimbriatum.

gracile.

Kansense.

minutum.

obesum.

pedunculatum.

poculatum.

punctatum.

semisulcatum.

subfuscum.

tuberculatum.

verrucosum.

Wrightii.

^i S

rt

+

+

+

+

+

+

+

<\

11

ho a ;

C c(
•= 1 bo

+

■

+

+

+

+
+

1

+

+

+

+

+

+ +
+

+1
+1

+ +

+

+

+

O ^

C \ 13
U

+

i +
+

+ +,+!

+1

+
+

+

+

+

+

+

+

S

+

II. CHLAMYDOPUS Speg. An. Mus. Nac. Buenos Aires, 6 :

189. pi. 4. f. 2, J. 1899

In describing this genus Spegazzini includes two species from
Argentina, C. clavatus, from which we cannot separate our New
Mexican material, and C.Amblaiensis with a distinct annulus (?), and
up to the present no other species of this genus have been reported.
It is evident, however, from the description and figure of Tylostoma
Mcyeniamnn Kl.* that that species, also, belongs to the genus
CJilamydopus. While the figure shows no trace of a volva, its
tapering stem with its broad attachment to the peridium, together
with its general habit would seem to indicate its relationship to

*Nov. Act. Caes, Leop. Carol. Nat. Cur. 19 : 243.//. j./ 4. 1843.

White: Tylostomaceae of North America 437

this genus The volva is of such a friable nature that it would be
preserved with difficulty even if the stem had not been detached
by the inattentive collector. Quite recently several specimens
have been found at Mesilla Park, New Mexico, which belong here.
In going over the material referred to Tylostoma, in tiie Ellis
collection, two other specimens, from different localities in New
Mexico, were found which though destitute of volva, evidently be-
long to the genus Chlainydopus though they may constitute another
species. For the present they have been placed with C. clavatiis.

I. Chlamvdopus clavatus Speg. /f^. c/V.

Peridium depressed globose, 1. 5-2 cm. high, 2.2-3.5 cm. in
diameter, smooth, leathery, lightish fawn- colored : mouth plane,
lacerate, irregular: stem 8-15 cm. long, much enlarged at the top
where it joins the peridium, 1.5-3 cm. in diameter at the top,
0.5-1.5 cm. at the base, sulcate, lacerate scaly, firm, solid, of the
same color within and without as the peridium : volva friable, cup-
like, with flaring sides, 1.5-2.5 cm. high, having remnants of earth
and sand adhering to it: capillitium very abundant, interlaced,
lightish yellow, hyaline, branched, sparingly septate, not swollen
at the joints, the free ends usually rounded, 5-7 ji wide : spores
subglobose, dark ferruginous, densely verrucose, 4-6 // in
diameter. (PI. 35, f 1-6.)

New Mexico : Mesilla Park, Cockcrell, Mesa, Garcia, Las Cru-
ces, Wooton. Sandy soil under mesquite.

Aside from its technical characters this species presents a
habit very unlike any species of Tylostoina found in this country.
Its clean smoothish peridium and stem are quite in contrast with
the usual condition found in the species of Tylostoina, where the
rough outer peridium, and usually the stem also, is frequently
covered with fragments of adhering soil. Also in the specimens
of this genus, the capillitium threads are not as distinct as those of
Tylostoina and have considerable white amorphous hyaline tissue
intermingled with them.

III. BATTARREA Persoon,* Syn. Meth. Fung. 129. pi. 3. f.

i-j. 1 801
It has been impossible to trace the history of Battarrea to its
very beginning as the Acta Anglica, in which Woodward makes

* This genus was named for Antonio Battarra, an early Italian mycologist, and
was first erroneously spelled Batarrea by Persoon. In 1804 the name was changed by
Palisot de Beauvais to Baitarea, but the correct spelling, Battarrea, was not adopted
until 1825 by Fries.

438 White: Tylostomaceae of North America

first mention and gives the first drawing of this plant, has been in-
accessible, but as early as 1785 Dickson* described Lycopcrdon
phalloidcs — " volvulatum stipilatum, pileo deflexo campanulato ;
supra pulverulento calyptrato, infra glabro libero " — which unques-
tionably was the original Battarrca. It was collected in Septem-
ber by D. Humphreys and T. Woodward at Norwich, Norfolk and
Bungay, Suffolk, England. In 1801, Persoon established the genus,
and named and figured one species, B. phalloidcs, which he de-
scribes as follows — " volulata stipitata, Pileus deflexus, campanu-
latus,villosus, puluerisstrato obsitus, a voluacalyptratus." In 18 14
Liboschitz f made the next mention, when he described a plant
which he named Dcndromyces Stevenii, from the river Volga, which
Fries in 1839 referred to Battarrea. De Toni % gives eight species,
and Fischer § mentions seven of these, while quite lately, in 1 899,
Spegazzini || described two new species from Argentina, making
the number of described species ten in all.

The first Battarrea reported in this country was B. phalloidcs '^'^
in 1873, said to have been collected by J. Torrey in the vicinity of
San Francisco, Cal., but strangely enough, though this is the well-
known European species, it has never since been found, to my
knowledge, in the United States. Owing to the lack of description
of the plant mentioned there is a question as to whether this plant
really belongs to the species indicated. The only other species so
far known in this country are B. attcnuata from Nevada, described
by Professor C. H. Peck ft in 1895, and B. Digucti, described in
the following year by M. Patouillard ^;J from Lower California.
Lastly some material has been sent to Professor L. M. Under-
wood from New Mexico and Arizona which proves to belong to
two undescribed species.

A list of the hitherto known species with their localities is
added :

* Plant. Crypt. Brit, i : 24. 1785.

f Beschreibuni^ eines neuendeckten Pilzes. 1S14.

J Saccardo, Syll. Fung. 7 : 65 ; 9: 270 ; 14: 260.

\ Engler & Prantl, Die nat. Pflanzenfam. i*** : 344. 1900.

II An. Mus. Nac. Buenos Aires 6 : 190, 191. 1899.

** Grevillea 2 : 35. 1873.

ft Bull. Torr. Club, 22 : 208. 1895.

JJJour. de Bot. 10: 251.7: 2. 1886.

White: Tvlostomaceae of North America 439

B. PHALLOiDES Pcrs. England, Italy, North America (?).

B. Gaudichaudii Mont. Peru.

B. GuicciARDiNiANA Ces. Italy.

B. Tepperiana Ludw. Australia.

B. Stevexii (Libosch.) Fries, Siberia.

B. DiGUETi Pat. & Har. North America.

B. attenuata Peck. North America.

B. MuELLERi Kalchbr. Australia.

B. GuACHiPARUM Speg. Argentina.

B. Patagoxica Speg. Argentina.

S$yuop»ii» of the ^orfli American iipecies

Stem hollow.

Plants more than 20 cm. high ; volva double, the inner layers split into numerous

leaves. I- B. laciniata.
Plants 20 cm. high or less.

Spores minutely verrucose ; volva simple. 2. B. Griffilksii.

Spores nearly smooth ; volva triple. 3. B. Digiieti.

Stem solid. 4. B. aitemiata.

I. Battarrea laciniata Underwood, sp. nov.

Peridium 5-6 cm. in diameter, 2-3 cm. high, smooth, mem-
branaceous, the upper portion sometimes having portions of the
outer coat of the volva adhering to it : the lower portion after the
spores have been shed is a yellowish-white and shows traces of
where the capillitium has sprung : stem 25-35 cm. long, woody,
hollow, stuffed with silky, thread-like fibers running about half
way down the center of the stem, outer coat fibrous, pealing, some-
times merely lacerate, at others shaggy, slightly tapering to the
base, about 2.5 cm. at the top and 1.5 cm. at the base : volva com-
plex, composed of a thick outer coating, and an inner set of very
numerous thin leaflets arranged more or less in layers about the
stem : capillitium whitish, hyaline, indefinite, irregular, somewhat
fascicular: cells 30-50 ,« long, 5-7/-^ wide, with raised annular
thickenings or nearly flat spiral markings : spores subglobose,
reddish-brown, 5-7 11 in diameter, almost smooth. (PI. 36, f 1-6.)

New Mexico : Mesilla Park, Cockerell.

This plant resembles B. Digued in some of its characters, but
it is distinguishable (i) By its much larger size, (2) By its having
no inner woody volva enclosing the lower portion of the stem, (3)
In the much larger number of leaflets which compose the inner
portion of the volva, and (4) In the different coloring and marking

440 White: Tvlostomaceae of North America

of the cells. The plants vary considerally as to size and roughness
of the stem.

2. Battarrea Griffithsii Underwood, sp. nov.

Peridium 2-3.5 cm. in diameter, 1-2 cm. high, smooth, mem-
branaceous, lower part flat, showing the line of dehiscence dis-
tinctly : stem hollow, equal, 1-1.5 cm. in diameter, 9.5-15 cm.
long, sulcate, pealing fibrillose : volva fairly well marked, com-
posed of a few appressed fibrillose blunt squamules : capillitium
4-10/7. wide, whitish, flattened, rather amorphous, branched, and
having a large number of cells mixed with it : cells darker than
the capillitium, with spiral markings and annular thickenings,
6-8 IJ. wide, 24-85 jx long : spores subglobose, minutely verrucose,
looking smooth except under very high magnification, reddish -
cinnamon colored, 4—5/^ in diameter. (PI. 37, f 1—6.)

Arizona, David Griffiths, to whom the species is dedicated.

3. Battarrea Digueti Pat. & Har. Jour, de Bot. 10: 251. //.

2. f. 1-6. 1 896

Peridium depressed globose, membranaceous : stem 15-20 cm.
long, 8—10 mm. wide : spores globose, ferruginous, 5—6 a in
diameter : cells loo-i 50 a long, 4-7 // wide, with darker yellow an-
nular markings : volva of three layers, the outer one woody, simple,
whitish, the inner woody and simple, and encircling the lower third
part of the stem, the third layer between the outer and inner volva
is composed of from 10 to 20 thin membranaceous leaflets.

Lower California, Diguct. In barren rocky soil.

No specimens of this species have been seen, but it has been
thought best to quote the above partial translation of M. Patouil-
lard's very full description.

4. Battarrea attenuata Peck, Bull. Torr. Club, 22: 208. 1895

" Exoperidium unknown : endoperidium 2 in. or more in breadth,
the basal part hard, thick, even and concave beneath, convex
above, and somewhat coarsely reticulated by the bounding walls
of broad shallow pits : stem 8 to 10 in. long, gradually attenuated
toward the base, hard, almost woody, solid, rough except at the
top, with rather coarse spreading or rcflexed scales, brown ex-
ternally, rusty brown within : spores globose, ferruginous, .0003
in. broad : threads of the capillitium destitute of spiral thickenings.

" Plant commonly growing in tufts of 3-5 individuals. Dry
sandy soil. Nevada. Collected by C. W. Irish ; communicated
by Dr. Thomas Taylor."

White: Tvlostomaceae of North America 441

The single specimen of this species known is at Albany and is
inaccessible at the present time, so the above original description
has been quoted verbatim.

IV. OUELETIA Fries. Ofversigt Kongl. Vetens. Akad. For-
handlingar, Stockholm, \ji. pi. 4. 1871
This genus was established on a single species, and as yet no
others are known. It is very rare, having been reported from
very few localities. It was discovered by Dr. L. Quelet, at
Herimncourt, France, and has since been reported from the en-
virons of Rouen (Saint-Saens), and Pont de Sochaux, France.
It has been found only once in the United States, at Trexlertown,
Penn., by Mr. William Herbst, and was reported by Professsor C.
H. Peck, in his 46th report of the State Botanist. He is of the
opinion that this plant was introduced into this country in some
way with tan bark, on which it usually grows.

I. QuELETiA MiRABiLis Fries, loc. cit.

Peridium globose, 2.5-3.5 cm. high, 3-4 cm. in diameter,
fragile, easily separating from the stem, rupturing irregularly, of
a reddish-brown color : collar irregular, of the same substance as
the peridium : stem 6.5-8 cm. long, 1.5 cm. wide at the top, 2
cm. at the base, fascicular, reddish -brown, within and without, like
the peridium, solid, lacerate, fibrillose, particularly at the base :
capillitium very abundant and interwoven, forming with the spores
a felt-like mass, reddish-brown, single threads whitish-yellow,
thick-walled, hollow as shown by the truncated ends, septa rare
or wanting, 5-9 fi wide, branches rather short, free ends rounded
and recurved : spores subglobose, coarsely warted, 4-6 p. in
diameter, some short-pediceled, inner portion breaking up and
issuing from the thin-warted coating which is then hyaline and
shrivelled. (PI. 38, f. 1-5.)

Pennsylvania : Trexlertown, Herbst. On spent tan bark.
Summer, after rains, forming circles.

V. DICTYOCEPHALOS Underwood, gen. nov.

Plants with the irregularly rupturing peridium closely attached
to the solid stem. Volva cup-like, persistent at the base of the
stem. Gleba composed of a mesh-like irregular tissue, in which
the capillitium threads are embedded.

442 White : Tvlostomaceae of North America

I. Dictyocephalos curvatus Underwood, sp. nov.
Outer peridium of a thick woody texture, bearing a definite
cup-like volva at the base of the stem, the upper portion being
carried up on the peridium, and either faUing off or remaining
adherent to it : inner peridium scleroderma-like, rough, dark brown,
scaly, rather flattened sideways, rupturing irregularly, 3-6 cm.
high, 5-8 cm. in diameter: stem 25-40 cm. long, 3-6 cm. in
diameter at the summit, 1.5-4 cm. in diameter at the base, con-
siderably flattened, twisted, solid, dark brown within and without,
sulcate, the outer surface- very uneven, and peeling : the collar in-
distinct, formed by the lower portion of the peridium adhering to
the top of the stem and becoming torn as the stem elongates : the
top of the stem is rounded and projects into the peridium forming
a pseudo-columella, of a yellowish-brown color, lighter than the
rest of plant, marked with irregular, reticulated pits, from the sides of
which the mesh-like tissue springs which forms with the spore mass
the main part of the gleba : capillitium 8-10 /i wide, mostly embed-
ded in the mesh-like tissue, bright yellow, cylindric, septate, not
much swollen at the joints, branched, the free ends rounded :
spores subglobose, warty, 5-7 />. in diameter. (PI. 39, 40.)

Colorado : Colorow, BctJicl.

Plants with a strong odor which in the dry condition much
resembles that of the dried bark of Ulnius fiilva. The spores of
these specimens first appeared to be of two kinds — darker warted,
larger ones, and smaller, smooth, light colored ones — but it was
soon seen that this was owing to the outer coat having been eaten
off by the quantities of small insects by which these plants were
infested.

These strange plants were found by Mr. E. Bethel at Colorow,
Col., in the month of August, 1897. In the notes sent with these
specimens to Mr. p:ilis, Mr. Bethel says : " These plants are very
odd-looking in their native haunts ; they grow on a soft alkaline
adobe soil. Some of them had lifted themselves entirely out of
the ground, while others had the stalk standing in about one inch
of soil. They presented a very fantastic appearance, as there was
little or no other vegetation about. * * * Some of the specimens
were very much bent, approximating a semicircle, others were
twisted like a corkscrew, with the portions of the stalk split and
bent back. I think the chief factor in lifting the plant out of the
ground is this twisting and bending back of the portions of the
stem during dessication."

White : Tvlostomaceae of North America 443

Explanation of Plates

In most cases the drawings have been made twice natural size and re(hiced one
half. The capillitium and spores were drawn with a camera lucida under double the
magnification noted.

Plate 31

Fig. I. Tylostoma pedunailatum, nat. size.

Fig. 2. Spores and capillitium, X '^1^-

Fig. 3. Spores, X S'O-

Fig. 4, 5. Tylostoma albicans, nat. size.

Fig. 6. Spores and capillitium, X lyo-

Fig. 7. Spores, X 3 lo-

Fig. 8. Tylostoma verrucosuni, nat. size.

Fig. 9. Spores and capillitium, X ^1^-

Fig. 10. Spores, X 31°-

Fig. II. Tylostoma iiiintitiim, nat. size.

Fig. 12. Spores and capillitium, X i?^-

Fig. 13. Spores, X 3io-

Plate 32

Fig. I. Tylostoma gracile, nat. size.

Fig. 2. Spores and capillitium, X I70-

Fig. 3. Spores, X310.

Fig. 4. Tylostoma obesiiiii, nat. size.

Fig. 5. Spores and capillitium, X IJO-

Fig. 6. Spores, X 3iO-

Fig. 7. Tylostoma A'ansense, nat. size.

Fig. 8. Spores and capillitium, X ^7°-

Fig. 9. Spores, X 3^°-

Plate 33
Fig. I. Tylostoma exasperatiim, nat. size.
Fig. 2. Spores and capillitium, X i?^-
Fig. 3. Spores, X3io-
Fig. 4. Tylostoma fibrillosiim , nat. size.
Fig. 5. Spores and capillitium, X I70-
Fig. 6. Spores, X3io-
Fig. 7. Tylostoma tuberculatum, nat. size.
Fig. 8. Spores and capillitium, X I70-
Fig. 9. Spores, X 3io-
Fig. 10. Tylostoma campestrc, nat. size.
Fig. II. Spores and capillitium, X 170-
Fig. 12. Spores, X 310-

Plate 34 .

Fig. I. Tylostoma puiictatum, nat. size.
Fig. 2. Spores and capillitium, X I70-
Fig. 3. Spores, X 3io-
Fig. 4. Tylostoma poculatum, nat. size.

Fig. 5. Spores and capillitium, X I70- I

Fig. 6. Spores, X 3io-

444 White: Tylostomaceae of North America

Fig. 7. Tylostoma fimbrialum, nat. size.
Fig. 8. Spores and capillitium, X ^70-
Fig. 9. Spores, X S'O-
Fig. 10. Tylostoma subfiisctim, nat. size.
Fig. II. Spores and capillitium, X 170-
Fig. 12. Spores, X 31°-

Plate 35

Fig. I. Chlamydoptis clavatus, nat. size.

F"iG. 2. Spores and capillitium, X ^1^-

Fig. 3. Spores, X3iO-

Fig. 4. Single spores, X 45°-

Fig. 5. Peridium showing lacerate mouth, and enlarged stem, nat. size.

Fig. 6. Section through peridium and stem.

Pl.a.te 36
Fig. I. Battarrea laciniata, y, nat. size.
Fig. 2. Spores and capillitium, X ^T^-
Fig. 3. Spores, X 3i°-
Fig. 4. Cells, X V-^-
Fig. 5. Cell after desiccation, X 3 10.
Fig. 6. Peridium, nat. size, showing attachment to the stem,

Plate 37

Fig. I. Battarrea Griffithsii, nat. size.

Fig. 2. Specimen showing under side of peridium, volva missing, nat. size.

Fig. 3. Peridium showing method of dehiscence, nat. size.

Fig. 4. Spores, capillitium and cells, X ^TO-

Fig. 5. Spores, X 3io-

Fig. 6. Cells, X 3io-

Pl.\te 38

Fig. I. Qiieletia mii-abilis, nat. size.

Fig. 2. Spores and capillitium, X ^70.

Fig. 3. Spores, X 3io-

Fig. 4. Spores, showing breaking up of inner substance, X 3iO-

Fig. 5. Section through the peridium, showing detachment of stem, nat. size.

Plate 39

Fig. I. Dictyocephalos curvatus, ^ nat. size.

Fig. 2. Lower portion of stem, and double volva, ;4 nat. size.

Fig. 3. Peridium, ruptured irregularly, with remnants of mesh-like tissue, ^ nat.

size.

Fig. 4. Portion of tissue with spores and capillitium embedded in it, X ^1^-

Plate 40

Fig. 5. Dictyocephalos cui-oatus, i{ nat. size.

Fig. 6. Spore, X 465-

Fig. 7. Inner portion of spore issuing from outer coat, X 465-

Fig. 8. Empty, hyaline spore coats, X 465-

Fig. 9. Smooth inner spores after issuing from outer coat, X 465-

Bull. Torr. Hot. Club, 28.

Pl. 31-

V. S. \V. Del.

TYLO STOMA

THE HELIOTYPE PRINTING CO.. BOSTON.

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PL. ^2.

TYLOSTOMA

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Pl. 33.

TYLOSTOMA

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PL. 34.

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TYLOSTOMA

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Pl-35-

»

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CHLAMYDOPUS

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PL. 36.

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B ATTAR RE A

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Pl. 37-

.<"«'»^iS

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.p-f^^^.^ii^'IK^,^

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BATTARREA

THE HELIOTYPE PRINTING CO., BOSTON.

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PL. 38.

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%

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QUELETIA

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Pl. 39.

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DICTYOCEPHALOS

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CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 18

SHRUBS AND TREES

OF THE

SOUTHERN STATES.
I, II, III, IV.

BY JOHN" K. SMALL

NEW YORK
1901

I

Nl.

[Reprinted from the Bulletin of the Torrey Botanical Club.]

Shrubs and Trees of the Southern States.— I.

Bv John K. Small.

TsuGA Caroliniana Engelm. Coult. Bot. Gaz. 6 : 223. 1881.

I have already reported this rare hemlock from Georgia* and
can now record a second station, at Tallulah Falls, several miles be-
low the first one. At the second station the trees grow in a more
accessible position and reach a better development, there being
plenty of soil. The station is towards the lower end of the caiion,
on the south side, where the bank slopes at an angle of 45° or
more, and about 1000 feet above the river. Owing to a drenching
rain which prevailed during my stay at Tallulah I could not ascer-
tain the extent of this grove.

PiNus PUNGENS Michx. f. Hist. Arb. Am. i: 61.//. 5. 1810.
I can now report this most locally distributed of our mountain-
inhabiting pines for the flora of Georgia, having encountered it in
the lower part of the caiion at Tallulah Falls. For the same rea-
son given in the foregoing paragraph, I am unable to tell the size
of the grove, but it is extensive and the trees are larger than I
have seen them elsewhere in the southern mountains.

Salix Wardii Bebb, Gard. & For. 8: 363. 1895.
At the Falls of the Yadkin river in North Carolina both Salix
nigra and Salix Wardii are plentiful, the black willow growing
along the water's edge on the south side of the river, while Salix

*Bull. Torn Club, 22 : 45,

62

Wardii occurs on the opposite shore hardly two hundred yards
distant, the two species being respectively confined to the north
and south sides of the river. The form of Salix Wardii occurring
there is peculiar ; the bushes are smaller than any heretofore ob-
served, diffusely branched and the branches decumbent or almost
prostrate, spreading radially and forming large, tangled mats,
seldom rising more than six inches above the ground.

QuERCUS NANA (Marsh.) Sarg. Gard. & Forest, 8: 93. 1895.
I was much surprised to find quantities of this scrub oak on the
summit of King's and Crowder's Mountains near the southern
boundary of North Carolina in the summer of 1894. The locality
is within several miles of South Carolina and about one hundred
miles cast of the Blue Ridge.

QUERCUS RUBRA L. Sp. PI. 996. 1/5 3-

The existence of the red oak in Georgia was unknown to bot-
anists before 1893. In that year I discovered a few trees just
south of the North Carolina boundary, on the summit of the
Thomas Bald, at an altitude of 5200 feet. The trees were stunted
and irregular, as is characteristic at high altitudes. Last year,
however. I found a remarkable development of the species in the
northwestern corner of Georgia, in Catoosa county. The species
abounds in the limestone "bottoms;" trees three feet or more in
diameter are not uncommon, their trunks, naked often for seventy-
five feet from the ground, are so straight that it is impossible to tell
which way they will fall when cut off at the base. The thick bark
is more or less mottled, whence the local name " Leopard Oak."

Celtis MississiPPiENSis Bosc, Encycl. Agric. 7: 577. 1822.
Although extending over most of the western part of Georgia,
this species of hackberry reaches its greatest development in the
rich limestone " bottoms " in the region east of Lookout Moun-
tain. Gigantic trunks, three or four feet in diameter, are very
common, and are covered with innumerable corky warts, which
range from one to two or even three inches in height.

Darbyaumbellulata A.Gray, Am. Journ.Sci. (II.) i : 388. 1846.
I have lately discovered a new station for this rare plant. It

63

grows in limited quantities on the south banks of the Yellow River,
near McGuire's Mill, Gwinnett county, Georgia. I found it in
company with its near relative, Comandra lunbellata.

Magnolia tripetala L. Sp. PI. Ed. 2, 756. 1763.
Another species new to the flora of Georgia, apparently never
found so far southeast of the Blue Ridge. I first encountered
some trees at the northern base of Stone Mountain, and later at
several localities near the mountain. The trees are small and
slender, and the species does not thrive as it does farther north.

Crataegus eluptica Ait. Hort. Kew. 2 : 168. 1787.
On several occasions, while collecting between Tallulah Falls
and Toccoa Falls and in the Nacoochee Valley in northern Georgia
I have observed numerous groves of Crataegus elliptua growing
on the barren slopes of low hills, usually above streams, at alti-
tudes varying from icxx>-i500 feet. After several seasons'
study of this form in the field I can see no reason for uniting it
with Crataegus flava as a variety, as has lately been done.* Be-
sides characters in the habit, the leaves, the fruit and seeds, which
serve to separate it specifically from Crataegus fiava, I find an ap-
parent trustworthy distinction in the bark of the trunk. The
bark of Crataegus flava is black and in high narrow ridges, while
that of Crataegus elliptica is a light brown and in thin broad scales.

Crataegus rotundifolia (Ehrh.) Borck. in Roem. Arch i : Ft. 3,

87. 1798.
This species of Crataegus is very common in the southern Al-
leghanies and extends southward almost to the Gulf of Mexico.
In 1895 I found it throughout the southern part of Georgia, where
its favorite situation is the low ridges in the pine barrens, where
the different species of hardwoods abound.

Cliftonia monophylla (Lam.) Britton, Bull. Torr. Club, 16: 310.

1889.
This curious and local plant forms remarkably dense thickets
in the swamps and districts bordering streams in the vicinity of
the Altamaha river, especially north of Jesup, Georgia. The

* Silva of N. A. 4 : 1 14.

64

stems there range from one-half of an inch to one foot in diame-
ter, and the thickets they form remind one of those made by the
growth of Kalinia latifolia and Rhododendron niaxUnum on the
higher mountains of North Carolina.

Acer leucoderme Small, Bull. Torr. Club, 22: 367. 1895.

When first described this species was thought to be confined
to the middle country <ii the Southern States, but my extensive
journeys in Georgia last season brought to light two new stations
in the foothills of the Blue Ridge and the AUeghanies ; one is the
deep caiion just below the precipice of Toccoa Falls, the other a
shallow cation along the Little Chickamauga creek near Ringgold.
Both stations are similar to the original, and at both the tree holds
all its characters.

Acer Floridanum (Chapm.) Pax, Engler's Bot. Jahrb. 7: 243'

1886.
This characteristic maple is apparently very common in the
river swamps of the Flint River in southwestern Georgia Last
summer I encountered a remarkable growth just below Albany.
It probably follows the river to its mouth, for I again met it in the
vicinity of Bainbridge. The trees are conspicuous on account of
their close white bark and very dark foliage. Although said to be a
small tree 1 measured many trunks that were three feet in diam-
eter.

Vaccinium hirsutum Buckl. Am. Journ. Sci. 45 : 175. 1844.
In April, 1893,1 discovered this local and little-known huckle-
berry on the southern cliffs of the caiion at Tallulah Falls, Georgia.
This apparently is the first collection since the original discovery
by Buckley in Cherokee County, North Carolina. In 1894 Prof.
A. Ruth found the shrub on the Cade's Cove Mountains in eastern
Tennessee, thus adding the third station and the third State in
which the species is known to exist.

Shrubs and Trees of the Southern States.— II.

By John K. Small.
I. New and Noteworthy Species.

TsuGA Caroliniana Engelm. Coult. Bot. Gaz. 6: 223. 1881.
Last fall I received specimens of this very ornamental hemlock
from two new localities in North Carolina. Mr. A. M. Huger
found groves of it at Banner's Elk, Watauga County, at an eleva-
tion of 1 300 meters and in the Linville Gorge, Burke County, at
about 575 meters above sea-level, the latter station, together with
that at Tallulah Falls, Georgia, and the New River, Virginia, rep-
resenting the lowest altitudes at which the species has been found.

HicoRiA GLABRA (Mill.) Brittou, Bull. Torr. Club, 15 : 284. 1888.

Among the many unique things that Stone Mountain affords
are some dwarf hickory trees, usually less than two meters in
height, bearing quite an abundance of fruit.

438

QuERCus MINIMA (Sarg.)

Qiiercus virens var. dentata Chapm. Fl. S. States, 421. i860.
'^oX. Q. dentata V)Z.xXx . 1794.

Querais Virgi>iianav2L.r. ininhna Sarg. Silva N. A. : loi. 1895.

A low shrub forming wide patches by the extensive spreading
of the underground stems. Branches erect or ascending, less than
I meter tall, solitary or several together, simple, or branched
above; leaves firm, obovate or sometimes oblong to oblanceolate,
3-10 cm. long, acute or apiculate at the apex, repand-serrate, or
the upper ones sometimes entire, those of the shoots often lobed,
all glabrous or finely tomentose beneath, gradually or abruptly
narrowed into short petioles which vary from 2-5 mm. in length ;
staminate aments very slender, 1-4 cm. long, tomentose ; acorns
solitary or several at the ends of peduncles which vary from I-3
cm. in length, or sometimes sessile; cups turbinate hemispheric,
about 1.5 cm. broad, white-tomentose, the bracts appressed, thick-
ened on the back, except near its edge where they form a fringe ;
nuts ovoid or elliptic, 1.5-1.8 cm. long, dark brown, glabrous.

Sandy sterile pine barrens, Florida, chiefly near the coast.
Flowers in March and April ; matures its fruit in the fall.

This peculiar oak cannot pose as a variety of Quercus Virgini-
ana under any reasonable considerations. It may be of interest
to note that it bears much the same relation to Querais Virginiana
as Castanea nana does to Castanea pnmila or Castanea dentata.
The habit of Quercus minima, with its underground stems, and
low erect branches which are usually much less than one meter in
height, is enough to separate it specifically from the gigantic forest
tree Quercus Virginiana. In addition to the differences in habit
just mentioned, the leaf types are characteristic and the nerves in
the leaves of Quercus minima are much more prominent than they
are in the live oak. The cups seem to furnish a diagnostic char-
acter, those of the Quercus minima being of a turbinate type, while
those of Quercus Virginiana are hemispheric.

Quercus geminata n. sp.

A shrub or small tree, 2-5 meters tall, with a maximum trunk
diameter of about 15 cm. Leaves narrowly oblong, elliptic, or ob-
long-oblanceolate, 3-6 cm. long, entire, obtuse or apiculate,
strongly revolute, mostly gradually narrowed at the base, glab-
rous and parchment-like above, finely tomentose and conspicu-
ously rugose by the prominent nerves beneath ; petioles 2-6 mm.

439

long; flowers not seen ; acorns usually 2 at the end of a pedun-
cle, which varies from I-4 cm. in length ; cups turbinate, i cm.
broad, tomentose, the bracts appressed, slightly thickened near
the base of the cup, fringed at the edge ; nuts ovoid or narrowly
oval, 1-1.7 cm. long, twice or thrice as long as the cups.

Sandy soil, chiefly in the scrub, Florida. Flowers in spring
and matures its fruit in the fall.

Mr. Nash, who collected and observed this plant during the
seasons of 1894 and 1895, assures me that it is perfectly distinct
from its relatives. This is doubtless a fact, and both the foliage
and fruit furnish excellent characters. The very prominently ru-
gose lower leaf-surfaces and the strongly revolute leaf-margins have
no parallel in Qtierais Virginiana. The acorns are always borne in
pairs at the ends of short stout peduncles ; the turbinate cups
with their constricted bases are diagnostic.

Celtis Georgiana n. sp.

A diffuse shrub with slender often 2- ranked branches, the leafy
twigs more or less pubescent. Leaves ovate, 2-5 cm. long, aver-
aging 2.5 cm. in length, or those on vigorous shoots sometimes
6 cm. long, acute, entire or sharply serrate above the middle, in-
equilateral, rounded or truncate at the oblique base, dark green,
scabrous and occasionally sparingly pubescent above, paler and
glabrous beneath, except for a few hairs on the nerves; petioles
1.5-4 nim. long, pubescent; pedicels usually slightly curved, 1.5-
4 mm. long, pubescent; drupes subglobose, sometimes broader
than long, 6-7 mm. in diameter, tan-color, smooth and glabrous,
or sometimes glaucous ; seeds obovoid-globose.

Along or near streams, north-central Georgia. Flowers in the
spring ; matures its fruit in September.

Collected by the writer, first in the Yellow River Valley, near
McGuire's Mill, Gwinnette County, in 1893, and in succeeding
years at many points about Stone Mountain and the contiguous
region.

A low species related to Celtis puinila, from which it may be
distinguished by its smaller merely acute leaves, the very short
pedicels and the smaller tan-colored drupes.

Celtis Helleri n. sp.

A much branched, wide spreading tree, sometimes lo meters
tall with a maximum trunk diameter of 1. 5 meters. Bark of the

440

trunk and main branches with numerous corky warts ; leaves
rather firm, the blades ovate to oblong, 4-7 cm. long, obtuse or
acute, crenate-serrate, especially above the middle, rounded or
subcordate at the base, deep-green and scabrous-pubescent above,
pale and tomentose beneath, slightly inequilateral, oblique at the
base ; petioles stout, 3-4 mm. long, tomentose ; pedicels sparingly
pubescent, curved, 1-1.5 cm. long; drupe subglobose, 7-9 mm. in
diameter, light-brown, translucent, smooth and shining ; seeds
globose, strongly 4-ribbed, prominently reticulated.

In dry ground near San Antonio, Texas.

A rather low tree with a short stout trunk varying from .5-1.5
meters in diameter, and a wide spreading top. The branches are
numerous and bulky. The original specimens were gathered by
Mr. Heller from trees growing in a strip of woodland between
the city of San Antonio and the San Antonio River, Texas, no.
1587.

ToxYLON POMIFERUM Raf. Am. Month. Mag. 2: 118. 1817.
Years ago the osage orange was planted on Paris Mountain,
South Carolina, for hedges and for ornamental purposes. For
many years the settlements have been neglected and deserted and
the tree has spread and established itself in an astonishing manner,
now appearing as if indigenous.

Albizzia Julibrissin Durazz. Mag. Tosc. 3: 11. 1772.
Although not indigenous, this tree now appears as if it were
native in the southern states. It grows along roadsides and here
and there through the pine woods much as the honey locust {Gled-
itsia triacanthos) does in many localities. It ranges from North
Carolina to Georgia, Florida and Alabama, where Prof. Underwood
collected specimens during the past summer. It is quite abundant
in southern Georgia.

Amorpha virgata Small, Bull. Torn Club, 21 : 17. //. 17 J- 1894,
In the spring of 1896 Dr. Charles Mohr sent me a specimen of
Amorpha virgata from the mountains of Madison county, Alabama,
thus extending the known geographic range of the species from
Stone Mountain, Georgia, to northern Alabama. Dr. Mohr gives
the altitude of this locality as 350 meters. While collecting at
different points along the eastern section of the Blue Ridge dur-

441

ing the summer of 1896, I was surprised to find the species both
on the slopes and summit of Paris Mountain near Greenville, and
on the slopes of Table Mountain. At the former locality it oc-
curred at an altitude of about 500 metres, and on Table Mountain
it ranged from 800-900 meters. Its characters hold perfectly.

LoNiCERA FLAVA Sims. Bot. Mag.,//. 1318. 1 8 10.

About two years ago I recorded* several new localities for this
handsome honeysuckle. Further exploration of the southern end
of the Blue Ridge has revealed additional stations. In the sum-
mer of 1894 I found some bushes on the upper slopes of Currahee
Mountain, an isolated peak nearToccoa, Georgia, and a little later
noticed several bushes on Stone Mountain. During the past sum-
mer I collected it on Paris Mountain, South Carolina, the original
locality, where it grows at several points along the rocky summit,
and later discovered a new station on the precipitous cliffs of Table
Mountain, in the same state. At the latter place the shrubs were
more robust and vigorous than at any of the other stations.

The finest flowering specimens I have ever seen were sent me
by Mr. A. M. Huger, who secured them on Tyron Mountain, Polk
County, North Carolina, last spring. Mr. Huger's discovery ex-
tends the range of the species into another state, but although we
now have specimens showing the species to range from North
Carolina to Georgia, it is not common at any of the localities, a
few bushes only existing at the different places,

II. The Genus Gaylussacia in the Southern States.

During several seasons I have had ample opportunity to study
this imperfectly understood group in the field and have made ob-
servations on all except one of the species recognized in the ap-
pended revision. As far as I can see, the forms hitherto con-
sidered as varieties of other species are abundantly distinct and
should be treated as species. Mr. Nash came to the same con-
clusion during his field-work in Florida. The diagnostic char-
acters are brought out in the following key and descriptions.

* Bull. Torr. Club, 21 : 305.

I.

G.

. dumosa.

2.

G.

hirtella.

3-

G.

nana.

4-

G.

tomentosa

5-

G.

frondosa.

6.

G.

ursina.

7.

G.

resinosa.

442

Corolla campanulate or globose campanulate ; leaves destitute of sticky resin
Stems horizontal, underground, the branches erect.
Pubescence consisting of gland-tipped hairs.
Twigs and racemes pilose.
Twigs and racemes bristly-hispid.
Pubescence consisting of simple non-glandular hairs.
Leaves glaucous, glabrous or nearly so.
Leaves densely tomentose, especially beneath.
Stems erect, the branches spreading.

Leaves leathery, obtuse or retuse ; drupe glaucous.
Leaves thin, acuminate and apiculate ; drupe black.
Corolla conic; leaves sticky with a resinous secretion.

I. Gaylussacia dumosa (Andr.) T. & G.

Vaccinhnn diimosum Andr. Bot, Rep. 8: ii2. 1794.
Gaylussacia dumosa T. & G.; A. Gray, Man. 259. 1848.

A low shrub, 1-5 dm. tall, with underground stems and erect
solitary or tufted branches ; the twigs, leaves and inflorescence
glandular-pilose. Leaves leathery, the blades oval, obovate or ob-
lanceolate, rarely linear-oblanceolate, 2-4 cm. long, apiculate at
the apex, ciliate, short-petioled, deep green above, paler beneath ;
calyx glandular, about 5 mm. broad, the segments triangular or
triangular ovate, acute, about as long as the tube ; corolla cam-
panulate, 5-6 mm. long, white or pink, wax-like, the segments
broadly ovate, more or less recurved and revolute; filaments short,
pubescent ; anthers longer than the filaments, prolonged into fili-
form tubes ; drupe globose, black, 6-8 mm. in diameter, commonly
somewhat pubescent.

In sandy soil, Newfoundland and along the coast to New
York, south to eastern Pennsylvania, North Carolina, Florida and
Louisiana. Spring ; matures its fruit in the summer.

2. Gaylussacia hirtella (Ait.) Klotzsch.

Vaccinmm hirtellinn Ait. Hort. Kew. Ed. 2. 2: 357. 181 1.

Gaylussacia hirtella Klotzsch, Linnaea, 14: 48. 1840.

Gaylussacia dumosa var. hirtella A. Gray, Man. 259. 1848.

A shrub, with underground stems, the branches, twigs and in-
florescence bristly-hispid, the tips of the hairs with minute glands ;
leaves firm, the blades oblanceolate-spatulate or elliptic, 3-6 cm.
long, apiculate, glandular-ciliate sparingly hispid above, short-
petioled ; racemes many-flowered ; calyx hispid, 6 mm. broad, the
segments triangular, rather acuminate, about as long as the tube;
corolla broadly campanulate, 7-8 mm. long, the segments broader
than long, the tips recurved, the edges revolute ; filaments short,

443

pubescent ; anthers longer than the filaments, prolonged into fili-
form tubes; drupe not seen.

In sand, Florida to Louisiana. Spring; fruit ripe in the
summer.

Certainly distinct from Gaylussacia diimosa, from which it dif-
fers in habit, size and leaf characters. The pubescence is always
diagnostic, the corolla is larger and much thinner than that of G.
dumosa, while the calyx- segments are longer and usually acumi-
nate.

3. Gaylussacia nana (A. Gray).
Gaylussacia frotidosa var. fiana A. Gray, Syn. Fl. N. A. Ed. 2.

2: Ft. I, 396. 1886.

A low glaucous shrub I-4 dm. tall, spreading by underground
stems. Leaves leathery, the blades elliptic, obovate or nearly
spatulate, 2-3 cm. long, obtuse or minutely apiculate at the apex,
glaucous on both sides, becoming bright green above, prominently
rugose and sprinkled with amber-colored resin beneath, short-
petioled ; racemes few-flowered ; pedicels slender, puberulent when
young; calyx glabrous, 3 mm. broad, the segments triangular,
acute, about as long as the tube; corolla globose-campanulate,
3 mm. long, the segments ovate, acutish, longer than broad; fila-
ments short, glabrous ; anthers longer than the filaments, pro-
longed into slender tubes ; drupes subglobose, 6-7 mm. in
diameter, rather dry, glaucous.

In sandy pine barrens, Georgia to Florida and Alabama.
March to April; matures its fruit in the summer.

Easily distinguished from Gaylussacia frondosa, with which it
has been associated, by its very glaucous foliage and strongly
rugose and much smaller leaves, besides its peculiar underground
stems.

4. Gaylussacia tomentosa Pursh.
Gaylussacia frondosa \2x. tomentosa A. Gray. Syn. Fl. N. A. 2 :

Ft. I, 19. 1878.

Gaylussacia tomentosa Pursh ; A. Gray. Syn. Fl. N. A. 2 : Pt.
1,19. As synonym. 1878.

A low shrub, spreading by underground stems, the foliage
ton entose with brownish hairs. Leaves leathery, the blades
oblong or elliptic, often slightly broadest above the middle, 2.5-7
cm. long, obtuse and apiculate at the apex or sometimes notched,
brow n-tomentose on both sides, densely so beneath, short-petioled;

444

racemes few-flowered ; pedicels 1-1.5 cm. long, much longer than
the bracts ; calyx glabrous, about 3.5 mm. broad, the segments
ovate, acute, about as long as the tube; corolla campanulate, 3.5
mm, long, the segments ovate, obtuse, about as long as broad, the
tips recurved, the edges revolute; filaments dilated, glabrous;
anthers longer than the filaments, prolonged into slender tubes;
drupes depressed-globose, 8-9 mm. in diameter, glaucous.

In sandy soil, Georgia and Florida. Spring ; matures its fruit
in the summer.

Like the preceding species, Gaylussacia tomentosa has under-
ground stems, but it differs from it in the brown-tomentose foliage,
more robust habit, larger leaves and different leaf-form. The fruit
of G. tomentosa is larger and much more fleshy than that of G. nana.

5. Gaylussacia frondosa (L.) T. & G.

Vacciniiim frondosiim L. Sp. PI. 351. 1753-

Gaylussacia frondosa T. & G. ; Torr. Fl. N. Y., i : 449. 1843.

An irregularly branched shrub 1-2 meters tall, with puberulent
twigs and young leaves. Leaves firm, the blades oblong-oblan-
ceolate, ovate, oval or obovate, obtuse or notched at the apex,
delicately revolute, short-petioled, bright green and glabrate above,
glaucous and sprinkled with minute golden globules of resin be-
neath ; racemes loose; pedicels long and slender; calyx gla-
brous, 3-4 cm. broad, the segments triangular, acute or acutish,
about as long as the tube ; corolla globose-campanulate, about 4
mm. long, green to purplish, the segments triangular, broa.der
than long, recurved and revolute ; filaments dilated, glabrous ; an-
thers longer than the filaments, prolonged into slender tubes;
drupe globose, 8-10 mm. m diameter, with a pale bloom.

In sandy soil. New Hampshire, south to Florida, Kentucky
and Louisiana. Spring ; matures its fruit in the summer.

6. Gaylussacia ursina (M. A. Curtis) T. & G.

Vaccinium ursinum M. A. Curtis, Am. Journ. Sci. 44 : 82. 1843.

Gayhissacia ursina T. & G. ; A. Gray, Mem. Am. Acad. (II.)
3: 49. 1846.

A straggling branching shrub, 6-1 5 dm. tall, with sparingly
pubescent twigs and young foliage. Leaves thin, the blades ob-
long, elliptic or oblanceolate, usually rhomboidal, 4-iOcm. long,
usually short-acuminate; apiculate, ciliate, deep green above, paler
beneath, pubescent on the nerves on both sides, obtuse or rounded
at the base, short-petioled ; flowers few, in lateral somewhat droop-
ing racemes ; calyx with numerous golden glands, about 3 mm.

445

broad, its 5 segments very low, obtuse, several times shorter than
the tube ; corolla globose-campanulate, greenish-white or tawny-
red, about 4-5 mm. long, its segments triangular, acutish, recurved,
revolute ; filaments dilated, pubescent, incurved at the apex, longer
than the anthers which have short tubes at the apex ; drupe glo-
bose, 10-12 mm. in diameter, black, shining, sweet.

In deep forests on the mountains. North Carolina to northern
Georgia. Spring ; matures its fruit in the late summer,

7. Gaylussacia resinosa (Ait.) T. & G.

Vacciniiim rcsinosiun Ait. Hort. Kevv. 2: 12. 1789-

Gaylussacia resinosa T. & G.; Torr. Fl. N. Y. i : 449. 1843.

A rigid branching shrub 3-12 dm. tall, its twigs and foliage
more or less pubescent and sticky with a resinous secretion when
young, leaves firm, the blades elliptic, oval or oblong, sometimes
broadest above the middle, firm, obtuse or apiculate, entire, cilio-
late, short petioled ; flowers in lateral drooping racemes ; pedi-
cels 2-8 mm. long, usually with two narrow bracts; calyx about
2 mm. broad, its 5 segments ovate, obtuse, about as long as the
tube ; corolla obconic, red or reddish-green, 5-6 mm. long, more
or less constricted near the apex, the segments ovate, spreading or
recurved, revolute, obtuse ; filaments winged, pubescent, shorter
than the anthers, each cavity of which is prolonged into a tube ;
drupes globose, 6-10 mm. in diameter, black or rarely white,
sweet.

In rocky woods and hillsides, Newfoundland to the Saskatche-
wan, south to Georgia. Spring; matures its fruit in the summer.

Shrubs and Trees of the Southern States.— III.

By John K. Small
1. HITHERTO UNDESCRIBED SPECIES

Prunus Cuthbertii

A tree 6 m. tall and sometimes 1.5 dm. in diameter near the
base, with tomentose twigs. Leaf-blades leathery, normally obovate,
varying to oval or fiddle-shaped, 4-9 cm. long, blunt or notched
at the apex, shallowy serrate, not markedly biglandular at the
often cuneate base, dull green above, pale or glaucescent beneath,
the midrib and petioles copiously tomentose and the lateral veins
slightly so : racemes 5-8 cm. long, terminating short leafy
branches, the rachis and pedicels pubescent like the twigs : pedicels
clavate, 3—5 mm. long during anthesis, becoming 8— ii mm.
long : sepals broader than long, obtuse, shorter than the hypan-
thium : petals suborbicular, about 2 mm. in diameter, crisped :
drupes subglobose, mostly 8-9 mm. in diameter, red.

In rich woods, near Augusta, Georgia.

A relative of Priimis scrotina differing in the decidedly obovate
blunt leaf-blades and the tomentose glandless petioles and tomen-
tose veins of the blades and the similarly pubescent racemes. The
flowers are smaller and the very blunt sepals are much broader
than the acute sepals of P. serotina. The fruit also furnishes a
character in its decidedly red color. The original specimens were
collected by Mr. A. Cuthbert in the vicinity of Augusta, Georgia,
on July 17, 1898, and June 2, 1899. Type in the herbarium of
the New York Botanical Garden.

Leucothoe platyphylla

A straggling shrub 0.5—1 m. tall, with spreading and drooping
branches. Twigs and branches finely pubescent : leaf-blades sub-
orbicular to oval, broadly oblong or oblong-ovate, 5—8 cm. long,
abruptly pointed or slightly acuminate, serrate with • very fine
spinulose teeth, especially above the middle : petioles stout, 4—8
mm. long, pubescent : racemes 2—5 cm. long, the rachis pubes-
cent like the twigs : pedicels 3—5 mm. long : calyx glabrous ;
longer sepals orbicular-ovate : corolla white, 6—8 mm. long : cap-
sule 5.5 mm. broad, 4 mm. high.

290

291 Small : Shrubs and Trees

In low thickets, Georgia to Alabama. Spring.

Leucothoe platypliylla differs from L. axillaris in its broader
leaf-blades, orbicular larger sepals, smaller corollas and larger cap-
sules.

The specimens on which the species is founded were collected
by the writer near the Ochlockonee River about Thomasville,
Georgia, in May and June, 1895. Type in the herbarium of Co-
lumbia University.

Hypericum splendens

An erect much branched shrub 0.5-1.5 meters tall, with 2-
edged glaucous twigs. Leaves firm, numerous ; blades oblong,
1.5-2.5 cm. long, obtuse or rounded at the apex, slightly revolute,
glaucous, especially beneath, narrowed at the sessile base : buds
conic : flowers very showy, several or many in terminal or axillary
cymes : sepals unequal, oblong, firm, apiculate, the outer about 8
mm. long, the inner 5 mm. long: corollas 3.5-4 cm. broad:
petals golden, cuneate, oblique, 1.5-2 cm. long: stamens very
numerous, orange-colored: styles 3, slender: capsules often
crowded, conic, 1.5 cm. high, acuminate at the apex, partially 3-
celled, about thrice as long as the sepals with 3 narrow wings and
6 obtuse angles: seeds cylindric, 1.3 mm. long, longitudinally
ribbed and transversely wrinkled, resembling ears of corn.

On granite slopes. Stone Mountain, Georgia. Summer.

This is perhaps the most showy North American Hypericum
and is related to H. ajirewn from which it differs in the smaller
firmer leaves, the more distinctly pedicelled flowers and the conic
buds.

The original specimens were collected by the writer on Stone
Mountain, Georgia, in July and August, 1893, and August 6,
1895. Type in the herbarium of Columbia University.

2. THE GENUS PTELEA IN THE SOUTHEASTERN UNITED STATES

PTELEA L.
The genus Pteka differs from Helictta, its only close relative in
North America, in its pubescent filaments, and in the fruit which
is winged all around and does not separate into distinct carpels.

Key to the Species

Parts of the flower usually in 5's : leaflets relatively small.

Filaments slightly pubescent at the middle : petals glabrouj within : Floridian.

I. y. Baldwinii.

OF THE Southern States 292

Filaments pubescent throughout : petals pubescent within : Texano-Mexican.

2. P. angtistifolia.
Parts of the flower usually in 4's : leaflets relatively large.
Samaras obovate.

Leaflets with sharply serrate blades : samaras 16-19 '"'"• lo"g> acute at the

base, the body merely glandular-dotted. 3. P. serrata.

Leaflets with entire or merely undulate blades: samaras 8-1 1 mm. long, or
rarely longer, rounded or truncate at the base, the body pitted.

4. /-". mici-ocarpa.
Samaras suborbicular.

Leaflets with entire or merely crenulate blades, the terminal one much longer

than broad, acute or slightly acuminate. 5- P- t>'ifoliata.

Leaflets with crenate or crenate-lobed blades, the terminal one only slightly
longer than broad, blunt.
Blades of the leaflets not lustrous : filaments nearly glabrous : anthers

rounded or retuse at the apex. 6. P. ^'Jumibifolia .

Blades of the leaflets lustrous above : filaments densely pubescent below :
anthers apiculate. 7. P. Toxicodendron.

I. Ptelea Baldwinii T. & G.
A shrub about 3 dm. tall, with irregularly branched stems.
Leaflets 3 ; blades oval or ovate, 1-2 cm. long, obtuse at both
ends, or the terminal one cuneate at the base, sessile, glabrous
except the midrib and the ciliate margins when they are young :
panicles few-flowered : flowers with the parts mostly in 5's: calyx
1.5 mm. broad: sepals oval, less than i mm. long, acutish, ascend-
ing : petals 4, oblong-oblanceolate, 4 mm. long, obtuse, undulate :
stamens shorter than the petals : filaments stout, hairy at the middle.

In the vicinity of St. Johns, eastern Florida. Spring.

2. Ptelea angustifolia Benth.
A little known species originally described from Mexico, with
pubescent foliage, wholly pubescent filaments and pubescent
petals, is said to occur in southern and western Texas.

3. Ptelea serrata sp. nov.

An irregularly branched shrub, 1-2 m. tall, with glabrous
foliage. Leaflets 3 ; blades thinnish, oval, elliptic or elliptic-obo-
vate, 2.5—7 c^- ^o^g. sharply acuminate, or rarely only acute,
rather shallowly but sharply serrate, deep green above, very pale
green beneath, the terminal one with a slender base : panicles
few-flowered: samaras obovate, 16-19 n^m- long, acute at the
base, the wing rather delicate, the body glandular-dotted.

On granite slopes, Stone Mountain, Georgia. Spring.

293 Small: Shrubs and Trees

Ptelea serrata is not closely related to any described species. It
differs from the other species in the sharply acute or acuminate and
sharply serrate blades of leaflets and the obovate samaras with their
acute bases.

The specimens on which the species is founded were collected
by the writer on Stone Mountain, Georgia on July 3, 1893. Type
in the herbarium of Columbia University.

4. Ptelea microcarpa sp. no v.

A shrub 1.5-3 m. tall, with branching stems and glabrous
foliage. Leaflets 3 ; blades firm, elliptic, oval or oblong -lanceo-
late, 4-10 cm. long, bluntly pointed or acutish, often slightly
acuminate at both ends, entire or nearly undulate, slightly paler
beneath than above, the terminal one not conspicuously narrowed
at the base : panicle many-flowered : samaras suborbicular or or-
bicular-obovate, 8-1 1 mm. in diameter, or rarely larger, rounded
or truncate at the base, the wings slightly crisped, the body pitted.

On limestone or granite ridges, Tennessee, Georgia and Ala-
bama. Spring.

Ptelea microcarpa is most closely related to P. serrata, but
differs in the firmer longer entire or merely undulate blades of
the leaflets, and the samara with its pitted body, and the wing
which is rounded or truncate at the base.

The original specimens were collected by officers of the Bilt-
more Herbarium (no. 4437''), at Nashville, Tennessee, August 9,
1897. The type is in the herbarium of the New York Botanical
Garden.

5. Ptelea trifoliata L.

An aromatic shrub or tree sometimes 8 m. tall, the foliage
glabrous or sometimes densely pubescent. Leaflets 3 ; blades
ovate, oval, elliptic, oblong, oblong-lanceolate or oblanceolate,
4-12 cm. long, acute or acuminate, entire or sometimes undulate
or partially crenulate : panicles many-flowered: sepals ovate, 1.5
mm. long, obtuse : petals narrowly oblong, 4-5 mm. long : sam-
aras suborbicular or oval-orbicular, 2-2.5 cm. long, rounded or
notched at the base.

In rich soil and on river banks. Long Island to Ontario, Min-
nesota, Florida and Texas. Spring. The form with pubescent
foliage is Ptelea trifoliata mollis M. A. Curtis and is most common
in the Gulf States.

OF THE Southern States 294

6. Ptelea rhombifolia Heller

A shrub 2-2.5 ^- t^iH. branching above, the foliage densely
pubescent. Leaflets 3 ; blades rhombic-ovate to rhombic-orbicu-
lar, 2.5-5 cm. long, blunt, crenate, dull green above, the terminal
one but little longer than broad : petals pubescent without, about
4 mm. long : filaments slightly pubescent near the base : samaras
nearly orbicular, 1.5-2.5 cm. broad.

In open woods, southern Texas. Spring.

7. Ptelea Toxicodendron sp. nov.

A branching shrub 1-2 m. tall, with glabrous foliage. Leaflets
3 ; blades oval or rarely oval-ovate, 1.5-6 cm. long, rounded or
blunt at the apex, crenate or somewhat crenate-lobed, dark green
and lustrous above, slightly paler beneath : panicles few-flowered :
petals about 5 mm. long, glabrous : filaments densely pubescent
below : mature fruit not seen.

In gravelly soil, near Kerrville, Texas. Spring.

A species most closely related to Pu/ea rhouibifolia, but easily
distinguished by its glabrous or nearly glabrous leaflets whose
blades are lustrous above, the filaments which are densely pubescent
below and the apiculate anthers.

The original specimens were collected by Mr. A. A. Heller at
Kerrville, Texas, and distributed in his Plants of Southern Texas,
under no. 1690. Type in the herbarium of Columbia University.

Shrubs and Trees of the Southern States.— IV.

By John K. Small
1. THE EBENACEAE IN THE SOUTH

Two types with as divergent characters as those possessed by
Diospyros Virgi7iiana and the so-called D. Tcxana, cannot with the
least degree of systematic order, be referred to the same genus.
The two plants are of wholly different habit, and the flower-struc-
ture of the two is so distinct that I cannot understand on what
grounds they have been associated with each other in the same
genus. The distinguishing characters of the two trees, which I
shall treat as two distinct genera, may be compared by means of

the following synopsis :

EBENACEAE

A family of about 6 genera and 275 species, most abundant in
tropical regions. Only the following representatives are now
known to occur in the United States.

Key to the Genera

Styles distinct : anther-sacs opening by longitudinal slits : filaments pubescent : pistillate
flowers without starainodia. I. DiOSPYKOS.

Styles united : anther-sacs opening by subapical pores ; filaments glabrous : pistillate
flowers with 8 staminodia. 2. Brayodendron.

I. DIOSPYROS L.

I. D. ViRGINIANA L. Sp. PI. 1057. 1753

In woods and fields, Rhode Island to Kansas, Florida and

Texas. As now limited this species may be an aggregate. One

or two additional species may be separable in the south Atlantic

and Gulf States.

2. Brayodendron *

I. B. Texanum (Scheele)
Diospyros Texana Scheele, Linnaea, 22: 145. 1849.
Along or near streams in river valleys, southern Texas and
adjacent Mexico.

* Named for Prof. W. L. Bray, head of the School of Botany, University of
Texas.

356

357 Small: Shrubs and Trees of the Southern States

2, HITHERTO UNDESCRIBED SPECIES
Quercus microcarya
A shrub, or a small tree sometimes 5 m. tall, with very smooth
branches and glabrous foliage. Leaves deciduous ; blades thin,
spatulate in outline, 5-10 cm. long, undulate or shallovvly lobed,
o-ray-green above, yellowish green beneath, short-petioled, the
small bristles terminating the lobes early deciduous : acorns sessile,
less than 10 mm. long ; cup deep saucer-shaped, 6-"] mm. broad ;
nut globose-ovoid, often nearly one-half included in the cup.
On granite outcrops. Little Stone Mountain, Georgia.
Related to Q. nigra, but more delicate in all its parts. The
leaf-blades are relatively thin and more irregularly lobed, while
the very small acorns, always, as far as I have observed, less than
10 mm. in length, with their relatively deep saucer-shaped cups,
are very distinct from the larger fruit of Q. nigra with its very
shallow cup. The fruit is much smaller than that of any of the
species of the group to which this plant belongs.

The original specimens were collected by the writer on Little
Stone Mountain, Georgia, September 1 1, 1894. Type in the her-
barium of the New York Botanical Garden.

Quercus fusiformis
A shrub 1.5-3 m. tall, with slender or switch-like stems or
branches. Leaves persistent; blades leathery, oblong or ovate-
oblong, 3-5 cm. long, entire or sparingly spiny-toothed on the
twigs, gray-green and lustrous above, paler and scurfy-tomentose
beneath, slightly revolute, rounded or truncate at the oblique base,
short-petioled : acorns numerous, solitary or several on slender
peduncles; cup turbinate, 10-12 mm. high, gray; nut fusiform,
2-2.5 cm. long, long, acute, conspicuously striate, thrice surpass-
ing the cup.

On arid limestone and granite hills, central Texas.
Quercus fusiformis belongs to the group formed by Q. minima,
O crcminata and Q. Virginiana. It is most closely related to
Q. Virginiana, but differs in the much elongated acorn with its
turbinate cup and fusiform nut.

The original specimens were collected on Lacey's Ranch near
Kerrville, Texas, by Mr. Howard Lacey during the years 1899-
1900, and given to me by Professor W. L. Bray, of the University
of Texas. Locally known as Live Oak. Type in the herbarium
of the New York Botanical Garden.

Small: Shkucs and Trees of the Southern States 358

Quercus Laceyi

A shrub or small tree, becoming 6 m. tall, with a rough deeply
and irregularly grooved bark. Leaves deciduous but rather tardily
so, numerous ; blades oblong and with 3-5 shallow lobes, or ob-
long-obovate and more prominently 3-lobed below the apex, 4-8
cm. long, olive-green and with a waxy lustre above, grayish and
slightly and minutely scurfy beneath, sometimes truncate or sub-
cordate at the base, short-petioled : acorns sessile or nearly so ; cup
shallowly saucer-shaped, very thick and corky ; nuts oblong to
oblong-ovoid, 15-19 mm. long, often depressed at the apex : seed
very bitter.

On the summits of Caprina limestone hills, north-central Texas.

Related to Q. brcviloba but differing markedly in the glabrous
leaves, the thick corky coarsely warty cup of the acorn and the
oblong or ovoid-oblong nut.

The original specimens were collected on Lacey's Ranch near
Kerrville, Texas, by Mr. Howard Lacey during the years 1899-
1900, and given to me by Professor W. L. Bray, of the University
of Texas. Locally known as Bastard Oak. Type in the her-
barium of the New York Botanical Garden.

Quercus Brayi

A large tree sometimes 18 m. tall, or more, with a pale flaky
bark. Leaves very numerous, deciduous ; blades thin, cuneate,
10-20 cm. long, abruptly acuminate at the apex, regularly and
coarsely sinuate-toothed nearly to the base, glabrous, with rela-
tively few, prominent and regularly placed lateral ribs, deep green
above, slightly paler and rather olive-green beneath ; petioles 1.5-
2.5 cm. long; acorns sessile or nearly so; cup hemispheric,
2-2.5 cm. broad, the lower scales somewhat warty on the back;
nut oblong or ovoid, 2.5-3 cm. long, about 1.5 cm. thick: seed
rather sweet.

In canons, central Texas.

Quercus Brayi h most closely related to Q. Michauxii, but differs
conspicuously in the membranous and glabrous leaf-blades with
their few coarse teeth and lateral ribs, and the shorter and warty-
thickened scales of the cup of the acorn. The original specimens
were collected on Lacey's Ranch near Kerrville, Texas, by Mr.
Howard Lacey in the years 1 899-1900, and sent to me by Pro-
fessor W. L. Bray. It is locally known as White Oak. Type
in the herbarium of the New York Botanical Garden.

359 Small: Shrubs and Trees of the Southern States

AEsculus austrina

A shrub several meters tall, with pubescent foliage and in-
florescence. Leaf-blades 1-3 dm. broad, long-petioled ; leaflets
3-5, firm or leathery, oval or oval-ovate, short-acuminate at both
ends or acute at the oblique base (except in the case of the
terminal one), lustrous and glabrate above, except on the tomentose
nerves, densely tomentose beneath, sharply serrate, commonly
quite long-petioluled : panicles 1-1.5 dm. long, tomentose:
flowers deep red, about 3 cm. long : calyx tubular, red, 10-14
mm. long, glabrous ; lobes ovate, rounded : petals minutely gland-
ular, those of the upper pair with oval blades about as long as the
slightly villous claws, those of the lateral pair with obovate blades
which are exceeded several times in length by thair claws : stamens
exerted ; filaments sparingly villous.

In low grounds, Louisiana.

Most closely related to AEsculus Pavia, from which it differs in
the foliage and flowers. It is AEsailus Pavia /3 discolor of Torrey
and Gray,* in part, but not the AEsculus discolor of Pursh.f Tor-
rey and Gray's disposition of this plant is the most nearly correct
interpretation we have yet had. Dr. Gray later referred it to his
AEscuhcs flava var. purpurasccus, a species with which it has but
distant relationship.

The specimen on which the species is founded was collected in
Louisiana by Dr. Hale and is preserved in the herbarium of
Columbia University.

Hypericum interior

A much branched shrub, several dm. tall, with sharply angled
branches and glabrous foliage. Leaves rather numerous ; blades
narrowly oblanceolate to narrowly linear-oblanceolate, 1-3 cm.
long, acute, slightly revolute, pale beneath, narrowed into slender
petioles : panicles many-flowered : sepals linear or narrowly ob-
long, 3-4 mm. long, acute, rather rigid, more or less spreading at
maturity : corollas yellow, about i cm. broad : petals obovate or
cuneate-obovate : capsules conic-ovoid, 5 mm. long.
Along streams, Tennessee and Texas. Summer,
A shrubby species related to Hypericum galioidcs, but more
corymbosely branched, and with broader leaf-blades. The sepals

*F1. N. A. I : 252.
t Fl. Am. Sept. 254.

Small. Shruds and Trees of the Southern States 360

are characteristic, being linear or nearly so, instead of manifestly
narrowed to the base. The following specimens belong here :

Texas : [no locality] , Vcatch.

Tennessee: near Dandridge, July, \Za,2, Rugd : — type, in the
herbarium of Columbia University.

Azalea Candida

A rigid shrub 1-2 m. tall, with wide-branching stems and
white-tomentose young foliage, or the twigs sometimes brownish.
Leaves numerous ; blades leathery, obovate, oblanceolate or ob-
long, 1-5 cm. long, acute or apiculate, ciliate, somewhat revolute,
thinly tomentose above, densely white-tomentose and somewhat
reticulated beneath, short-petioled : corymbs several-flowered :
pedicels canescent and copiously glandular-pubescent : calyx-
lobes pectinate-ciliate : corolla rose-colored or pinkish, 3-3.5 cm.
long: capsules 1.5-2 cm. long, canescent, curved, longer than
their pedicels, often twice as long.

In hammocks and river swamps, southern Georgia. Spring.

Related to Azalea nndiflora, but easily distinguished by the
copious soft pubescence of the twigs, the white-tomentose leaf-
blades, the smaller flowers and the smaller softly and closely
pubescent capsules. Collected by the writer along the Withlo-
cooche River about Valdosta, Georgia, June 6-12, 1895. Type
in the herbarium of Columbia University.

Dendrium Hugeri

An evergreen shrub 2-4 dm. tall, with erect much branched
stems. Leaves mainly alternate ; blades leathery, oblong, 1-1.5
cm. long, lustrous and dark green above, paler beneath, obtuse,
revolute, somewhat obliquely narrowed into petioles 1-2 mm. long:
flower-clusters dense : bracts oblong-ovate, 3 mm. long, obtuse :
pedicels 5-10 mm. long, minutely glandular: calyx nearly gla-
brous ; lobes lanceolate, about i mm. long, acute : corolla white ;
lobes ovate, 4 mm. long, obtuse : filaments club-shaped, as long as
the corolla-lobes: capsules ovoid, 4-4.5 mm. long, glabrous or
nearly so, obtusely lobed, twice as long as the calyx-lobes, ab-
ruptly contracted into the stoutish style which is about as long as
the capsule-body.

On cliffs and rocky mountain summits, North and South Car-
olina. Spring and early summer.

This overlooked species is most closely related to Dcndrunn

361 Small: Shrubs and Trees of the Southern States

buxifolvun o{ \hQ Atlantic pine lands ; it is, however, larger through-
out and more erect. It may readily be separated by the large
leaves, the filaments, which are about as long as the corolla-lobes,
and the large capsule, which is about twice as long as the calyx-
lobes.

North Carolina : Top of Hibriten Mountain, Lenoir, Sept.,
1896, A. M. Hiigcr. Table Rock, 5. B. Buckley; Small &

Heller, 1 89 1.

South Carolina: Top of Table Mountain, August, 1896, /.
K. Small: — type, specimen in the herbarium of Columbia Uni-
versity.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 19.

THE AMERICAN SPECIES OF LIMNORCHIS
AND PIPERIA. NORTH OF MEXICO

BY PER AXEL RYDBERG.

NEW YORK
1901

[Reprinted from Bulletin Torrey Botanical Club, 28: 605-61S. 25 Nov., 1901].

The American Species of Limnorchis and Piperia, north of Mexico

By Per Axel Rydberg

In revising the manuscript of the Orchidaceae for Dr. Brit-
ton's Manual of the Flora of the Northern States and at the same
time that of my Catalogue of the Flora of Montana and the Yel-
lowstone Park,* I came to the conclusion that the genus Habe-
naria as treated in America was an altogether too diversified
genus and contained species of very distant relationship. In
Europe, as a rule, several genera are admitted. Prof. PI Pfitzer,
who prepared the Orchidaceae for Engler and Prantl's Die natiir-
lichen Pflanzenfamilien, places the North American Hahcnariae in
the following genera : Cocloglossiim, Gymnadcnia, Platanthera,
Perularia and Habaiaria. Kraenzlin in his Orchidaccaniui Gen-
era et Species, admits Habenaria, Gyninadenia and Plataiithera.
Both these authors place Habenaria in a separate subtribe from
the rest. The true Habenariae are represented in North America
by only a few species in the Gulf States, West Indies, Mexico and
Central America. They are characterized by the long filiform
appendages of the stigmas, anthers and petals. Platanthera as
understood by Pfitzer is still a very complex genus and had he
known the American forms as well as those represented in Europe,
he undoubtedly would have restored Rafinesque's Blepliariglotds for
our fringed orchids and established a few more genera. During
last July I stayed a few days in London and met Mr. Rolfe, the
authority on orchids at Kew. He expressed the opinion that
Habenaria must be split up, not only so that Gvuinadenia and
Platanthera should be taken out, but that all three genera should
be subdivided. In one instance he went farther than I had dared,
viz., expressing the opinion that Orchis I'oiundifolia perhaps should
be removed from Orchis.

When doing my work referred to above, I came to the conclu-
sion, that if Coeloglossnni and Pendaria should be admitted, certain
other groups represented wholly by American species must be

*Mem. N. Y. Bot. Garden, Vol. 1.

605

606 RvDBERG : The American Species of

taken out of Plataiithcra and Gyninadcnia. I therefore described
in my Catalogue of the Flora of Montana the new genera Lysiclla
^.nd Litniwrchis diwd in Dr. Britton's Manual Gymnadcniopsis* In
the latter I also reestablished BlepJiariglottis Raf In the Bulletin
of the Torrey Botanical Club,t I added another genus, Pipcria.
Of 'these, no species belonging to Gyninadoiiopsis, Bephariglottis
or Pipcria are mentioned by Pfitzer, which shows that he was not
well acquainted with these- plants.

This revision is based on the material found at the New York
Botanical Garden together with a few specimens cited from the
Canby Herbarium. A few of the Habcnariae, described from
Mexico and Central America may belong to Liiniiorchis or Pipcria.
I have not seen, however, any species from there referable to either
genus, but as several are unknown to me I have limited my work
to the North American species growing north of Mexico. One of
the Siberian specimens in the Columbia University herbarium is
evidently a Liiniiorchis, but as it is wrongly named and rather
scrappy, I have left it without consideration.

The illustrations are drawn by the writer and represent two
views of each species on a scale twice the natural size.

LIMNORCHIS Rydb. Mem. N. Y. Bot. Card, i: 104. 1900

Leafy-stemmed plants with elongated fusiform root-like tubers
and fleshy-fibrous roots : flowers whitish or greenish or tinged with
purple ; upper sepal ovate to almost orbicular, erect, 3— 7-nerved
but usually 5 -nerved ; lateral sepals from linear to ovate-lanceolate,
ireQ from the lip, 3-nerved, seldom 4— 5-nerved, spreading or often
somewhat reflexed ; upper petals erect, usually slightly shorter
than the upper sepal, from narrowly to broadly lanceolate, 3-nerved,
oblique at the base and semi-cordate, that is cordate on the lower
side ; lip entire, usually indistinctly nerved, flat or slightly con-
cave, reflexed, free, not clawed, from linear to rhombic-lanceolate,
obtuse ; column short and thick ; anther-cells parallel, opening
in front ; stigma broadly triangular ; ovary sessile, in fruit elon-
gated elliiDSoid.

The mode of propagation in many orchids is very peculiar.
Near the base of the stem is produced a short offset, tuber-, cbrm-
or root-like in appearance, usually fleshy. In the upper portion

* Wrongly printed as Gymnandeniopsis.
•j- 28 : 269. 1901.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO 007

is a hidden bud, from which the stem of the following year will
spring, the lower portion is often root-like, elongated, sometimes
cleft and functions more or less as a root. What this offset should
be called I do not know. It has been called both tuber and corm.
The latter name is not applicable from the way in which it arises.
This would indicate the name tuber, for it is really formed at the
end of a short subterranean branch, but it has only one bud and
this is directed towards the place from which the offset sprang.
The distal end, which in common tubers usually bears most of the
buds, is here often root-like. This is especially the case in Liui-
norchis, and here the offsets are usually taken for roots. It may
be that the larger portion thereof really is a fleshy primary root
(if the term primary root can be applied to an offset). The secon-
dary roots are developed the next year at its top from the base of
the bud.

The first species of Limnorchis was described as Orchis liyper-
borea Linn.* The description was drawn from a specimen from
Iceland collected by Konig. About the same time a plant also
collected by Konig (and probably the same as the type of 0. Jiy-
perborea) was figured in Flora Danica without a binominal name.
The description accompanying this is a little faulty because it
does not agree with the plate nor with any plant known. This
description was copied by Retzius f and he gave it the name
Orchis Koenigii. Orchis liyperborca L. is fairly common on Iceland
and Greenland but rare on the American continent. Another
plant closely related to it but larger and with shorter and more
clavate spur has been confounded with it. In preparing the re-
vision of Dr. Britton's manual I had not distinguished the two and
the description of the spur refers principally to the larger plant.
At my visit to the Botanical Garden at Copenhagen last sum-
mer, I first saw good material from Greenland and Iceland, and
then I noticed the difference between the two plants. The larger
plant has been named Platanthcra hyperborca [i major Lange \ and
Platantlicra Koenigii Lindl ; § but it is evidently not Orchis
Koenigii Retz.

* Linn. Mant. I2I. 1767.
f Fl. Scand. I: 16S. 1779.
JConsp. Fl. Groenl. I18. 1880.
I Gen. et Sp. Orchid. PI. 286. 1835.

608 Rydberg : The American Species of

In Habenaria liypcrborca R. Br. or PlatantJiera Jiyperborea
Lindl. there have been included several other forms, as for instance
Orchis Hiironensis Nutt.* {PlatantJiera Hiiroiicnsis Lindl. /. c. 288),
O. dolicliorhiza Fisch.f and Habenaria borealis fi viridiflora Cham.|
All three,' I think, are good species and they were so regarded by
Lindley. The first is common in the northeastern America. In
Britton's manual it appeared under the name LininorcJds Huronensis
(Nutt.) Rydberg. § I also described another species of the same
group as L. media. That there were at least three distinct forms
of the L. hyperborca group in the northeastern United States, I have
not been the only one to notice, for Professor Peck || recognized
three growing in the same meadow. I did not see his report be-
fore the page proofs of the manual had already gone to the printer,
but had come to the same conclusion independently. As the L.
Jiyperborea of my treatment there contains in my opinion now two
species, as I have stated before, the number of the group in the
northeastern United States becomes four, and, together with three
species of the L. dilatata group, our eastern species become seven.

Habenaria borealis /3 viridiflora Cham, is common throughout
the Rocky Mountain region, the Columbia valley and the northern
part of tl\e Pacific coast. It is without doubt closer related to the
true L. Jiypoborea than any of the other species and differs mainly
in the light green flowers and the short clavate spur.

OrcJiis dol ichor Jiisa (^PlatantJiera Kocnigii /9 dolicJiorJiiza Lindl.
/. c. 287), I have not seen, but from the description it must be
distinct from either. This was from the island of Unalaska.

The second in order of the North American species oi Livinor-
cJiis described was published under the name OrcJiis dilatata Purshlf
{LimnorcJiis dilatata Rydb. **). It is a common plant of the north-
eastern part of this country. Most of the specimens so named from
the Rockies and the west coast belong \.o Habenaria borealisQ\\-AVCs..W

*Gen. N. Am. Pi. 2 : 189. 1818.

fLind. 1. c. 287, as a synonym.

X Linnaea 3 : 28. 1828.

\ Britt. Man. Fl. Northern States and Can. 294. I901.

II See Peck, Regent's Rep. 50 : 126. 1897.

^Fl. Am. Sept. 588. 1814.

** Britten, Man. Fl. N. St. 294. 1901.

ff Linnaea 3 : 28. 1828.

LlMNORCHlS AND PiPEKIA NORTH OF MEXICO 609

{Liiniiorchis dilatatifonnis Rydb.*). This differs from L. dilatata
mainly in the shorter, more clavate spur, the dull or greenish white
flowers and the usually smaller size. It approaches much the
form referred to above under L. hypcrborca viz. Habcnaria borcalis
{iviridijJora Cham., and in fact connects the L. Jiypcrborea and the
L. dilatata groups. The other specimens from the West labeled
Habenaria dilatata belong to L. Iciicostacliys and a few of the new
species described below. No specimen of L. dilatata has been
seen from any place west of Minnesota.

Lindley in his Genera and Species of Orchideous Plants de-
scribed the following species which fall w'ithin the limits of Liin-
norcJiis : Platanthcra co)ivallariacfolia, P. leticostachys, P. gracilis, P.
stricta and P. graminca. While staying at Kew in July this year,
I had the privilege of seeing Lindley's herbarium as well as Doug-
las! plant preserved at Kew, which has enabled me to verify my
determinations.

P. convallariacfolia was described from specimens collected in
Kamtchatka by Fischer. It is a species resembling a slender L.
dilatata, but the lip is linear, not at all dilated at the base, the
spur is filiform and about the length of the lip and the lower leaves
are oblaTiceolate and obtuse. In the Torrey herbarium there is a
specimen communicated by Lindley. This is evidently a cotype
of that in Lindley's own collection for the two are as like as they
can be. In Torrey's herbarium there is also another specimen,
collected on Unalaska, which I also refer here.

P. Icitcostachys was described from a specimen collected by
Douglas. This is in the herbarium of the Kew Gardens. In
this specimen the spur is slender, filiform, not at all thickened and
not very obtuse at the apex. There is a form also common in
the Columbia valley with the same habit but with the larger spur
much more clavate and very obtuse at the apex. If this should
be regarded as a form of L. Icucostachys or as a distinct species is
a question of which opinions may differ, but it is, however, im-
portant to know what the original P. leiicostachys was.

P. gracilis is a slender plant with the white flowers of L. dila-
tata, but the lip is linear and the spur decidedly clavate. In
reality it is nearest related to P. convallariacfolia, from which it dif-

*Mem. N. Y. Bot. Gar. i : 105. 1900.

610 Rydberg : The American Species of

fers mainly in the spur. It is not at all the plant that has gone
under the name Habenaria gracilis in the United States. This is
the following species, belonging to another group of the genus.
P. gracilis is confined to Alaska and subarctic America.

P. stricta is the tall species, with greenish or purplish flowers,
linear lip and short saccate spur, that Watson and others have
called Habenaria gracilis. This is common in the Columbia
region and the valleys of the Rockies as far south as Colorado.

P. graminea resembles P. gracilis in habit, but the leaves are.
narrower and the lip dilated at the base, the spur is longer and
more clavate. Its home is Alaska. It was first collected by
Menzies, and later by Miss Cooley and by Gorman. A single
specimen from the lower St. Lawrence region evidently is to be
referred here also.

Hooker's Flora Boreali-Americana contains the same species
as Lindley's Genera.

Watson, in the Botany of California, has in his second section
of Habenaria, which section would correspond to Limnorchis, the
following species : H. Icucostachys (Lindl.) S. Wats., H. sparsiflora
S. Wats., H. pcdicellata S. Wats., H. hyperborea R. Br., H. Copcri
S. Wats., and H. gracilis S. Wats.

In H. Icucostachys, he includes H. Tkiirberi K. Gray,* but the
flowers of this species are greener, the dilated portion of the lip
oval instead of rhombic and the spur longer. Watson states the
characters of Gray's H. TImrbcri were mostly drawn from the vari-
ety mentioned (/. e., the same as H. sparsiflora), a statement which
is not correct.

H. sparsiflora Wats, is a good species, which, together with H.
brevifolia Greene t and two more species, constitute a distinct
group, perhaps a subgenus, distinguished b)' the very broad and
thin connective.

H. pcdicellata Wats. /. c. I have not seen, but from the descrip-
tion it cannot belong to Limnorclds.

H. hyperborea of the Botany of California is the same as H.
borcalis [i viridiflora Cham, and not O. hyperborea L.

* Proc. Am. Acad. 7 : 389. 1868.
t I'mc. .Am. Acail. 12: 276 1876.
J B .t (Jaz. 7 : 218. 1881.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO Oil

//. Coopcri Wats. /. c. is not a LiuinorcJds, but a Pipcria.

//. gracilis Wats, is not the same as PlatantJicra i^raci/is Liiid-
le}', but P. stricta of the same author.

Kraenzlin's treatment in his Orchidaccannn Genera et Species
is altogether unsatisfactory. He admits only two species, Platan-
tJicra gracilis and P. hypcrborea, the latter with several varieties.

His P. gracilis is, however, not P. gracilis Lindley but Habe-
naria gracilis Wats. {P. stricta Lindl.).

Under P. liypcrborca a var. gejinina he cites naturally also
Habenaria borcalis (i viridiflora Cham, as a synonym, but he has
printed viridis instead of viridiflora. I take this, however, as a
distinct species.

P. convallariacfolia he has reduced to a variety, P. hypcrborea
/5 var, convallariacfolia. In the same manner, he calls //. ddatata,
y var. dilatata Lindl. (I do not think that Lindley ever made it a
variety ; Kraenzlin is very loose in citing authorities.) Under this
variety he has as synonyms among others : Platanthcra Hiironcnsis
Lindl., Habenaria pedicellata S. Wats., H. Cooperi S. Wats., P.
<rravnnea and P. borcalis Reichenb. Of these the last named is the
only one, that can be referred here with any reason, for that, which
is the same as Habenaria borcalis Cham., belongs to the dilatata
group, while P. Hiironcnsis belongs to the hypcrborea group and H.
pedicellata and H. Cooperi belong to distinct genera.

Under his o var. leiicostachys he has as synonyms besides P:
Iciicostachys Lindl., also Habenaria brcvifolia Greene, P. GJiies-
brechtiana Rich. & Gal., P. sparsiJloraS. Wats., P. Thurbcriv. Grayi
S. Wats, and H.flagcllaris S. Wats. Of these Habenaria brcvifolia
Greene and H. sparsifiora S. Wats, can under no conditions be
referred to //. Iciicostachys on account of their broad connective.
The habit of H. birvifolia alone should throw it out ; there is
scarcely a more distinct species in the whole family. As far as
H. sparsifiora S. Wats, is concerned, it would have been more rea-
sonable to refer that species to his H. gracilis, i. e., H. stricta Lindl.;
the habit and lip in the two are the same and the flowers have some
resemblance. The main difference is in the spur and the connective.'
It is also to be noticed that Watson never published any Platan-
thcra sparsifiora but a Habenaria sparsifiora, nor any var. Grayi of
either Platanthcra or Habenaria Thurberi. In the citation of the

612 Rydberg : The American Species of

place of publication of P. Ghicsbrechtiana Rich, et Gal., he has Ann.
Sc. Nat. (1845) I. 30, when it should have been III. 3: 30.
Haboiai'ia flagcUaris S. Wats, has never been published.

A work with such numerous and glaring mistakes reflects little
credit to the author, and how can it be trusted ? He has given the
following remarks under his Platantlicra Jiypei'borea : " Es is ein
ganzlisch fruchtloses Bemiihen, die unzahligen Formen dieser weit-
verbeiteten Art unter Diagnosen von besonderen Species zu fix-
ieren. Je tiefer man in die Formenreihe eindringt, desto mehr
verschwimmen die bei der Betrachtung einzelner fiir constant gehal-
tenen Merkmale. Durch die Liberalitat amerikanischer Institute
und Botaniker verfiigen die europaischen Sammlungen iiber ein
reiches Material, welches die kritische Beurteilung sehr wohl ermog-
licht. Alle bisher beschriebenen 'Arten' stellen ihre Berechti-
gung auf rein habituelle Merkmale, was bei Orchideen stricte zu
verwerfen ist."

It is evident that notwithstanding the statement given above the
author has not tried very hard to bring the different forms under dis-
tinct diagnoses and he has not penetrated very deep into the series
of forms when he did not find the broad thin connective of Habc-
naria sparsiflora and H. breviflora ; perhaps he had seen neither.
It is strange if the herbaria of continental Europe should be rich
in North American orchids when our own are far from rich and
the representation at Kew is very meager. And still if the author
had examined the American material when at Kew, he would have
been able to correct some of his mistakes. None of the species
so far as I know have been merely based on characters in habit,
for in all, the spur, the lip, and the stamen have been described
more or less completely, and these are the only organs that give
any reliable characters. The habit is mostly the same in all and
can be used as a diagnostic character only in a few cases.

In discussing H. brevifolia Greene, the author remarks : " Der
Samler und Autor der Pflanze vergleicht sie mit Platantliera oder
nach seiner Schreibart Habenaria sparsiflora Wats. Der Vergleich
is nicht glijcklisch, denn PL sparsflora ist eine noch dazu wenig
charakteristische Form von hyperborea.'" Prof. Greene's comparison
of H. brevifolia with H. sparsiflora was the only logical one, for the
latter was the only known species with the flower-structure of H.
brevifolia, although this has much larger flowers.

LiMNORCHIS AND PiPERIA NORTH OF MEXICO 613

Synopsis of Species

Lip more or less lanceolate, /. <•., somewhat dilated near the base.
Flowers greenish or purplish.

Spur not twice as long as the lip. (Hyperbore.^e. )

Spur decidedly clavate, thickened and obtuse at the apex, shorter than
the lip.
Petals purplish ; spur only one half to two thirds the length of the
lip, very saccate.
Lip linear or nearly so, 5-7 mm. long ; ovary slightly curved ;

spike usually elongated. I . L. stricla.

Lip lanceolate, fleshy, 4-5 mm. long ; ovary strongly curved ;
spike usually short. 2. L. ptirpiirascens.

Petals greenish ; spur almost equalling the lip.

Petals ovate, three fourths as long as the upper sepal ; spur very

saccate. 3. L. hrachypetala.

Petals lanceolate, only slightly shorter than the upper sepal ;
spur less saccate.
Lips less than 5 mm. long, lanceolate.

4. L. viridiflora.
Lip over 5 mm. long, ovate-lanceolate.

5. L. major.

Spur slender, scarcely thickened at all towards the apex, often acutish,
equalling or slightly exceeding the lip.
Plant tall and stout ; flowers comparatively large ; sepals 4-6 mm.

long. 6. L. media.

Plant slender ; flowers small ; sepals 2.5-4 mm. long.
Flowers almost erect in a rather lax spike.

7. L. Iluronensis.
Flowers lateral in a dense spike. 8. L. hypcrborea.

Spur about twice as long as the lip. (Behringiaxae. ) 9. Z. Behringiana.
Flowers white or nearly so.

Spur about equalling or only slightly exceeding the lip. (DiLATATAE.)
Spur usually shorter than the lip and clavate. lO. L. borealis.
Spur equalling or slightly exceeding the lip, slightly or not at all clavate.
Plant over 3 dm. high.

Bracts all longer than the flowers, the lower three times as long ;
lips about I cm. long, the dilated part oval.

11. L. folios a.

Upper bracts shorter than the flowers, the lower at most twice as

long ; lips 5-7 mm. long, decidedly rhombic at the base.

Petals broadly lanceolate ; flowers inodorous or nearly so.

12. L. dilatata.
Petals narrowly linear-lanceolate ; flower very fragrant.

13. L. fragrans.
Plant less than 3 dm. high ; petals narrowly linear-lanceolate.

14. L. leptoceratitis.
Spur from Vj^ to % longer than the lip. (Leucostachyae. )

Spur filiform, mostly acutish.

Plant 3 dm. or less high ; spur 7-8 mm. long ; lip 5-6 mm.

14. L. leptoceratitis.

614 Rydberg : The American Species of

Plant S-8 dm. high; spur 10-14 mm. long ; lip 7-9 mm.

Dilated portion '• f the lip oval ; spur almost % longer than the

lip. 15. L. Thurberi.

Dilated portion of the lip decidedly rhombic ; spur scarcely y,
longer than the lip. 16. L. leucostachys.

Spur clavate at the apex.

Leaves lanceolate or oblanceolate ; stem tall and stout; lip decid-
edly rhombic at the base. 16. L. leucostachys robmta.
Leaves narrowly linear ; stem slender ; lip not rhombic at the base.

17. L. ^rominifolia.
Lip linear, /. e., not at all dilated at the base.
Spur usually shorter than the lip.

Flowers purplish or greenish ; spur very short and decidedly saccate.

I. Z. strict a.
Flowers white ; spur almost equalling the lip. (Convallariaefoliae. )
Spur clavate. 18. L. gracilis.

Spur filiform. 19. L. convallaria;folia.

Spur much exceeding the lip ; flowers greenish or purplish.

Connective narrow; spur almost twice as long as the lip. (Arizon-

ICAE. ) 20. L. Arizonica.

Connective broad.

Leaves elongated, lanceolate or linear; spur 8-IO mm. long.
(Sparsiflorae. )
Spike comparatively dense. 21. L. ensifolia.

Spike very lax.

Lip about 6 mm. long; spur about 8 mm., somewhat

clavate. 22. L. laxijiora.

Lip about 8 mm. long; spur over 10 mm., filiform.

23. L. sparsiflora.
Leaves very short, ovate ; spur almost 1.5 cm. long. (Brevi-
FOLIAE. ) 24. L. brevifolia.

§HvPERBOREAE : Ustially small or middle sized species, spike
usually dense, except in the first species ; flowers small, greenish
or tinged with purple, especially the petals ; spur less than twice
as long as the lip ; lip except in L. stricta lanceolate, /. c, some-
what dilated at the base but not at all rhombic ; connective nar-
row and anther-sacs close together.

I. LiMNORCHis STRICTA (Lindl.) Rydb. Mem. N. Y. Bot.

Garden i : 105. 1900
Platantliera stricta Lindl. Gen. & Sp. Orchid. PI. 288. 1835-9 ;
Habenaria gracilis Wats. Proc. Am. Acad. 11 : 277. 1876 ; Habc-
naria hyperborca Rothrock, Wheeler's Rep. 4 : 265. 1878 ; Habe-
naria saccata Greene, Erythea, 3 : 49. 1 895 ; Habenaria stricta
Ryd. Bull. Torr. Club, 24: 189. 1897; PlatantJicra gracilis
Kraenzlin, Orch. Gen. et Spec, i : 639. 1899. Not Lindl.

LiMXORCHIS AND PlPERlA NORTH OF MEXICO

615

Tall and strict 3-10 dm. high: tubers fusiform, .5-1 cm. in
diameter: lower leaves oblanceolate, obtuse, 5-12 cm. long, 15-25
mm. wide : the upper lanceolate, acute :
spike usually very long, 1-3 dm., and
lax : bracts linear-lanceolate, the lower
much longer than the flowers, often 3-4
cm. long: flowers 12-14 mm. lons" •

sepals green, rather thin ; the upper

ovate, erect, 4-5 mm. long ; the lateral

ones lanceolate, obtuse, 5-6 mm. long :

petals purplish, lanceolate, acute ; lip Fig. i.

linear, obtuse, 5-7 mm. long, thick,

purple ; spur one-half to two-thirds as long as the lip, purplish,

very saccate, /. c, thickened and round at the apex. (Fig. i.)

The structure of the flowers places this species in the L. hypcr-
Iwrca gxow^, although the lip is linear, scarcely at all broadened at
the base. This character and the lax spike would place it near
L. laxiflora, but that is distinguished by the long spur and broad
connective. L. stricta is common in damp places, swamps,
etc., from Alaska to Washington, Wyoming and Montana. It
has also been collected in Colorado at Twin Lakes, 1873, /<r^////
Wolf, p6j, and Manitou, 1896, Fra/. Clements.

Type : " America boreali-occidcntalis , Douglas^

2. LiMNORCHis PURPURASCENS Rydb. Bull. Torr. Club, 28 : 269.

1 90 1

A rather stout plant, 3-5 dm. high : tubers elongated, fusiform :
leaves ovate to lanceolate, acute, 6-ib cm. long, 15-30 cm. wide,

dark green : spike dense ; bracts lanceolate,
the lower exceeding the flowers : these 10-12
mm. long : lateral sepals oblong, linear to
lanceolate, obtuse, 4-5 mm. long ; the upper
broadly ovate, erect, obtuse, tinged with
purple : petals slightly shorter, erect, pur-
ple, lanceolate ; lip broadly linear-lanceo-
late, obtuse, about 5 mm. long, purplish,
thick, slightly dilated at the base ; spur
one-half to two-thirds as long as the lip. slightly curved and
strongly saccate : ovary strongly curved. (Fig. 2.)

This species resembles the preceding in the short spur and
purplish flowers, but the habit and form of the lip are more like
that of L. liyperborea. L. purpurascens is a rare plant, growing on

Fic.

616

Rydberc; : The American Species of

damp wooded hillsides at an altitude of 2700-3000 m. The fol-
lowing specimens belong here :

Colorado: Iron Mountain, i goo, Rydbcrg & ] rccland, 6414
(type in herb. N. Y. Bot. Gard.) ; Georgetown, 1878, M. E. Jones,
J 14; Como, South Park, 1895, C. S. Crandall ; Mouth of Chey-
enne Caiion, 1896, E. A. Bcsscy.

3. LiMNORCHis BRACHYPETALA Rvdb. Bull. N. Y. Bot. Garden

2: 161. 1901.

Stem slender, striate, 4-5-leaved, 1-2 dm. long : tubers elon-
gated fusiform, 7-8 mm. thick : lower leaves oblong, obtuse, 4-6

cm. long, strongly nerv^ed ; the upper lan-
ceolate, acute : spike short, 5 cm. long ;
bracts linear-lanceolate, the lower 2-3 times
as long as the flowers : flowers greenish or
brownish, 8 mm. long : upper sepals about
2 mm. long, nearly orbicular, slightl}' trun-
cate and indistinctly 3 -toothed at the apex ;
lateral sepals spreading, oblong, obtuse, nearly 3 mm. long :
petals round ovate, acute, slightly over i mm. long ; lip very
narrow, a little dilated at the base and near the apex, acute : spur
clavate, almost saccate, nearly straight, about equalling the lip in
length. (Fig. 3.)

In habit and flower most like L. Jiyperborca, but with a nar-
rower lip, shorter petals and shorter and thicker spur, which re-
sembles those of the two preceding but is comparatively longer.
L. brachypctala grows in wet places in Alaska and the Yukon
Territory.

Yukon Territory : Bennett City, 1 899, R. S. Williams (type
in herb. N. Y. Bot. Gard.); above Fort Selkirk, 1899, /.i>. Tarlc-
ton, 116.

Alaska: Unalaska, 1891, /. M. Macomi, 142.

4. Limnorchis viridiflora (Cham.)

Habenaria borealis /9 viridiflora Cham. Linnaea, 3 : 28. 1828 ;
Habe?taria hvpcrborea S. Wats. Bot. Cal. 2:134. 1 880 ; in part and
subsequent authors ; Limnorchis Jiypcrborca Rydb. Mem. N. Y.
Bot. Garden, i : 104. 1900.

LiMNORCHIS AM) PiPKRIA NORiri Ol- MkXICO

617

Fir,. 4.

Stem 2-5 dm. high : tubers elongate fusiform, 5-7 mm. thick :
lower leaves 5-15 cm. long, 2-3 cm. wide, oblanceolate, obtuse ;
the upper lanceolate, acute : spike short
and dense ; bracts linear-lanceolate, the
lower slightly exceeding the flowers :
flowers 10—12 mm. long, light green:
upper sepal broadly ovate, erect, 3—4
mm. long ; lateral ones 5-6 mm. long,
lanceolate, acutish or obtuse, spreading ;
petals erect, lanceolate, acute, slight!)'
shorter than the upper sepal ; lip lan-
ceolate, obtuse, less than 5 mm. long ; spur decidedly clavate,
curved, about equalling the lip. (Fig. 4.)

This species is closely related to the true L. Jiypci'lwrca, and is
its representative in the west. It differs only in the lighter green
flower and the^thicker and shorter spur. It is also related to L.
horcalis on the other hand, and it is no wonder that Chamisso
made it a variety ^of that species, as it differs only in the green
flowers and more saccate spur. Intermediate forms are met with ;
these [are 'perhaps of hybrid origin. L. viridiflora is common in
the Rocky Mountain regions and the Cascades. Its range ex-
tends from southern Alaska to North Dakota, central Nebraska
and Colorado, and is the common plant of the West, known as
Habenaria Jiypcrborea. Numerous specimens have been examined.

Type : " Unalaschca," Chamisso.

5. Limnorchis major (Lange)
Platanthci-d~^Kocnign Lindley, Gen. & Sp. Orchid. PI. 286.

1835. Not [6>rr///5 Kooiigii Retz. 1779; PlatantJiera dilatata

Torr. Fl. N. Y. 2 : 276, in part. 1843 ;
Habcnaiia Jiypcrborea Gray, Man. Ed.
5: 500, in part. 1867; Platanthcra
hypcrborea (i major Lange, Consp. Fl.
Groenl. 118. 1880; Limnorchis Jiypcr-
borea Rydb. ; Britton, Man. Fl. N. St.
294. 1 90 1, in part.

Stem stout, 2—6 dm. high : leaves
lanceolate, mostly acute, 5-30 cm. long,

1.5-4 cm. wide: spike dense, 5-15 cm. long; bracts lanceolate,

the lower slightly exceeding the flowers: flowers 12—14 mm.

long : upper sepal broadly ovate, obtuse, 5 mm. long ; the

Fig. 5.

618

RvDHERG : The American Species of

lateral ones ovate-lanceolate, obtuse, spreading : petals lance-
olate, acute, oblique, a little shorter than the sepals ; lip 5-6 mm.
long, ovate-lanceolate, obtuse ; spur almost equalling or some-
what exceeding the lip, distinctly clavate and curved. (Fig. 5.)

This is the most common species in Greenland, but also found
on the North American continent. It differs from L. hypcrborca
in the stouter habit, the larger flowers, the broader lip and the
clavate spur. It is more common than the specimens cited below
seem to indicate, but I have cited here only the specimens in our
herbaria, as these are the only ones presently at hand.

Greenland: Godhaven, 1878, L. Kinnlcin ; and a specimen
by an unknown collector from Hooker's herbarium.

Maine : Norway, S. O. Smith.

Wisconsin : Milwaukee, /. A. Lapliaui.

New York : North Yonkers, 1887,/. F. Poggenbiirg ; Torrey;
A. Gray.

6. LiMNORCHis MEDIA Rydb.; Britton, Man. N. St. 294. 1901

Habcnaria dilatata Torr. Comp. 3 1 8, in part. 1826. Not Orchis
dilatata Pursh. 181 3 ; Habenaria hypcrborca Gray, Man. Ed. 5 :
500, in part. 1867, and subsequent authors.

Stem very stout, 4-8 dm. high :
tubers narrowly fusiform, 5-7 mm. thick :
leaves lanceolate, acute, 1-2 dm. long :
spike long, densely flowered ; bracts
large, longer than the flowers, with sca-
brous margins : flowers divaricate, about
15 mm. long : upper sepal ovate, obtuse,
about 5 mm. long ; lateral ones lance-
olate or oblong, 5-6 mm. long : petals
green or purplish, lanceolate, acute ; lips
lanceolate, obtuse, about 6 mm. long ; spur filiform, /. c, not at
all clavate, curved, slightly longer than the lip. (Fig. 6.)

This is nearest related to the preceding, but still stouter, and
differs in the narrower lip and more slender, not clavate spur.
In habit it is intermediate between L. major and L. dilatata, and
has been confused with both. The name refers to this fact and
not to the size, for it is the largest species of the hypcrborca
group.

Canada : Quebec, Mrs. Pcrcival. (Type in Torrey Herba-
um.); Notre Dame chez Lac, iZ'^'j , JoJin Northrop.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO

619

Vermont: Willoughby, 1892, H. H. Rusby.
New York: John Torrey ; Silver Bay, Lake George, 1901./.
F. Kemp.

Wisconsin: Milwaukee, 1881, Dr. Hassc.

Minnesota: Minnehaha Creek, 1890, E. A. Mcarns, 77.

South Dakota : Harney Peak, 1 892, Rydbcrg, 102S (in part).

7. LiMNORCHIS HuRONENSis (Nutt.) Rydb.; Britton, Man. LI. N.

St. 294. 1 90 1

Oir/iis Hiironensis Nutt. Gen. N. Am. PI. 2 : 189. 1818 ; Plat-
anthcra dilatata Beck. Bot. 347, in part. 1833. Not Orchis dilatatn
Pursh. 1 8 14; Platanthcra Huroncnsis Lindl. Gen. et Sp. Orchid.
PI. 288. 1835; Habcnaria hypcrborca Gtz.y,MdiX\. Ld. 5: 500, in
part ; PlatantJiera hypcrborca y var. dilatata Kraenzlin, Orch. Gen.
&: Sp. 640. 1 899, in part.

Stem slender. 2-4 dm. high : lower
leaves oblanceolate, obtuse ; the upper lan-
ceolate, acute, 5-10 cm. long, 1.5-2 cm.
wide : spike long, lax ; bracts lanceolate,
the lower often twice as long as the
flowers : flowers light green, almost erect :
upper sepal ovate, 2-3 mm. long; lateral
ones 2-4 mm. long, oblong to lanceolate,
obtuse : petals narrowly lanceolate, slightly
shorter than the sepals, oblique ; lip linear-lanceolate, 3-4 mm.
long ; spur filiform, not clavate, strongly curved and slightly longer
than the lip. (Pig. 7.)

This species is nearest related to L. hypcrborea, and distin-
guished by the erect flowers, the narrower lip and the longer pod.
It is also usually taller with a laxer spike.

Type : " In wet places on the islands of Lake Huron and
Michigan."

Canada: Quebec, Mrs. Pcrcival.

Vermont: Monkton, 1879, C. G. Pringlc.

Massachusetts: Shelbourne, 1889, Geo. F. Taylor.

Connecticut: Canaan, 1866, W. H. Lcggctt.

New York: Utica, 1833,/^/'. Gray; Albany, L. C. Beck;
Jamesville, 1 890, L. M. Underwood ; Silver Bay, Lake George,
1 90 1,/. F. Kemp; Catskills, 1887, G. Rauipsbcrger.

Fig. 7.

620 Rydberg : The American Species of

Michigan: Long Lake, 1890, Bcardsley & Kofoid.
Minnesota: Chisago Lake, 1892, B. C. Taylor.
Wisconsin : Madison, S. H. Watso)i.

8. Limnorchis hyperborea (L.) Rydb. ; Britton, xMan. II. X. St.

294, in part. 1901

Orchis hyperborea L. Mant. 121. 1 767 ; Habenaria hyperborea
R. Br. Hort. Kew. Ed. 2:5: 193. 1813 ; Plataiitlicra Jiypcrborca
Lindl. Gen. & Sp. Orchid. PI. 287. 1835.

Low and slender, 1.5—4 <^^^- l^ig^^ • tubers
almost fleshy-fibrous, slightly thicker than the
roots : lower leaves oblanceolate, obtuse, 5- 10
cm. long, 1-2 cm. broad ; the upper lance-
olate, acute : spike rather dense ; bracts linear-
lanceolate, the lower slightly longer than
the flowers : flowers light green, 8-12 mm.
long : upper sepal broadly ovate, obtuse,
3-4 mm. long ; lateral ones lanceolate: petals
lanceolate, slightly shorter than the sepals ; lip lanceolate, obtuse,
nearly 5 mm. long ; spur filiform, strongly curv^ed and a little ex-
ceeding the lip. (Fig. 8.)

In habit L. viridiflora and this species are much alike and the
former would have been merged in the latter, if all the western speci-
mens (/. e., L. viridiflora) seen, had not had more or less clavate spur,
a character not seen in Icelandic, Greenlandic and eastern specimens.
Type: " Islandia, K'dnigy

Iceland: Gardas, 1888, Prof. & Airs. Sprague Smith.
Canada : Big Swamp, 1 866, /. Macoun.
New Brunswick : Kennebeccosis, 1871,/. Fow/er.
Vermont: Willoughby, 1892, //. H. Riisby.
New York : Kirkville, Onondaga county, 1891, L. M. Cnder-
ivood, 3155.

§ Behringianae: Low plants, i- 1.5 dm. high : spike short and
dense: flowers greenish-purple ; lip oblong-lanceolate ; spur fulh'
twice as long as the lip ; connective narrow.

9. Limnorchis Behringiana sp. nov.

Stem low, 1-1.5 dm. high, about 3-leaved : tubers elongated
fusiform, about 5 mm. thick: lower leaf ovate-lanceolate, about 5
cm. long and 1.5—2 cm. wide ; the upper lanceolate and smaller:
spike dense, 3-4 cm. long ; bracts linear-lanceolate, the lower about

LiMNORCHlS AND PiPERIA NORTH OF MEXICO

621

1^^

twice as long as the flowers : flowers purplish, about I2 mm. \om
upper sepal ovate, obtuse, 4-5 mm. long ; the lateral ones oblong
petals equalling the .sepals, broadly-
lanceolate ; lip about 5 mm. long ;
spur fully ID mm. long, filiform.
(Fig. 9)

This species is not closely re-
lated to any of the other species.
The type specimens were labelled
Habeiiaria gracilis Wats. With
this, /. e., L. stricta it has scarcely
anything common except the color of the flowers and such char-
acters as are found in all species of LimnorcJiis. L. Bchringiana
has not been found in North America but I mclude it in here, as it
would] likely be found in Alaska, as L. convallanaefolia from the
same region has been.

Asia: Behring Island, 1891, British Behring Sea Commission,
i^j (type in herb. Columbia Univ.).

§Dilatatae: Plants usually middle sized : flowers white or in
the first species yellowish-white ; lip lanceolate, more or less dis-
tinctly rhombic dilated near the base ; spur equalling or slightly
longer than the tip ; connective narrow.

10. Limnorchis borealis (Cham.)
Habcnaria borealis Cham. Linnaea, 3 : 28. 1828

Plataiithera

dilatata Lindl. Gen. and Sp. Orchid. PI. 287; in part. 1835;
Habcnaria
Habcnaria

dilatata Coult. Man. Rocky Mts. 342. 1885;
dilatatiforuiis Rydb. Bull. Torr. Club, 24: 189.
1897 ; Platanthera liypcrborca y var. dilatata
Kraenzlin, Orch. Gen. et Sp. 640. 1899, in
part ; Limnorchis dilatatiformis Rydb. Mem.
N. Y. Bot. Garden, i: 105. 1900.

Tall and leafy, 4-8 dm. high : tubers
elongated fusiform, 5-8 mm. thick : lowest
leaves oblanceolate and obtuse ; the rest
lanceolate, acute, 5-15 cm. long, 1.5-2.5
spike often rather dense, 1-2 dm. long; bracts lan-
ceolate, the lower often much exceeding the flowers : flowers
10-14 mm. long, white or sometimes yellowish or greenish white:
upper sepal ovate, obtuse, 4-5 mm. long ; the lateral ones oblong-

FiG. 10.

cm.

wide

622

RvDBERG : The American Species of

lanceolate, spreading : petals lanceolate, slightly shorter than the
sepals; lips rhombic-lanceolate, obtuse, about 5 rnrn. long; spur
usually shorter than the lip, and more or less clavate. (Fig. 10.)

This species represents L. dilatata in the Rocky Mountain re-
gion, but differs in the smaller flowers and shorter, more clavate
spur. It also approaches L. viridiflora, from which it differs in
the whiter flowers and more rhomboid lip. Where growing
together, intermediate forms are often found, perhaps of hybrid
origin. L. borealis is common from Alaska to Washington and
Colorado.

Type : "Unalaschca," Chamisso.

Fig. II.

1 1 . Limnorchis foliosa sp. nov.

Stem stout and very leafy, about 3 dm. high : leaves linear-
lanceolate, attenuate, i — 1.5 dm. long: spike short and dense, less

than I dm. long ; bracts linear-
lanceolate, attenuate, about three
times as long as the flowers : these
about 1 5 mm. long, white : upper
sepal broadly linear, obtuse, about
6 mm. long ; lateral ones narrowly
lanceolate, about 9 mm. : petals
narrowly linear-lanceolate, attenu-
ate ; lip about 10 mm. with an oval
base and an almost linear lower
half, obtuse : spur about the length
of the lip, filiform, not at all clavate.
(Fig. II.)

This species has the habit o{ Pcnilaria flava and Cocloglossuui
bractcatimi, but the flower is of the typical LiinnorcJns t}^pe and
places it nearest to L. dilatata. The flowers, however, are larger
and the lip less rhomboid.

Al.\ska : Nagai, 187 1-2, M. \V. Harrington (Ball's Fxplora-
tion ; type in the Columbia herb.).

12. LiMXOKCHis DiL.vr.ATA (Pursh) Rydb.; Britton, Man. Fl. N.

States, 294. 1 90 1

OrcJiis dilatata Pursh, Fl. Am. Sept. 588. 18 14; Habcnaria
dilatata Hook. Exot. Fl.//. gj. 1825 ; Plata ntlicra dilatata Lindl.;
Reck. Rot. N. & M. St. 347, in part. 1833 ; Platanthcra hyper-

LljVINORCHIS AND PiPERIA NORTH OF MeXICO

623

Iwrca y var. dilatata KraenzHn, Orch. Gen. ct Sp. 640, in part.

1899.

Stem slender, tall, leafy, 3-6 dm.
high : leaves lanceolate, 7-20 cm. long,
the lower obtuse, the upper acute :
spike 5-25 cm. long, usually rather
lax ; bracts linear-lanceolate, the lower
twice as long as the flowers : flowers
white, 15-18 mm. long: upper sepal
ovate, obtuse, about 6 mm. long, the
lateral ones lanceolate, acutish : petals
acute, lanceolate ; lip lanceolate with

a rhomboid base, about 7 mm. long ; spur longer than the lip,
filiform, scarcely at all clavatc. (Fig. 12.)

L. dilatata is common from Newfoundland to Saskatchewan,

south to New York and Nebraska.

Type : " In Labrador. Co/inastcr, v. s. in Herb. Dirkson.''

Limnorchis dilatata linearifolia var. nov.

Stem slender; leaves linear,' attenuate, 5-10 cm. long, 5-7 mm.
wide.

Very different from the species in general habit, but the flowers
are exactly the same, and intermediate forms are not lacking.

New York : Bridgewater, Dr. Gray (type in herb. Torrey).

Michigan : Portage of Kee-w\ana-wa, Dr. Peters.

13. Limnorchis fragrans Rydb. ; Britton, Man. Fl. N. States,

294. 1 901

Stem slender, 2-3 dm. high : tubers narrowly fusiform, about
5 mm. thick : leaves linear, about i dm. long and 8-10 mm. wide,

acute : spike slender, lax ; bracts lanceo-
late, acuminate ; the lower longer than the
flowers : flowers pure white, very fragrant :
upper sepal ovate-lanceolate, obtuse ; the
lateral ones linear-lanceolate, acutish,
strongly veined : petals narrowly linear-
lanceolate, equalling the sepals ; lip lanceo-
late with an ovate-rhomboid base, about 5
mm. long ; spur filiform, not at all clavate,
curved, slightly exceeding lip. (Fig. 13.)
Closely resembling the variety of preceding species, it is dis-
tinguished by the narrow petals and sepals and by the fragrance.

624

RvDBERG : The American Species of

The two sheets of the orginal collection in the Columbia Herbarium
are the only ones seen.

Vermont: Willoughby Mountain, 1892, H. H. Rnsby (type
in herb. Columbia Univ.).

14. LiMNORCHis LEPTOCERATiTis Rydb. Bull. N. Y. Bot. Garden,

2 : 162. 1901

Stem slender, 2-4 dm. high : tubers slender, slightly tliicker
than the fleshy fibrous roots : lower leaves oblong, obtuse, 4-8

cm.

long,

I -I.; cm. wide, the

upper

linear-lanceolate, acute : spike short, less
than I dm. long ; bracts linear-lanceo-
late, the lower somewnat longer than
the white flowers : sepals 3-4 mm. long,
lanceolate : petals linear or narrowly
linear-lanceolate, about equalling the
sepals ; lip lanceolate, somewhat rhombic,
dilated at the base ; spur very slender, filiform, strongly curved
forward, a little exceeding the lip. • (Fig. 14.)

This is nearest related to the eastern L. dilatata and L. fra-
irrans, but differs in the smaller size, smaller flowers and shorter
leaves. In habit it resembles most a depauperate L. borcaHs, but
the spur is different.

Yukon Territory : Bennett City, 1899, R. S. JVi//iams {type
in herb. N. Y. Bot. Card.).

Alaska: Unalaska, 1891./. J/. Macon n : southern Alaska,
1883,/. Albert Rudkin.

British Columbia: Tulameen River, 1900, /. F. Kcuip.
§ Leucostachvae : Tall and stout plants : flowers pure white or
slightly greenish in L. Tliurbcri ; lip lanceolate with a more or less
rhombic or oval dilatation near the base ; spur more than y.^ longer
than the tip ; connective narrow : spike long and dense.

15. Limnorchis Thurberi (A. Gray) Rydb.

Habcnaria Thurberi A. Gray, Proc. Am. Acad. 7: 389. 186S;

Habcnaria IcucostacJiys S. Wats. Bot. Cal. 2 : 134, in part. 1880 ;

Coville, Cont. U. S. Nat. Herb. 4: 201. 1893 ; Platanliicra Jiypcr-

borca d var. Iciicostacliys Kraenzlin, Oich. Gen. ct Sp. 640, in part.

1899.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO

625

Fir.. 15.

Stem stout and leafy, 4-6 dm. high : tubers elongated fusi-
form, 6-7 mm. in diameter : leaves lanceolate to linear-lanceo-
late, 1-2 dm. long, 1.5-3 cm. wide,
acute : spike long and dense ; bracts
linear-lanceolate, attenuate, about
equalling the flowers : these dull
white, 16-20 mm. long: upper
sepal ovate, obtusish ; the lateral
ones lanceolate, acute : petals lan-
ceolate, about equalling the sepals ;
lip 7-8 mm. long, lanceolate, ob-
tuse, the dilated portion ovate ; spur
filiform, curved, about -,< longer than
the lip.

This has been merged into A. Icucostachys, which it resembles
in habit, but the spike is denser, the flowers are dull or greenish
white, and the form of the lip is different.

Type : " Arizona, Thurber (925)."

New Mexico: 1851-52, C. Wright, igoo.

California: Thouias Bridges, ^56 ; Mammoth, 1891, Covillc &
Funston, 1822 ; Miner's Ditch, Nevada county, 1867, N. J.Davis,
J 7 ,• Canoe Creek, /. .S". Newberry; Nevada, 1893, Michcner &
Bioletti ; Sisson, 1897, H. E. Brown, j^ 20 ; Marin county, 1873,
Edtvards; Bernardino Mts., 1882, S. B. & W. F. Parish, ij2i ;
Fresno county, 1900, Hall & Chandler, ijO.

16. LiMNORCHIS LEUCOSTACHYS (Lindl.) Rydb. Mem. N. Y. Bot.

Garden, i : 106. 1900.

Platanthera leiicostachys Lindl. Gen. & Sp. Orchid. PI. 288;
Habenaria leiicostachys S. Wats. Bot. Cal. 2: 134. 1880; Pla-
tanthera hypcrborea o var. leiicostachys Kraenzlin, Orch. Gen. et
Sp. 640, in part. 1899.

Stem stout and tall, 6-10 dm. high: lower leaves oblanceo-
late, 1-2 dm. long, 1.5-3 cm. wide; the upper lanceolate, acute:
spike 1-3 dm. long, rarely very dense ; bracts linear-lanceolate, the
lower exceeding the flower : these purely white, i 5-20 mm. long :
upper sepal ovate, obtuse, about 5 mm. long ; lateral ones lanceo-
late, acute, j-^ mm. long : petals lanceolate, attenuate, a little
shorter than the upper sepal; lip lanceolate with a decidedly
rhombic base, about 8 mm. long ; spur filiform, scarcely clavate,

626

RvDBERG : The American Species of

acutish, about half longer than the lip, less than i mm. thick.
Fig. 1 6.)

Lindley describes this as having the
lip half as long as the spur, but I have
not seen any with so long a spur, not
even in Douglas' specimens at Kew.

TvPE : "In ora occidcntali Aiiicricae
scptcntrioiialis, Dcniglns {Jiab. s. sp. coinui.
Soc. Hort.)r

Alaska: Ankow River, 1892, /'".
Funstofi, j2.

Washington : Snoqualmie, 1892, J/.
Parker; Wallawalla to Kuskuski, 1838-42, Wilkes Exp., J26.
Idaho: 1892, Isabel Mulford ; Priest Lake, 1900, D. T. Mae-
Dougal, 7 and ijS.

Utah: i^yi, Palmer, 461.

Nevada: E. Humboldt Mountains, 1868, S. Watson, 1134-
California: Sierra county, 1S74, L:inin)i \ Plumas county,
1 88 1, Mrs. Austiti ; Yosemite Valley, 1866, Bolander, 4.^36.

Fig. 16.

Limnorchis leucostachys robusta var. nov.

Very stout and tall, 4-10 dm. high, leafy: spike long and
dense, 1-3 dm. long : spur usualK' clavate, over i mm. thick,
obtuse, about one-third longer than the
lip. (Fig. 17.)

This may be distinct from L. leuco-
stachys but the onl}' characters distin-
guishing it from the type is the thicker
spur and the denser spike. More field
work is needed in order to settle its
relationship to L. IcucostacJiys. The fol-
lowing specimens are in our herbaria :

Oregon : 1871, Elihu Hall, 505.

Washington: 1889, R. S. J'asey, yj (type in herb. N. Y.
Bot. Gard.) ; Chevalis River, 1897, Frank //. Laud\ iigga;
Olympia, 1898, A. A. & E. G. Heller, 4046 : Falcon Valley, 1893,
W. H. Suksdorf, ijj6 ; Upper Valley of the Nesqually, 1894,
O. D. Allen ; Yakima Region, 1883, F. Tweedy ; Palace Camp,

Fig. 17,

LlMNORCHIS AND PiPERIA NORTH OF MeXICO

627

I SS^, Mrs. Baihy Willis; Seattle, 1890, C. \\ /"/>•/' ; Cascade
Mts., 1882. Brandcgi'c, ^80.

British Columbia: Vancouver Island, 1887, /<V/// Macouii ;
Tulameen River, 1900, Prof. J. F. Kemp.

Idaho : Valley of Clearwater River, 1892, Sandberg, Mcxc-
Dougal & Heller, 288.

17. Limnorchis grarainifolia

PlatantJiera graininca Lindle)', Gen. & Sp. Orchid. PI. 289.
1835 ; not Hahoiaria graniinca Sprcng. Syst. 3: 690. 1826; nor
P. graininca Lindley, /. c. 292 ; Platanthcra Jiypcrborca y var.
dilatata Kraenzlin, Orch. Gen. et Sp. 640, in part, 1890.

Very slender, 3-4 dm. high : leaves
narrowly linear, 5-10 cm. long, 3-4 mm.
wide : spike short ; bracts lanceolate,
shorter than the flowers : these white,
12—15 mm. long: upper sepal ovate, 4
mm. long, the lateral ones lanceolate :
petals narrowly lanceolate, slightly
shorter than the upper sepal ; lip lanceo-
late, obtuse, dilated near the base, but
not rhombic ; spur one-third to one-
half longer than the lip, clavate towards the ape.x, acute. (P'ig. 18.)

This plant is nearest related to L. Icucostachys, but differs in the
slender habit, narrow leaves and the form of the lip. The type is
in Lindley's herbarium, where I saw it. The specimens from
Lower Canada I cannot distinguish from the Alaskan specimens,
although the occurrence of the species on the east coast is exceed-
ingly remarkable.

Type : "In ora occidcntali Amcricac scptcntrionalis, Mcnzics
{Jiah. s. sp. conini. ccl. Mensiesy

Alaska: Douglas Island, 1891, Grace E. Coolcy ; Gorman's
Lake, 1895, M. IV. Gorman, 7./.

Canada: Mouthof Riviere du Loup, Wm. Canby.

% Convallariaefoliae: Slender plants about 3 dm. high: spike
short and rather lax : flowers white ; lip linear, not at all dilated
at the base ; spur about equalling the lip ; connective narrow.

18. Limnorchis gracilis (Lindl.).
PlatantJiera gracilis Lindley, Gen. & Sp. Orchid. PI. 288.
1835-

Fic. 18.

628

Rydberg : The American Species ok

Fig. 19.

Slender, about 3 dm. high : leaves oblong to lanceolate, 5-10
cm. long, 1-1.5 cm. wide, the lower obtuse, the upper acute : spike

slender and lax, about i dm. long ;
bracts lanceolate the lower slightly
exceeding the flowers : these white,
10-12 mm. long : upper sepal broadly-
oval, about 4 mm. long, the lateral
ones linear-lanceolate, acute : petals
linear-lanceolate ; lip linear, obtuse,
6—7 mm long ; spur clavate, only
slightly curved, a little shorter than
the lip. (Fig. 19.)
This resembles L. stricta in the linear lip and the lax spike,
but has white flowers and slender spur. Watson confused the
two and following him, most authors have used the name Habc-
naria gracilis for L. stricta. No one who has seen Lindley's type
could confuse the two.

Type: "In ora occidciita/i Ainericac scptcntricmalis, Menzics ;
Observatory inlet. Herb. Hooker {hab. s. sp. comm. eel. Menzies)."
Alaska: Back Bay, 1895, M. IV. Gorman, f,2 (in Colum-
bia Herbarium) ; Sitka, i8cji, IV. G. Wright {1359) (?)•
Subarctic America: 186 1-2, /. S. Onion.

19. Limnorchis convallariaefolia (Lindl.)

Platanthera convallariaefolia Lindl. Gen. & Sp. Orchid. PL 287.
1835; Platanthera hyperborea (i,VA.\. cojiv all ariaef alia Kraenzlin,
Orch. Gen. et Sp. 640. 1899.

Stem slender, about 3 dm. high : leaves
oblong-lanceolate or oblanceolate, 4-8 cm.
long, 1-2 cm. wide, the lower obtuse, the
upper acute : spike short, less than i dm.
lone ; bracts lanceolate or linear-lanceolate,
equalling or the lower exceeding the whitish
flowers : upper sepal almost orbicular or
broadly ovate, 3-4 mm. long ; lateral ones

lanceolate, obtuse : petals lanceolate, acute ; lip linear, about 6
mm. long, obtuse ; spur filiform, curved, almost equalling the lip.
(Fig. 20.)

This is closely related to L. gracilis, and differs scarcely in any
respect except that the flowers are smaller and the spur is longer
and more slender. It was described from specimens collected by

Fig. 20.

LiMNOKCHIS AND Pll'KKIA NORTH OF MEXICO

629

Fischer in Kamtchatka. The t\'pe is in Lindley's herbarium and
a cotype in Torre>-'s. The Unalaska specimen also collected by
Fischer, I cannot distinguish from the Asiatic.

Type : "In KamtcJiatka, Fischer {hab. s. sp. couim. ccl. Prcs-

cott)r

Asia : Kamtchatka, Fischer, ex. herb. Liudley.

Alaska : Unalaschca, Fischer.

§ Arizoxicae : Rather stout, 5-7 dm. high : spike long and lax :
flowers light greenish ; lip linear, not at all dilated at the base ;
spur filiform, almost twice as long as the lip ; connective narrow.

20. Limnorchis Arizonica sp. no v.

Stem tall, leafy : lower leaves oblong, about 5 cm. long, obtuse,
the middle and upper ones linear, the former 2-3 dm. long, 2-3
cm. wide : spike slender, lax, many-flowered
1.5-3 cm. long; bracts linear-lanceolate, the
lower slightly longer than the flowers : these
12-15 mm. long, greenish or purplish : upper
sepal broadly ovate, 3-4 mm. long ; lateral
ones ovate-lanceolate : petals lanceolate, acute,
slightly shorter than the upper sepal, 4-5
mm. long. (Fig. 21.)

This species resembles somewhat both L.
stricta and L. sparsiflora. From the former
it is easily distinguished by the long slender spur and from the
latter by the many-flowered spikes, smaller flowers and narrow
connective. It grows in rich canons of Arizona.

Arizona: 1891, Nealley (type in herb. N. Y. Bot. Gard.);
Santa Rita Mts., 1881, C. G. Pringlc ; Ft. Huachuca, 1893, T. E.
Wilcox ; Weber Creek, 1887, E. A. Mearns, ijj.

§ Sparsiflorae : Plant slender, light green : spike elongated :
flowers greenish with strongly reflexed sepals ; lip linear ; spur
slender, much longer than the lip ; connective very broad : leaves
normal.

2 1 . Limnorchis ensifolia sp. nov.

Platantliera liyperborea o var. leucostacJiys Kraenzlin, Orch.
Gen. et Sp. 640, in part. 1899; not P. leiicostachys Lindl.

Stem strict, 3-4 dm. high, light colored, few leaved: leaves
linear-lanceolate, attenuate, slightly falcate, 1-1.5 dm. long, 1-1.5
cm. wide : spike short and not very lax ; bracts lanceolate.

Fig. 21.

G30

RvDUERG : The American Species of

acuminate, light green, about equalling the flowers : these greenish,
about 1 5 mm. long : upper sepal broadly ovate, about 5 mm.

long ; lateral sepals about 7 mm. long,
oblong-lanceolate : petals narrowly lan-
ceolate, acute ; lip linear, about 8 mm.
long, thick ; spur filiform, about i cm.
long. (Fig. 22.)

This is nearest related to L. sparsi-

flora, but the spike is shorter and denser,

the upper sepals and petals larger and the

bracts shorter and broader, and light

colored. The type was growing at an

altitude of about '2600 mm.

Arizona: Mt. Humphrey, 1897, R. E. Kutizc (type in herb.

N. Y. Bot. Gard.) ; Willow Springs, 1874, Rothrock, 36g.

Utah : " Central Utah," 1875, C. C. Parry, 8c} ; Rock Creek,
1877, Dr. E. Palmer, 460 (in part).

New Mexico: Silver City, 1880, E. L. Greened

Fig. 22.

'>9

Limnorchis laxiflora sp.

nov.

Stem slender, 4-6 dm. high : lower leaves oblanceolate, obtuse,
8-10 cm. long, 1-1.5 cm. wide ; the upper linear-lanceolate, acute:
spike very slender and lax, 1-2 dm. long, few-
ilowered ; bracts linear-lanceolate, usually
equalling or shorter than the flowers : these
greenish, 10-12 mm. long : upper sepals broadly
obovate, obtuse, about 4 mm. long ; lateral
ones broadly lanceolate, acutish : petals almost
equalling the upper sepal, lanceolate ; lip
linear, obtuse, about 6 mm. long ; spur about
8 mm. long, slightly clavate, a little longer
than the lip. (Fig. 23.)

This is closely related to and has been mis-
taken for L. sparsijiora, which, however, has larger flowers, longer
spur, not at all clavate, longer bracts and more slender stem.

Oregon: Coast Mountains, 1884, Thomas Howell (type in
herb. Columbia College); Interior of Oregon, 1838-42, Wilkes
Expedition ; 1871, Elihu Hall, 504.

Utah : Rock Creek, 1877, Dr. E. Palmer {460 in part).

Fig. 2-

* This is labelled Habenaria Jiagellaris Wats.

LiMNORCIIlS AND PiPERIA NORTH OF MEXICO

631

Colorado: Uncompahgne Mountains near Los Pinos, 1878,
IVm. F. Flint.

23-

Limnorchis sparsiflora (S. Wats.) Rydb.

Fir.. 24.

Habcnaria sparsiflora S. Wats. Proc. Am. Acad. 12 : 276.
1877; PlatantJi:ra Iiyperborea d var. IcucostacJiys Kraenzlin, Orch.
Gen. et Sp. 640, in part. 1899.

Stem tall and slender, 4—6 dm. high :
lower leaves oblanceolate, obtuse, 1—2 dm.
long, 1.5—3.5 cm. wide; the upper lanceo-
late, acute : spike long, slender and lax,
2—3 dm. long ; bracts linear-lanceolate,
usually exceeding the flowers : these light
green, very delicate, about 15 mm. long:
upper sepal broadly ovate, 4-5 mm. long;
the lateral ones lanceolate, acute, about 6
mm. long : petals narrowly lanceolate,
acute ; lip linear, about 8 mm. long, obtuse ; spur filiform, about
1 cm. long. (Fig. 24.)

This is characterized by the long, slender, lax spike, delicate
green flowers and long bracts. The type was collected by Bo-
lander, near Mariposa Grove, California.

Oregon: Kerbyville, 1884, T. Hoivcll.

California: Donner Lake, 1865, /. Torrey, jii ; Pine Ridge,
Fresno county, 1900, Hall & Chandler, ijg ; Truckee River,
1893, C. F. Sonne; Mt. Dyer, 1879, ^- ^^- Austin; Sierra county,
1874, Lenimon.

New Mexico: 1881, MongoUon Mountains, H. H. Rnsby.

§ Brevifoliae : Like the Sparsiflorae, but leaves very short,
much reduced : flowers nearly twice as large.

24. Limnorchis brevifolia (Greene) Rydb.

Habenaria brevifolia Greene ; Coulter, Bot. Gaz. 6:218. 1 88 1 ;

Platantlicra Iiyperborea o var. leucostachys Kraenzlin, Orch. Gen.

et Sp. 640, in part. 1899.

Stem strict, leafy, 3-6 dm. high : leaf-blades ovate, 2-4 cm.
long and about i cm. wide, acute : spike elongated, 1—2 dm.
long; bracts similar to the leaves but smaller, about equalling the
flowers: these green, 2-2.5 cm. long: upper sepal ovate, about
5 mm. long ; lateral sepals linear, acute, about 8 mm. long : petals

632

Rydberg : The American Species of

linear-lanceolate, acute, a little shorter than
the upper sepal ; lip linear, obtuse, about
lO mm. long; spur filiform, about 15 mm.
long. (Fig. 25.)

This is characterized by its large flowers
and reduced leaves.

Type : " On dry southward slopes of the
Finos Altos Mountains, New Mexico, in
i88o."

New Mexico : White Mountains, 1 897,
E. O. Wooton, 5./^ ; Finas Altos Mountains,
E. L. Greene.

Mexico: State of Chihuahua, Sierre Madre, 1887, C. G.
Pringle, IJ74.

Fig. 25.

FIPERIA Rydb. Bull. Torn Club, 28: 269. 1901

Somewhat leafy-stemmed plants, but the leaves are usually
near the base and withering at or before the anthesis, the stem-
leaves being reduced and bract-like : tubers spherical or rounded
ellipsoid; flowers greenish or white ; sepals and petals i-nerved
or very obscurely 3-nerved ; the upper sepal ovate or lanceo-
late, erect ; the lateral ones spreading, linear to lanceolate, their
bases united with the claw of the lip ; upper petals free, lanceolate
or linear-lanceolate, oblique, but not cordate ; the blade of the lip
linear-lanceolate to ovate, obtuse, truncate or hastate at the base,
concave but with a longitudinal low ridge in the middle produced
by the undulation of the lip ; the claw, if it can be called so, united
with the bases of the lower sepals, bordered with an erect margin
which connects the lip with the column ; anther cells unusually
large for the size of the flower, parallel, opening nearly laterally :
stigma a small beak in the angle between the anther-cells ; ovary
sessile, ellipsoid in fruit.

The principal differences between Limnorchis and Piperia are
the following :

Limnorchis

FlPERIA

Tubers rounded.

Tubers elongated fusiform,
root-like.

Stem leafy ; leaves remaining Stem leafy at the base ; leaves
until fruit is set. withering at or before anthesis.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO 633

LiMNORCHIS PiPERIA

Sepals 3-7-nerved ; petals dis- Petals and sepals i -nerved or
tinctly 3- nerved, without gibbosi- obscurely 3-nerved, with small
ties. gibbosities at their bases.

Lateral sepals free. Lateral sepals with their bases

adnate to the claw of the lip.
Lip flat or concave, without Lip with a more or less dis-
median ridge, not truncate at the tinct median ridge ; blade trun-
base, free. cate or hastate at the base ; a

margin connecting the claw with
the column.
Anther-cells opening in front. Anther-cells opening laterally.

The first .species of Piperia was originally described as
Spirantlies Utialascliensis Sprengel * and generally known as Habc-
naria UnalascJiensis Wats. In tlie Kew Index this species is
referred to Henniuium coiigcstuvi, a plant from northern India.
We have no specimen of the latter, but the identity is so unlikely
that I did not think of making a comparison when I visited Kew
last summer. P. UnalascJiensis \sx\o Herminhnn, however, for that
genus is distinguished by the total absence of a spur besides by
different structures of the column. Kraenzlin recognizes both Pla-
tantJiera Unalaschcnsis and P. foctida. It is true that no one seems
to know exactly what 5^;m;////r5 UnalascJiensis '^^^xQ:\^\^t\ really was,
but I know of no plant except the present species, which agrees with
Sprengel's short description. Kraenzlin placed P. Unalaschcnsis
among the scapose species and P. foctida among the foliose, but
in the description he characterizes the latter as having the leaves
near the base. I cannot find any character on which to base a
segregation of two species. The publication of Platanthera foctida
Geyer, is generally given as Hook. Journ. Bot. 7 : 376. 1855 ;
but there is no description, only the name and references to the lo-
cality where it was collected. The first real publication seems to
have been in King's Report, where Watson gives it as a synonym
under Habenaria foctida there described. Gymnadenia hmgispica
Durand f has been referred to this species, but this must be a mis-

* Syst. 3 : 708. 1826.
fPl. Pratten, loi. 1855.

634 RvDBERG : The American Species of

take as that species was described as having a spur longer than the
ovary, which is not the case with P. Unalascheiisis.

The second species of Pipcrin was first described as PlatantJiera
elegans Lindl.* from specimens collected by Douglas in north-
western America. It seems from the description as if more than
one species had been included therein as for instance " cmile squa-
inis parvis ranicntaceis, spica longa densa cylindracca." Although
this species has scale-like stem leaves, they are not by far so con-
spicuous as in the species that Bolander took for PL elegans and
that the spike is described as dense does not very well fit this spe-
cies. Not having any authentic specimen I naturally thought
that the name Plataut/iera elegans Lindley belonged to the species
named Habenaria elegans by Bolander or else the one that I have
described here as Piperia multiflora. I therefore described the
lax-flowered species under the name P. elongata. At Kew last
summer, I found only one specimen collected by Douglas and
this I refer to my P. elongata although its spike is more dense
than in my type, but not denser than some specimens of the same
species in our herbaria. It is, therefore, best to pass my P. elon-
gata into synonymy for the present.

With regard to Habenaria elegans Bolander it must be stated
that it was not properly published before it appeared in the Botany
of California, for in Bolander's Catalogue of the Flora of San
Francisco it is a nouicn nudum, without description or synonyms.
What Bolander's plant was is not unknown, for there is one speci-
men in the Torrey herbarium collected and named by Bolander.
This can scarcely be distinguished from Habenaria Michacli
Greene,t although it has a longer, less crowded spike.

A further study has revealed that several other species than
the ones enumerated in the place of publication of Piperia, must be
included in that genus, viz., Habenaria Michaeli Greene, and Gyni-
nadenia longispica, both mentioned above, Habenat'ia Cooperi
Wats.:}: and Habenaria niaritinia Greene. § Concerning H. Cooperi
Wats, it may be remarked that it was placed by Watson in that

*Gen. & Spec. Orchid. PI. 285. 1835.
t Man. Bay Reg. Bot. 306. 1894.
jProc. Am. Acad. 12: 276. 1876.
§ Piltonia, 2 : 298. 1892.

LlMNORCHIS AND PiPERIA NORTH OF MeXICO 635

section of Haboiaria which now constitutes LininorcJiis, but his de-
scription of the flower is the best hitherto given of any species of
Pipcria.

Synopsis of Species

Spur less than twice as long as the lip.

Stem leafy only at the base ; lip oblong ; spur slender, slightly exceeding the
lip. \. P. Unalaschensis.

Stem more or less leafy ; lip ovate.

Spur slightly exceeding the lip, very saccate. 2. P. Cooperi.

Spur nearly twice as long as the lip, only slightly clavate.

3. P. lancifoiid.
Spur 2-3 times as long as the lip, filiform.

Lip linear to lanceolate.

Spike very lax ; lip 4-5 mm. lon^ ; spur about 8-10 mm.

4. P. li'ptopetahi.
Spike dense ; lip ab^ut 6 mm. long ; spur 15-18 mm. long.

5. P. 1)1 u tijlora.
Lip ovate or ovate-lanceolate.

Spike elongated, lax ; leaves withering at anthesis.

Bracts linear-lanceolate ; stem leafy only at the base.

6. P. e Cleans.
Bracts ovate-lanceolate ; stem usually leafy. 7. P. longispica.

Spike short and very dense ; flowers crowded ; leaves withering before
anthesis.
Petals and sepals about 4 mm. long ; petals purplish or greenish.

8. P. Miclmeli.
Petals and sepals about 5 mm. long ; petals white. 9. /'. iiiaritiina.

I. PiPERIA Unalaschensis (Spreng.) Rydb. Bull. Torr. Club, 28 :

270. 1 90 1

Spirantlics Unalaschensis Spreng. Syst. 3: 708. 1826; Habc-

naria Scliisc]imareffianaQ\\?in\.\Jim\-Ae.^,2>'- '^9- 1828; Hermmiinn
UnalascJikcnsc ^€\<z\\&\\h. Fl. Germ. 13: 107.//. 6^; PlatantJicra
Scliisclimareffiaiia Lindl. Gen. & Sp. Orchid. 286. 1835; Habe-
naria foctida S. Wats. Bot. King's Exped. 5: 341. 1871 ; Plat-
antJicra foctida Geyer. as a synonym under the preceding ;
Habenaria Unalaschensis Ws-is. Proc. Am. Acad. 12: 277. 1877;
Montolivaea Unalaschensis Rydb. Mem. N. Y. Bot. Gard. i : 107.
1900.

Stem strict, slender, 3—5 dm. high, leafy only near the base :
basal leaves oblanceolate, obtuse or acutish, i — 1.5 dm. long, .8-3
cm. wide, withering at anthesis or soon after ; stem leaves bract-
like, linear-lanceolate to lanceolate, attenuate, .5—1 cm. long : spike
long and lax, 1—3 dm. long; bracts lanceolate to ovate-lanceolate,

G36 Rydberg : The American Species of

one-half to two-thirds as long as the flowers : these greenish, dis-
tinctly in spirals, 8-io mm. long : petals and sepals 2-4 mm. long,
upper sepal ovate, acutish ; lateral ones oblong-lanceolate, ob-

tusish : petals purplish green, somewhat fleshy,
lanceolate, oblique at the base ; lip oblong,
obtuse, slightly hastately lobed near the
base ; spur filiform or slightly clavate, a little
exceeding the lip but shorter than the ovary.

,^^^ (Fig- 26.)
^ ■ P. Unalasdiensis is common from south-

ern Alaska and Alberta to California and Colorado, and grows in
damp woods.

Type: '' Ins. Aleut."

The following locality is remarkable, being far out of the sup-
posed range :

Quebec: Jupiter River, Anticosti, i^d>2), John Macoiin.

2. Piperia Cooperi (S. Wats.)

Habcnaria Cooperi S. Wats. Proc. Am. Acad. 12 : 276. 1876.

Tall and strict, 3-10 dm. high, leafy below : basal leaves ob-
long lanceolate, acute, about i dm. long ; lower stem leaves lan-
ceolate, attenuate, 10-15 cm. long; the
upper reduced and bract-like, lanceolate,
1-2 cm. long : spike strict and lax, 1-3
dm. long ; bracts ovate-lanceolate, acumi-
nate, two-thirds as long as the flowers :
these yellowish-green, about 10 mm. long :
sepals and petals about 4 mm. long, ob- fig. 27.

tuse ; upper sepal ovate ; lateral sepals and
petals oblong lanceolate ; lip ovate, rounded at the apex, somewhat
hastate at the truncate base, but the lobes rounded, rather thick and
with prominent ridge in the middle ; spur thick, decidedly clavate,
about equalling the lip, but much shorter than the ovary. (Fig. 27.)

I have not seen the type, collected by Cooper at San Diego,
California, but the plants cited below agree with the description
and were collected in the same region. It is from these I have
drawn the description and figure.

Type: "On Clay hills near San Diego, California; Dr. J. G.
Cooper."

California: San Diego, 1884, C. R. Orciitt ; Point Loma,
1897, T. S. BnDuicgce.

LiMNORCHIS AND PiPERIA NORTH OF MeXICO

637

Fig. 2S.

3. Piperia lancifolia sp. nov.

Stem stout, 3-5 dm. high, the lower portion leafy; basal leaves
and lower stem-leaves lanceolate, attenuate, 10-15 cm. long, 1-2
cm. wide, withering after anthesis : spike many-flowered, but lax,
2-3 dm. long; bracts ovate, acute, striate,
about two-thirds as long as the flowers, or
the lower almost equalling them : flowers
greenish, 11-13 mm. long: upper sepal
ovate, obtuse, about 4 mm. long ; the lateral
ones slightly longer, oblong-lanceolate :
petals lanceolate, obtusish, oblique at the
base; lip about 4 mm. long ; blade round
ovate, rounded at the apex, truncate at the

base, scarcely at all hastate, thick, with prominent median ridge ;
spur filiform, slightly clavate, almost twice as long as the lip and
about equalling the ovary. (Fig. 28.)

This species is closely related to the preceding, differing in the
longer less clavate spur, scarcely at all hastate lip and broader
bracts. It is growing in cafions.

California: Sierra Santa Monica, 1892, H. E. Hassc, ^dyj
(type in herb. N. Y. Bot. Card ).

4. Piperia leptopetala sp. nov.

Stem slender, 3-4 dm. high, leafy only at the base : basal leaves
two, oblong-lanceolate, obtuse or acutish, about i dm. long, 1.5-

2.5 cm. wide, withering at anthesis or
soon after ; stem leaves all reduced
and bract-like, few, lanceolate, acute :
spike slender, lax, 1-2 dm. high ;
bracts lanceolate to linear-lanceolate,
acuminate, about half as long as the
flowers : these greenish, about i cm.
long : upper sepal lanceolate, obtuse,
about 4 mm. long ; lateral ones nar-
rowly lanceolate, acute, about 5 mm.
long: petals narrowly lanceolate, acute,
about equalling the upper sepal ; blade of the lip lanceolate,
obtuse, hastately toothed at the base ; median ridge rather obscure ;
spur filiform, not clavate, about twice as long as the lip and longer
than the ovary. (Fig. 29.)

In habit this species resembles most P. Unalaschensis, but the
sepals and petals are narrower and the spur is almost twice as long.

Fig. 29.

638

Rydberg : The American Species of

California : Mountains east of San Diego, 1850, C. C. Parry
(Mex. Bound. Surv., type in herb. Columbia Univ.) ; Plumas
county, 1875, Mrs. Austin.

Washington: Nesqually River, 1838-42, Wilkes Exp., 146,

in part.

5. Piperia multiflora sp. nov

Montolivaca elcgans Rydb. Mem. N. Y. Bot. Garden, i : 106,
in part as to specimen cited. 1900; not Reichenb. 1881 ; Piperia
elegans Rydb. Bull. Torr. Club, 28: 270, in part. 1901.

Stem stout, 4-6 dm. high, 3-4-
leaved only near the base : basal leaves
oblong, oblanceolate or obtuse or
acutish, 1-1.5 dm. long, 2-3 cm. wide,
withering at anthesis ; lower stem
leaves lanceolate, acute ; upper stem
leaves much reduced and bract-like,
lanceolate or linear-lanceolate, scat-
tered, attenuate, 1-2 cm. long : spike
very dense, 1-2 dm. long; bracts
linear-lanceolate, almost equalHng the
flowers : these greenish-white, spread-
ing, about 1.5 cm. long: upper sepal
lanceolate, acute, 4-5 mm. long ; lateral
sepals and petals linear-lanceolate, acutish or obtuse ; lip almost
linear, obtuse, only slightly hastate at the base, about 6 mm.
long; median ridge very low; spur 15-18 mm. long, filiform,
not clavate. (Fig. 30.)

This species is perhaps nearest related to P. elcgans, differing
however in the dense spike, the spreading flowers, and narrow
petals and sepals.

Washington: Gray's Harbour, 1838-42, Wilkes Exp ed., 1^4
(type in herb. Columb. Univ.); Cascade Mountains, \^%2, Brande-

ge^. 4-75-

Montana: Mission Range, 1883, W. M. Ca?iby,joy.

California : Monterey, Wm. Rich.

6. Piperia elegans (Lindl.) Rydb. Bull. Torr. Club, 26 : 270

in part. 1901

Platanthera elegans Lindl. Gen. & Sp. Orchid. PI. 285. 1835 ;
Montolinaea elegans Rydb. Mem. N. Y. Bot. Card, i : 106, in

LiMNORCHIS AND PiPEKIA NORTH OF MeXICO

C39

part, as to synonym. 1 900 ; not Reichenb. 1 88 1 ; Pipcria dongata
Rydb. Bull. Torr. Club, 28: 270. 1901.

Stem slender, strict, 4-7 dm. high : tuber ellipsoid, about 2 cm.
long, I cm. in diameter : basal leaves 2 or 3 ; blades lanceolate or
oblanccolate or rarely oval, acute or
obtuse, 8-15 cm. long, 1-3, sometimes
even 5 cm. wide ; stem-leaves much re-
duced, lanceolate, acuminate, 5-10 mm.
long : spike long and usually lax, 1.5-3
dm. long ; bracts lanceolate, acuminate,
from half to fully as long as the flowers :
these greenish-white, about i cm. long :
sepals about 5 mm. long ; the upper
lanceolate, acute ; the lateral ones
linear-oblong or lanceolate, obtuse :
petals lanceolate, acute ; blade of the

lip broadly or ovate-lanceolate, slightly hastate and truncate at
the base ; median ridge rather indistinct ; spur filiform, 10- 1 2 mm.,
about two and a half times as long as the lip and longer than the
ovary. (Fig. 31.)

Type: "In America horcali occidoitali, Douglas {Jiab. s. sp.
connn. Soc. Plort.).''

British Columbia: Vancouver Island, \^?>j, John Alacoiin.

Washington: W. Klickitat county, 1885, W. N. Siiksdorf.

Oregon: Grave Creek Hills, 1887, Thomas Howell; 1871,
Elihu Hall, jo6.

Idaho: Priest Lake near lower end, 1900, D. T. MacDoiigal,
168 ; Priest River valley, 134; Kootenai county, 1887, /. H.
Sandbcrg ; Wiessner's Peak, 1892, Samiberg, MacDougal &
Heller, 584 (broad leaved).

California: Santa Lucia Mountains, 1898, R. A. Plaskett,
16 J ; 1872, Mrs. Bancroft.

7. Piperia longispica (Durand)

Gymnadenia longispica Durand, PI. Pratten, loi. 1855.

Stem stout, 3—7 dm. high, more or less leafy below : tuber
ellipsoid, 3-4 cm. long, about 1.5 cm. thick, basal leaves and lower
stem leaves 2-4, lanceolate, acute, 1-1.5 dm. long, 2-3.5 cm. wide,
withering about the time of anthesis : upper stem leaves reduced,
1-3 cm. long, lanceolate : spike many-flowered, but not dense, 1—3
dm. long; bracts ovate-lanceolate, .5—1 cm. long, acuminate:
flowers greenish, about 1.5 cm. long : upper sepal ovate, obtuse.

640

Rydberg : The American Species of

about 5 mm. long ; lateral sepals oblong-lanceolate, obtuse : petals
broadly lanceolate ; blade of the lip ovate-hastate, distinctly

auricled, and truncate at the base ;
spur filiform, two and a half times as
long as the lip. (Fig. 32.)

I have not seen the type of Gym-
nadoiia longispica, which was de-
scribed from a fragmentary specimen
collected by Pratten, /. c, it consisted
only of the upper portion without any
leaves. Durand's species has been re-
garded as P. UnalascJiensis but this
cannot be so, for Gymnadenia long-
ispica was described as having a spur longer than the ovary,
which throws it out of P. UnalascJiensis. As the spike is described
as long and lax it must have been either P. cicgans, P. Icptopetala
or the species described here. As this is the only species with
broad bracts, the name G. longispka must belong here, unless
it is a species unknown to me. The only discrepancy is that the
spur is described as clavate, which is scarcely the case.

California: Mokelumne River, Mr. Rich ; Monterey, 1850,
C. C. Parry (Mex. Bound. Surv.) ; Cucamonga Mts., 188 1, 5. B.
& IV. F. Parisli, 1157 ; Santa Monica range, 1892, H. E. Hasse.

Fig. 32.

8. Piperia Michaeli (Greene)

Habcnaria clegans Boland. (Cat. PI. San Franc. 29) ; S
Bot. Calif. 2: 133. 1880; not PlatantJicra cicgans Lindl
Habenaria Michaeli Greene, Man. Bay-
Reg. Bot. 306. 1 894 ; Montoliveia elegans
Rydb. Mem. N. Y. Bot. Gard. i : 106,
in part as to synonym, 1900; Piperia
elegans Rydb. Bull. Torr. Club, 28 :
270, in part as to synonym.

Stem stout, at flowering time leafless,
2-3 dm. high : tuber ellipsoid, 2-4 cm.
lonQ-, 1-1.=; cm. wide: basal leaves
elliptic or oblanceolate, about 1.5 dm.
long, 4 cm. wide, withering before blooming ; stem-leaves
ous, small and bract-like, lanceolate to ovate-lanceolate, acu

Wats.

1835;

Fig. 33.

numer-
minate,

LiMNORCHIS AND PiPERIA NORTH OF MeXICO

641

1-2 cm. lonfj : spike very dense, 5-15 cm. lonp^: flowers about
I cm. long, greenish : upper sepal ovate, about 4 mm. long ;
lateral ones oblong-lanceolate, obtuse : petals lanceolate, acute ;
blade of the lip ovate, obtuse, scarcely hastate, obtuse or truncate
at base ; spur filiform, about two and a half times as long as the
lip and a third longer than the ovary. (Fig. 33.)

Type : " Open hills, under oaks, etc., from near Livermore
southward."

California : San Luis county, 1886, G. W. Michael ; Stans-
bury Valley, 1876, Harry Edwards ; San Francisco, 1838-42,
Wilkes Expedition, 1554, in part ; " California," Bolander.

Washington: Gray's Harbor, 1838-42, Wilkes Expedition,
1554, in part ; Nesqually River, 146, in part.

9. Piperia maritima (Greene)

Habenaria maritima Greene, Pittonia, 2 : 298. 1892.

Stem stout, 2-3 dm. high, at flowering time leafless : tuber
almost spherical, 2-2.5 cm. in diameter: leaf -blades oblong,
obtuse or acute, about i dm. long,
3-4 cm. wide, withering before anthe-
sis ; stem leaves reduced and bract-
like, numerous, ovate-lanceolate, acu-
minate, 1-2 cm. long : spike short and
very dense, 4-10 cm. long; bracts
about two-thirds as long as the flow-
ers : these 10—12 mm. long, white:
upper sepal ovate, obtuse, about 4
mm. long ; lateral sepals oblong-lance-
olate : petals broadly lanceolate ; blade
of the lip elliptic, obtusish, scarcely
hastate, somewhat truncate at the base,
as well as the petals and sepals, white, thin ; spur filiform, about
2.5 times as long as the lip and longer than the ovary, (Fig. 34.)

This differs from all the other species in its stoutness and
white flowers. It resembles most P. Michaeli in habit, the short
spike, numerous scale-like stem leaves and the absence of root
leaves at the time of flowering.

Type : " On dry hills near the sea at Point Lobos, near San
Francisco."

California: San Francisco county, 1892 and 1893, MicJiener
& Biolctti.

Fig. 34.

642

Rydberg : The American Species of

CROSS-REFERENCES AND INDEX

Blephariglottis Raf. , 605-6.
COELOGi.ossuM Hartman., 605.
Gymnadenia. , 605-6

longisfica Durand = P. longispica.
Gymnadeniopsis Rydb., 605-6.
Habenaria Willd., 605.

boreal is Cham. = L. borealis. ^

— li viriJiflora Cham. = L. viridi-
flora.
brevifolia Greene = L. brevifolia.
Cooperi S. Wats. = P. Cooperi.
dilatata Coult. = L. borealis.
— Hook. = L. dilatata.
— Torr. = L. media.
dUatatiformis Rydb. = L. borealis.
elegnns Bolander = P. Michaeli.
foetida S. Wats. = P. Unalaschensis.
gracilis S. Wats. = L. striata.
graminea Spreng. = Plathantheragra-

minea Lindl.
hyperborca R. Br. -= L. hyperborea.
— Gray = L. major, L. media, L.

Huronensis.
— Rothr. = L. striata.
— Wats. ^=. L. viridiflora.
leucostachys S. Wats. = L. Thurberi,

L. leucostachys.
marilivia Greene = P. mariiima.
Michaeli Greene = P. Michaeli.
pedicellata S. Wats. = ?.
saccata Greene ^ L. striata.
Schischinareffi-ana ^ P. Unalaschensis
sparsiflora Wats. =- L. sparsiflora.
strirta Rydb. ^ L. striata.
Thurbei-i A. Gray = L. Thurberi.
Unalaschensis Wats. = P. Unalas-
chensis.
Herminium., 633.

Unalaschetise = P. Unalaschensis.
LiMNORCHis Rydb., 606.
Arizonmca Rydb., 629.
Behringiana Rydb., 620.
BOREALIS (Cham.) Rydb., 621.
BRACHYPETALA Rydb., 6r6.
BREVIFOLIA (Greene) Rydb., 631.
CONVAT.I.ARTAEFOI.IA (Lindl.) Rvdb.,

628.

LiMNORCHis Rydb.

DILATATA (Pursh) Rydb., 622.

— LINEAR! FOLIA Rydb., 623.

dilatatiforviis Rydb. =: L. borealis.
ENSIFOLIA Rydb., 629.
FOLIOSA Rydb., 622.
FRA(iRANS Rydb., 623.
GRACILIS (Lindl.) Rydb., 627.

GRAMINIFOLIA Rydb., 627.

Huronensis (Nutt. ) Rydb., 619.
HYPERBOREA ( L. ) Rydb., 620.
hypcrb rea Rydb. = L. viridiflora, L.

major.
LAXIFLORA Rydb., 630.

LEPTOCERATITIS Rydb., 624.

LEUCOSTACHYS (Lindl.) Rydb., 625.
— ROBUSTA Rydb., 626.

MAJOR (Lange) Rydb ,617.

MEDIA Rydb , 618.

PURPURASCENS Rydb., 615.

SPARSIFLORA (S. Wats. ) Rydb., 631.

STRICTA (Lindl.) Rydb., 614.

Thurberi (A. Gray) Rydb., 624.

VIRIDIFLORA (Cham.) Rydb., 616.
Lysiella Rydb., 606.
MONTOLIVAEA Reichenb. , 635.

ei.egans Reichenb., 639.

elegans Rydb. ^ P. elegans, P. mul-
ti flora, P. Michaeli.

Unalaschensis Rydb. = P. Unalas-
chensis.
Orchis L. , 605.

dilatata Pursh r=L. dilatata.

dolichorhiza =^1., 608.

Huronensis Nutt.= L. Huronensis.

hyperborea L. ^ L. hyperborea. -,

Koenigii Retz. = L. hyperborea.

ROTUNDIFOLIA Pursh., 605.

PERULARiALindl., 605.
PiPERiA Rydb., 632.

Cooperi (S. Wats.) Rydb., 636.

ELEGANS (Lindl.) Rydb., 638.

elegans Rydb. = P. multiflora, P.
Michaeli.

elongata Rydb. — P. elegans.

LANCIFOLIA Rydb., 637.

LEPTOPETALA Rydb., 637.

LiMNORCHIS AND PiPERIA NORTH OF MEXICO

643

Pli'ERiA Rydb.

LONGISPICA (Durand) Rydb., 639.
MARITIMA (Greene) Rydb., 641.
MiCHAELi (Greene) Rydb., 640.
MULTIFLORA Rydb., 638.
Unalasciiknsis (Spreng. ) Rydb. 638.
Platanthera Rich., 605.

convallariaefoUa Lindl. = L. conval-

lariaefolia.
dilalata Beck = L. Huronensis, L.
dilatata.
— Lindl. = L. borealis, L. dilatata.
— Torr. = L. major, L. dilatata.
gracilis Kraenzlin := L. striata.

— Lindl. - = L. gracilis.
graminea (Spreng.) Lindl., 627.
graminea Lindl. = L. graminifolia.
foetida Geyer=P. Unalaschensis.
Huronensis Lindl. ^^ L. Huronensis.
hyperborea Lindl. = L. hyperborea.
— a genidna Kraenzlin ^L. hyper-
borea, L. viridiflora, L. major,
L. media.

Platanthera Rich.

— ,3 convallariaefoUa Kraenzlin =

L. convallariaefolia.
— ,3 major Lange == L. major.
— y dilatata Kraenzlin = L. dila-
tata, L. Huronensis, Habenaria
pedicellata S. Wats., P. Coop-
eri, L. graminifolia, and L.
borealis.
— y leucostachys Kraenzlin = L.
leucostachys, L. brevifolia,
Platanthera Ghiesbrechtiana
Rich. & Gal., L. sparsiflora,
L. Thurberi, L. en.sifolia.
Koenigii Lindl. = L. major.

— /? dolichorhiza Lindl. = Orchis
dolichorhiza =: ?.
leucostachys Lindl. = L. leucostachys.
Schischtnareffiana = P. Unalaschen-
sis.
stricta Lindl. = L. stricta.
Unalaschensis Kraenzlin ^ P. Una-
laschensis.

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 20.

SEEDS AND SEEDLINGS OF ARISAEMA Till-
PHYLLUM AND ARISAEMA DRACONTIUM

BY ROSINA J. RENNERT.

NEW YORK
1902

[Reprinted from Bulletin Toekey Botanical Club, 29 : 37-5L 28 Jan., 1902].

I Reprinted from the Bitlletin of the Torrey Hotanical Club, 29: January, 1902.1

Seeds and Seedlings of Arisaema triphyllum and Arisaema

Dracontium

By Rosina J. Rennert
(Plate 3)

Some interesting facts in the germination of the seeds of Aris-
aema Draconiuiui have been recently described by Dr. MacDougal.
and the author has devoted a greater part of the last year to the
comparative study of the morphology and physiology of the seed-
lings of this species and of A. triphyllum. Some attention was
also given to seedlings of a hybrid between the two species and
those of A. flavum from Sikkim, India.

It was previously found by Dr. MacDougal that the plumule
of A. Dracontium emerges from the cotyledonary sheath only in
a small proportion of the seedlings, and that only a few of those
emerging exhibited a development sufficient to carry the lamina
of the first leaf up into the air and unfold it to the light.*

According to Braun, the closely related species Arum macu-
latum displays a similar behavior. He says " So spielt auch an
der Keimpflanze die Niederblattbildung eine grossere Rolle, als
bei Calla, in dem nach Irmisch auf dem Cotyledon zunachst
mehrere unterirdische vegetirende Niederblatte folgen bevor ein
Laubblatt, und zwar erst im zweiten Jahre zum Vorsche in kommt."t
This statement seems to have escaped the notice of Scott and Sar-
gant who have recently made an extended study of these seed-
lings.!

The seedlings of A. triphyllum, on the other hand, send up a
plumule as soon as the hypocotyl is firmly established in the soil,
while the hybrid between A. Dracontium and A. triphyllum dis-
plays a curiously intermediate type of germination. The hypo-

* MacDougal, D. T. Seedlings of ^ma,fw«. Torreya, 1 : 2. 1901.

t Braun. Ueber das Vorkommen mehere Hullblatter am Kolben von Arum
maculatum L. , Calla palustris, und Richardia Africana Kth. Verb. bot. Ver. Brand-
enburg, I : 94. 1859.

% Scott and Sargant. On ihe Development of Arum maculatum from tbe Seed.
Annals Bot. 12:399. 1898.

37

38 Kennert : Seeds and Seedlings of Arisaema

cotyl, in this case, increases in size at first without sending up a
plumule, and it is only after the seedling has exhausted the entire
food supply of the endosperm and has separated from the seed
that the plumule appears.

Like A. tripliyllum the Indian species, A. flavurn, follows the
normal type of germination, and its plumule arises shortly after
the hypocotyl emerges from the seed. The seedling is only about
one half the size of that of A. tripliyllum.

As the general structure of the seedling of A. triphyllum re-
sembles most closely that displayed by A. Dracontium, this species
was chosen as a basis for comparison in the study of the curious
variations of A. Dracontmvi.

The fruits of A. Dracontmvi and A. triphylliini are very simi-
lar. In both the scarlet berries, which have a sweetish, slightly
acrid taste, are closely crowded upon the spadix. The berries of
A. Draconthun are larger and fleshier than those of A. tripJiyllum
and contain from three to six seeds, while those of A. triphyllum
are one- or three- seeded. In shape the seeds of both are some-
what ovoid. At the hilum, the seed of A. Dracontitini is generally
puckered into two or three ridges and the short stalk of the funic-
ulus is conspicuous, while in A. triphyllum the seed is flattened in
the hilar region, but also bears a prominent funiculus. Elsewhere
the seeds are well rounded, except when flattened by the pressure
of other seeds in the same berry. Both seeds bear a rudimentary
aril which consists of a small disc-shaped fleshy mass situated imme-
diately inside the coats at the hilar end of the seed, and extending
as a core down the funiculus. The average size of the seeds of A.
Dracontium is 3.5 mm. in transverse and 4.25 mm. m longitudinal
diameter, while the average seed of A. triphyllum is 4 mm. in trans-
verse and 3.5 mm. in longitudinal diameter, almost exactly revers-
ing the proportions of A. Dracontium. When only a few are pro-
duced in a berry, the seeds of A. Dracontium are larger in both
axes than those of A. triphyllum, which develop singly. When,
however, a number of seeds mature in the same berry they develop
in such a position as to exert a lateral pressure upon each other
and in consequence the transverse diameter remains shorter. In
these cases the seeds of A. Dracontium do not attain the size of
even the smallest of A. triphyllum. On the whole, however, A.

TRIPHYLLUM AND ArISAEMA DrACONTIUM 39

Dracontiinn has the lar<^cr seed. The surface of both seeds is
finely pitted and sometimes minute reddish spots are to be seen
upon them. A. tripJiylltiui has a yellowish color and A. Dracon-
tiinn a reddish-yellow blush. This color is due to substances in
the wall of the inner coat of the seed, as the testa is translucent.

In the general microscopic structure the seeds present only
slight dissimilarities. The outermost row of cells in the testa of
both species are brick-shaped and are set close together with their
long axes at right angles to the radius of the seed. In both
the walls of this outer row of cells are mucila<rinous, swelline
greatly when they come in contact with water. In A. Dracontitim
they are slightly larger and their outer walls are considerably
thicker. A cuticle covers this row of cells in both species. Some
of tile cells below the outer row of the testa contain tannin,
others are filled with mucilage in which raphide cells are im-
bedded, still others have a red coloring matter within them. It
is to these latter cells that the dotted appearance sometimes noted
on the seeds is due. The tegmen consists of three rows of tabu-
lar cells with very thick mucilaginous walls. In A. Dracontitim
these walls are orange -colored, and in A. triphylluni yellowish in
color and so serve to give each seed its distinctive appearance.
There is a cutinized lamella on either side of the tegmen cells.

The seeds of both plants are orthotropous and the embryo is
imbedded in an abundant endosperm. In A. triphyllum the endo-
sperm appears entirely homogeneous to the naked eye, while in A.
Dracontiiim there is a horny layer next the seed coats. Upon mi-
croscopic examination, however, but slight differences are evident.
The outer portion of the endosperm of A. Dracontiiun consists of
3 to 5 rows of cubical aleurone cells with walls capable of great
swelling in water. These cells contain granular proteid globules,
a single crystalloid, and also a small amount of fat. The aleurone
cells of A. triphylluni are of exactly the same character, but are
generally only from two to four rows deep, and also a trifle smaller.
Within the layers of aleurone cells, making up by far the greater
part of the endosperm, are large starch-bearing cells which have
thin cellulose walls. Numerous crystalloids are imbedded in the
starch of these cells. These crystalloids are slightly more nu-
merous in the seeds of A. Dracontiinn. In both cases, however,

40 Rennert : Seeds and Seedlings of Arisaema

they are very abundant near the aleurone cells and become scarcer
toward the embryo at the center of the endosperm. It will be
seen that the only difference between the two seeds lies in the size
of the cells which compose the aleurone layer and the seed coats.
It follows from the greater size of the seed of A. Dracontiiim that
this species has a food supply slightly greater than that of A. tri-
pJiylliim.

Both resting embryos occupy a position near the base of the
seed. In fact the aleurone cells near the micropyle are obliterated
and the base of the hypocotyl of each embryo is close to the teg-
men at this point. The cavity in which the embryo lies, but does
not entirely fill, corresponds in general shape to that of the embryo
and extends as an axis from the micropyle to the hilar region
through the center of the seed. Its sides are formed of starch-
bearing cells, aleurone cells close to the cavity at the hilar end,
while the tegmen bounds it at the opposite end as described above.
The embryos are club-shaped and small in proportion to the
amount of endosperm contained in the seeds. In A. Dracontiuni
they are 2 mm. long and .75 mm. thick, while those oi A. tripliyl-
hun are slightly larger, measuring 2.75 mm. in length and .5 to
.875 mm. in thickness. In A. triphylhim three fourths of the
length of the embryo consists of cotyledon, the remainder is taken
up by the hypocotyl. This relation is different in A. Draconthun,
for the cotyledon in this case bears a slightly greater proportion to
the entire length of the embryo. A slight constriction marks the
insertion of the cotyledon on the hypocotyl, and a very small pro-
tuberance at the base of the hypocotyl is the incipient radicle.

Five regions are already differentiated in the resting embryo of
each species, the dermatogen, procambium, root cap, meristem
and fundamental parenchyma. The dermatogen is continuous
with the epithelial layer which covers the entire embryo and, at
the slit for the emergence of the plumule in the cotyledonary tube,
is deflected back to line the plumular cavity. In the plumule of
A. tripliylluui dermatogen is already clearly differentiated but in
A. Dracontiuni it is not so evident. The dermatogen is composed
of a single row of narrow cells with their long axis at right angles
to the longitudinal axis of the embryo. They are filled with
aleurone grains and each contains a crystalloid. The procambium

TRIPHYLLUM AND ArISAEMA DrACONTIUM 41

Strands which foreshadow the bundles have already been laid down
in the cotyledon, stem-bud and hypocotyl. Near the tip of the
cotyledon the procambium strands branch and anastomose irregu-
larly and therefore no definite number or arrangement of them
can be determined, but at the base the number becomes constant
and each strand occupies a regular position. The plan of arrange-
ment in both species is similar. In cross section the strands mark
out' a crescent in which the largest is at the middle of the bow. In
A. tyiphylltan which has regularly five strands, two lie on either
side of the central ones, while the cotyledon of A. Dracontium
shows the same arrangement of its five principal strands, but has
in addition one or two smaller ones. When there are two, these
extra strands occupy positions on either side of the large middle
trace. When only one extra strand is retained, it invariably is at
the right. In the tubular portion of the cotyledon, the largest
strand is in the thickest part of the wall. The strands are com-
posed of six or seven rows of narrow elongated cells which have
their long axis four to six times the length of their radial, and
parallel with the longitudinal axis of the embryo. In the stem-bud,
procambium strands have also been differentiated. In A. Dracon-
tium the strands in the plumule are faint and there is some varia-
tion in the number of the procambial regions. They are never
more numerous than three and at times only the pro-cambium
of the future midrib has been laid down. In A. triphylliim five
distinct procambium regions can be distinguished in the plumule.
Procambium strands are also present in the hypocotyl and occupy
fully one half of its entire diameter. They are continuous with
the procambium of the stem-bud and form at the base of the
hypocotyl the rudimentary root stele. At the base of the hypo-
cotyl the root-cap may be distinguished, made up of several rows
of rather large cells which stretch across its entire base and are
continuous at the sides with the dermatogen of the embryo. In
the outer rows the cells are empty. Meristem tissue is to be found
at the stem-bud and at the base of the hypocotyl where the root
takes its origin. These cells are very small and nearly isodia-
metric. Their nuclei are large and fill almost the entire cell.
Parenchyma cells make up the rest of the embryo. They are two or
three times the size of the epidermal cells and are packed with

42 Rennert : Seeds and Seedlings of Arisaema

starch. Scattered through the inner tissue are large cells filled
with proteids in which a crystalloid also is embedded. These pro-
teid cells are distributed through the tissues in those regions where
the pro-cambium strands first make their appearance.

The embryos of the two species are similar in shape, position
in the seed and general structure. A. Dracontiinn differs from A.
fnpliyllmii in four particulars : (i) Its size, (2) The proportion
which the length of its cotyledon bears to its hypocotyl, (3) The
number of procambium regions to be found in the cotyledon and
(4) The state of differentiation of the procambium strands in the
plumule. Here already the tendency in A. Dracontiiun to delay
the development of the plumule is evident.

The Germination of the Seedlings

In both species when water reaches the seeds the mucilaginous
walls of the outer row of cells of the testa swell, while the cells
below, pulled out by the swelling outer row and pushed out by the
increase in size of the mucilaginous tegmen and the walls of the
aleurone cells, separate and show an irregular alternation of large
and small cells, which are, of course, empty. The seeds now in-
crease considerably in size by the absorption of water, and the action
of the moisture slowly reduces the outer row of the testa cells to
a gelatinous film about the seeds. The mucilaginous tegmen is
also soon dissolved, while all that remains of the seed coat is the
inner part of the testa. The tannin in the vesicles contained within
this becomes diffused through the whole of the cellulose coat and
makes it more resistant to the disintegrating action of the soil.
This part of the coat alone adheres to the seed until the last par-
ticle of endosperm is dissolved. The mucilaginous cells of the
seed coat and the thick walls of the aleurone cells serve to in-
crease the absorption of water, while the outer row of the testa is
instrumental in bringing about the attachment of the seed to the
soil.* The cellulose layer of the testa, impregnated with tannin,
protects the endosperm during germination. When water reaches
the embryo in the seed it swells and fills the cavity in which it lies,
so that its epithelial cells are in contact at the sides with the starch

•* Klebs. Heilrage zur Morphologic und Biologie der Keimung. Untersuch. a. d.
Botan Institut z. Tubingen, i : 536. 1885.

TRIPHYLLUM AND ArISAEMA DrACONTIUM 43

cells of the endosperm. Here the diastase formed by the epithe-
lium of the embr)-o can begin to act upon the stored starch of the
seed and a proteolytic enzyme to dissolve the crystalloids. The
starch of the endosperm is absorbed before the crystalloids, and
when the cells in contact with the embryo have been emptied of
their contents the cell walls are pushed back by the advancing
growth of the cotyledon and the enzyme acts upon the contents
of the cells next outside. The proteids of the aleurone cells do
not become soluble until late in the history of the seedling. The
cell walls of the endosperm do not seem to be acted upon by any
enzyme, and are wholly intact until the seed separates from the
seedling.

While the epithelial cells are secreting the enzymes, the coty-
ledon increases in length and by its elongation pushes the hypo-
cotyl, bearing the stem-bud, through the seed coats at the nii-
cropyle. A. Dracontkini requires fully a month longer than A.
h-ipliylhnn for its hypocotyl to protrude. When the hypocotyl
has wholly emerged from the seed it is directed down into the
ground at right angles to the plane of the cotyledon, which is still
in the seed. This is effected by means of a bend in the cotyledon
at the place where it leaves the seed. This downward pull of the
hypocotyl serves to tilt the seed up, and is sufficient when the seed
has not been planted deep to break through the ground and bring
the seed to the surface. In all cases it loosens the earth about the
seedling, and so renders it easier for the pumule to make its way
through the soil.

A great part of the cotyledon is confined within the seed during
its entire development. The length which the cotyledons attain
varies in both species, but those oi A. Dracontinin are on the aver-
age shorter than those of A. tripJiylluin. The average length of
the A. Dracoiitiinn cotyledons outside of the seed is 3 mm., while
those of A. tripltyl.iiin reach an average length of 7 mm. The
number of the cells in the cotyledon is not increased by its elonga-
tion, for its entire growth is due to the increase in size of the
already existing cells. The epithelial cells, which in the resting
embryo had their axis perpendicular to the longitudinal axis of the
embryo, now in the region of the cotyledon have this relation com-
pletely changed. Their long axes are parallel with the longitudinal

44 Rennert : Seeds and Seedlings of Arisaema

axis of the embryo. The cells of the parenchyma have also taken
on an elongated form. The only region in the cotyledon where
new cells are added is that occupied by the procambium and fibro-
vascular bundles. Here the cell division is very active. Radial
growth takes place only as the contents of the endosperm cells is
absorbed. The shape, arrangement of cells and manner of elon-
gating are exactly similar in the cotyledons of A. Dracontmm and
A. triphylhnn. It is only in the fibrovascular system that any dif-
ference between them has been discovered. Here in both cases,
as in the procambium strands in the embryo, the bundles near the
tip, branch and anastomose so that their distribution is irregular.
At the base, however, the number of bundles is constant. A. Dra-
contium as a rule possesses six bundles : one of these strands is very
weak and occupies the same position as the sixth procambium strand
in the cotyledon, /. e., at the right of the largest central bundle. The
left hand strand which corresponds to the seventh in the resting
embryo disappears before the cotyledon has proceeded very far
with its development. The cotyledon of A. tripliyllujii has five
bundles arranged as in the resting embryo. In a few instances
chlorophyl has been found in that part of the cotyledon of A.
Dracont'mni which protrudes from the seed both when the plumule
was functional and when it was not. These cases, however, are
uncommon.

Up to this point there is almost absolute identity in the germi-
nation of these two species : the only difference between them lies
in the various sizes which the cotyledons attain. After this stage
in the germination has been reached, however, various differences
in the order of development of the organs become apparent.

The rudiments of all the structures of the mature corm are
present in the hypocotyl of both A. triphylhan and A. Dracontiiwi
and the formation of the corm is brought about by the enlarge-
ment of the base of the hypocotyl and the laying down in it of the
definitive tissues. In both it increases in bulk at least to a small
extent before the first primary root is sent out. The hypocotol
of A. Dracontmm enlarges at a very much more rapid rate in the
first stages of germination than does A. triphyLlum and generally
attains a considerable development before the root appears. A
seedling o{ A. Dracoiitiiim with two roots each only i mm. in length

TRIPHYLLUM AND ArISAEMA DrACONTIUM 45

may have a corm 4 mm. in diameter while in a well-developed
seedling of A. tripJiylliiDi with a strong system of roots and a
plumule well grown, the corm may be only 175 mm. in diameter.
The enlargement of the corm of A. triphylliiDi becomes marked
only after the leaf is well established as a photosynthetic organ.
In those seedlings of A. Dracontiuni in which the plumule does
not become functional the corm, when its growth is completed,
averages 4.5 mm. in diameter. When, however, the seedling has
a functionating leaf the corm may attain a diameter of 6 mm.
The corm of ^. triphyllum may have a diameter of 5.75 mm. at
the end of the season, but in many cases it is much less.

The enlargement of the hypocotyl is due to the increase both in
size and number of its cells. As its growth is rather radial than
longitudinal, the epithelial cells become isodiametric in shape.
The storage cells increase in size and in number in all directions
and so maintain the same relative dimensions. The starch grains
contained in these cells are several times as large as those of the
endosperm of the seed. The procambium cells of the hypocotyl
are very active, dividing both in their longitudinal and transverse di-
ameters and the fibro-vascular system soon becomes differentiated.
Raphide cells are numerous in the corm and in all parts of the
seedlings of the two species. As the corm enlarges, a layer of
periderm arises on the outside, from which are cut off tangentially
five or six rows of empty flattened cells. In addition to the pro-
tection they afford the corm, these periderm cells are instrumental
in effecting a separation between the cotyledon and the corm, after
the food supply of the endosperm has been absorbed. They
grow between the cotyledon and the corm and separate the two
by proliferating cells from their surface. In several instances
chlorophyl has been formed in the two or three rows directly be-
neath the layers of periderm. The only difference in the develop-
ment of the corm of the two species lies in the earlier enlargement
of the hypocotyl of A. Dracontium and the greater average size of
its completed corms.

The time at which the roots of A. Dracontium appear varies
markedly. In some cases the hypocotyl enlarges greatly, becom-
ing, as was described above, about 3 mm. in diameter before the
first root arises. In other instances the roots are developed as

46 Rennert : Seeds and Seedlings of Arisaema

soon as the hypocotyl has found its way into the ground. Seed-
hngs of A. Draconiiiim vary greatly also in the number and kinds
of roots which they send forth. Those with the most well de-
veloped system have two primary roots from the base of the
hypocotyl and later a thick root arising adventitiously from the
the nodes. This secondary root, becomes contractile and trans-
versely ridged for part of its length. One of the primary roots
also often becomes contractile and helps to draw the corm deeper
in the ground. A secondary root, however, is developed only in
those seedlings which also send up a plumule and a few even of
these have none. The seedlings in which the plumule has not
started into activity have as a rule two primary roots, one of which
becomes thickened and assumes the function of contraction. In
some cases neither of the primary roots is contractile and in others
only one short thin root is produced. A direct correlation be-
tween leaf and root development is here evident. Variations in
the structure as well as the number of the roots occur. The con-
tractile roots which are for the most part simple have been ob-
served occasionally to be branched. In another instance (the
single case in which two contractile roots occurred in the lot of
A. Dracontiinn seedlings observed) two contractile roots were fused
for part of their length. The root system oi A. tripliylhnn is very
much better developed than is that of A. Dracoiitmm, for the roots
are both longer and more numerous. The seedlings have always
two and often three primary roots which reach a length of 5 cm.
and in addition after the seedling is well under way, three thick
adventitious roots appear, budding from the nodes. These roots
are contractile and show the same transverse ridges as the con-
tractile roots of A. Dracontium. In some cases they pull the
hypocotyl down as much as 2 or 3 cm. below the level at which it
germinated. After the secondary roots become firmly established,
the primary roots grow no more and the greater part of the entire
root function is discharged by the contractile roots which bear root
hairs near their tips and become about 7 cm. long.

The only difference in microscopic structure between the con-
tractile, secondary and primary roots consists in the greater radial
elongation of the cells of the inner cortex of the contractile roots.
In A. Dracontium the root stele of the primary root shows a diarch

TRIPHYLLUM AND ArISAE.MA DrACOXTIUM 47

structure and the secondary roots are tetrach as a rule. Some,
however, have been observed to be triarch ; in A. tripliylhim both
primary and contractile roots are triarch. After the contraction
of the roots the outer cortex cells are very much strained and
twisted, but the radial elongation of the inner cells protects the
stele from any distortion. The roots of A. Draconthini are more
variable in all respects than those of A. tripliyllum.

The stem-bud of A. Dracontiiim and A. triphyllinn have the
same structure in the resting embryo, with the exception that the
procambium stands in the bud leaves have reached a more
advanced stage of differentiation in A. triphyllinn. During
the first season's growth, however, the structure of the bud in
the two species has an entirely different history. Like the
roots the plumules of the A. Dracontium seedlings vary greatly
in the state of development which they attain. As a rule the
plumule of A. Dracontinui develops to at least a small degree
after germination, forming a lamina and a short petiole which al-
though it may break through the cotyledonary sheath, yet rarely
reaches the air and is often represented only by a rudiment 2 mm.
in length remaining permanently enclosed by the cotyledon and en-
wrapping in its turn the stem-bud of the next year. This undeve-
loped plumule is cut off at the end of the season by the formation
of a periderm between it and the corm in the same way as the coty-
lendon is separated from the corm. About 10 per cent, of the
seedlings of A. Draconthini produce functional plumules. They
appear at different stages during the growth of the corm, gener-
ally when it has already attained a considerable size and in some
cases only after the seedling has entirely separated from the seed.
Its method of emergence from its enclosing cotyledon is exactly
the same as that of A. triphylluui which produces a plumule nor-
mally. As soon as the root has secured a firm hold on the soil
and the hypocotyl is only slightly enlarged, the plumule with its
blade recurved parallel to the petiole and rolled in at the mar-
gin breaks through the tubular part of the enclosing cotyledon
through a longitudinal slit which is already present in the resting
embryo. The plumule becomes green rapidly while the petiole
at the base of the lamina straightens out and the folded blade un-
rolls and grows larger.

48 Rennert : Seeds and Seedlings of Arisaema

The functional plumules of A. Dracontiuvi exhibit a tendency
toward degeneration. A comparison of the plumules of the two
species will well demonstrate the particular instances. The fully-
grown leaves of A. triphyllum are ovate, the largest being T)^ mm.
in length and 30 mm. in width, while the smaller are 25 mm.
by 13 mm. The functional leaves of A. Dracontium vary greatly
in the shape and size of the lamina and in the length of the petiole.
In outline they range from broadly elliptical to nearly circular and
all bear a mucronate point at the tip as does A. tripliylliim. The
largest leaves of ^. Dracontium are about 25 mm. in length by 22
mm. in width, but the greater number are smaller, some being
only 6 mm. by 3 mm. The margin of nearly all leaves of A. tri-
phyllnvi are very finely serrulate ; while some leaves of A. Dracon-
titini display a tendency to serrulation, more often they are entire.
The upper surface of the A. triphyllum leaf is covered with faint
whitish lines ; this appearance is due to the presence of elongated
raphide cells in the mesophyll directly beneath the epidermis. A.
Dracontium has numerous raphide cells in the mesophyll but they
are not so large as those of A. triphylbim and cannot be distin-
guished without the aid of the microscope. The under surface of
the leaves of A. triphylhiin is covered with a waxy bloom and the
upper surface also shows some wax. In A. Dracontium the wax is
very much thinner on the under surface and barely discernible on
the upper. As a consequence the leaf oi A. Dracontium wilts very
quickly after being removed from the plant. The development of
stomata on both leaves is about the same. The leaves of both
species are pinnately net- veined. In A. triphyllum one pair of the
lateral branches is more strongly marked than the others, fore-
shadowing distinctly the plan of the mature trifoliolate leaf. In A.
Dracontium the plan of venation is more generalized. The lateral
veins have all equal value and are neither so well marked as some
nor so faint as other lateral veins of the leaf of A. triphyllum.
The mesophyll of the A. Dracontium leaf is not so well developed
nor so well supplied with chlorophyll as is that of A. triphyllum.
Those plumules of A. Dracontium which become assimilatory
organs are functional for a shorter period than those of A. triphyl-
lum, for they wither upon the plant much sooner. Tho.se of long-
est duration last not more than three months while some send up

TKIPHYLLUM AND ArISAEMA DrACONTIUM 49

a petiole only, the blade being shriveled before it expands. Mon-
strosities seem to be very frequent in the leaves of A. Dracontiiun,
for instance, the displacement of the mucronate point from the tip
to the middle of the under surface of the midrib or the multipli-
cation of these points upon the under surface. The blade is also
often lobed upon one side and in one instance both sides of the
leaf showed this peculiarity.

The petioles of the leaves of both species are streaked with
red, the color being most abundant near the base. The petiole of
A. Dracontium is considerably shorter than A. triphylbim and the
fibro-vascular system is weaker on the whole although some stems
of ^. Dracontium and A. tripJiylluvi show scarcely any difference.
In the arrangement of tissue within the stem there is exact simi-
larity. The bundles are arranged to form an open cylinder gen-
erally, in A. Dracontium, three on each side of the largest bundle.
Those stems which attain the greatest development may have in
addition a bundle running through the center of this cylinder,
while in A. triphvUum the fibro-vascular system may include still
another bundle, completing the cylinder and making nine bundles
in all.

The development of the buds goes on during the growth of
the seedling. Each rudimentary leaf arises as a hollow elevation
enclosing the next youngest. At the end of the season the stem-
bud of A. Dracontium has four bracts which enwrap the bud
at the center and A. tripJiyllum agrees exactly with this plan in
structure. All but the innermost of these enclosing bud leaves
remain as rudiments upon the corm protecting the bud. The
fourth grows large enough to protect the leaf as it pushes through
the soil the second year, but remains at its base as a membraneous
sheath.

The second season all A. Dracontiiiui corms, like those of ^.
tripliyllwn, produce one trifoliolate leaf enclosed at the base by a
sheath which, in those cases where no plumule is produced, must
be regarded as the first leaf sent up by the plant. The functional
leaf of the second season is trifoliolate normally, but is subject to
great variation, such as the whole or partial fusion of two of its
lobes or the complete obliteration of one of them. Even when the
leaf is regularly trifoliolate great variations occur in the shape of

50 Rennert : Seeds and Seedlings of Arisaema

the individual leaflets. In some, the side leaflets may be narrowly-
lanceolate while others may approach an elliptical shape. The
mid leaflet is generally broader than the lateral ones, but here too
a variety of forms may occur. At the end of the second season's
growth there is still a slight diflerence in the size of the corms of
those plants of A. Dracontuun which have borne a plumule the first
year and those that have not. The root systems are, however, simi-
lar; six adventitious roots are developed in each. Except in the
retarded development of its plumule, the development of the stem-
bud of A. Dracontiinn agrees with that shown by A. tripJiylluui.
The repression of the plumule does not seem to be correlated with
any variations in the stem-bud of the second season, since varia-
tions of leaf form occur as frequently in second-year plants which
produced a functional plumule the first season.

The principal differences between the seedlings of A. Dracon-
tiiivi and A. tripJiylluin consist not only in the reduction and vari-
ability of the roots, the variation and repression of the plumule on
the part of ^. Dracontiiini, but also in the precocious enlargement
of the corm. This difference arises as soon as germination starts ;
in A. Draco)itiwn the foodstuff of the endosperm is employed
directly to build up the hypocotyl at the expense of the devel-
opment of root and plumule. In A. inphyHuJii, however, as
soon as the hypocotyl breaks through the seed coats, the roots
are sent out and attain some development at once, the plumule
then appears, and in normal cases it is only after this assimilatory
organ is well established and the seedling has separated from the
seed that the hypocotyl begins to enlarge. In A. triphylhini the
endosperm furnishes the material which is necessary to bring the
root and plumule to an advanced stage of development, and the
food material for the hypocotyl is supplied by the assimilation of
the plumule. The early enlargement of the corm and the repres-
sion of the plumule can scarcely be held to be due to a patholog-
ical condition of the seed or to an unfavorable environment, as the
seeds planted were perfectly sound and the plumules in the em-
bryos of those from the same lot which were examined showed no
evidence of the attacks of parasites or any abnormality, while the
conditions under which the plants were grown corresponded to the
normal environment of these plants as was well demonstrated by

TRIPHVLLUM AND ArISAEMA DrACONTIUM 51

the vigorous growth of .L tripJiylluin which was subjected to the
same conditions.

On the other hand the small proportion of functional plumules
developed by seedlings of A. Dracoiitiuui and, in the cases in which
a leaf was produced, the great variations in shape, the small size, the
generalized form and venation, the delicacy of the lamina and the
weak development of wax upon it, the shortness of the petiole and
the frequent appearance of monstrosities, all are indications of de-
generation and lead to the conclusion that this seedling is losing
its power to produce a normal assimilating plumule. This con-
clusion is borne out by an examination of the internal structure.
The petioles of the smaller leaves of A. triphylhim show the same
arrangement of fibrovascular bundles as the largest of A. Dra-
contiiivi and a series of increasingly smaller plumules which may
easily be obtained from a number of seedlings shows a successively
more generalized development of fibrovascular system until in the
least developed system it becomes reduced to a mere rudiment. The
highly organized fibrovascular system of the hypocotyl, which is
of very little service to the seedling without a plumule and only a
weakly developed root system, is still retained and bears evidence
that the present type of seedling which brings a plumule to various
stages of imperfect development may be a degeneration from a con-
dition like that of the seedlings of A. tripliyllum with a plumule
normally functional.

So far as has been observed this type of germination is of no
benefit to the seedlings. It does not seem to enable the corms to
escape from the danger which they incur of being torn up by
animals, for the development of the seedling without a strong
root system of course precludes any great downward pull upon
the corm by the contraction of its roots. Those plants of A.
Dracoiitmni which develop a leaf and correlated with this a
stronger root system have invariably been found buried deeper
than the plants w^hose plumule is not functional, for although in
the plumuleless seedlings there is no upwardly directed force to
bring the corm near the surface such as the plumule exerts in
forcing its way through the ground yet on account of the weak-
ness of the root system the tendency is such as to keep the corms
very near the depth at which the seeds germinated. The repres-

62 Rennert : Seeds and Seedlings of Arisaema

sion of a plumule, of course, prevents the destruction of young
seedlings by grazing animals ; but the well- developed plumule
contains enough raphide cells to protect it from attacks of this
kind and the advantage which the plants enjoy in this respect is
largely over-balanced by positive disadvantages. This variation,
therefore, can scarcely have been perpetuated because of the ad-
vantage as a means of defense which plumuleless seedlings pos-
sess. As far as can be discovered at present this variation is of
no benefit to the plant but, on the contrary, there are great disad-
vantages in this method of germination, which tend to throw the
plant entirely upon the food supply of the parent form for another
season.

The germination of A. Draconthmi consists essentially in the
conversion of the hypocotyl of the embryo into the corm of the
seedling by the transfer of the food material contained in the
endosperm. In some aroids,* this resorption of the endosperm
and conversion of the base of the hypocotyl into a corm takes
place before the embryo leaves the seed ; that is to say, the de-
velopment of the seed is not completed until a bulbiform embryo
with true fibro- vascular bundles and no trace of cotyledon has been
formed. An example of this type is Spatliycma foctida. When
the seed of this plant germinates the stem-bud pushes out of the
micropyle and breaks through the ground first ; later adventitious
roots spring from the nodes. No primary root is produced. It
will be seen that the only difference between the germination of
this seed and the sprouting of the first year's corm of A. Dracon-
tium is the fact that in the former case the seed coats which sur-
round the bulbiform embryo must be penetrated by the stem-bud.
If the corm formation of the seedling of A. Dracontiiun took
place within the seed coats (and this could be brought about
simply by the arrest of the elongation of the cotyledon) the two
cases would be exactly similar. In fact the cotyledon of A.
Dracontuiiii is varying in just this direction, for it often displays a
tendency to be shorter than the cotyledon of A. tripliyUiiui as
has been pointed out above. The delay in germination at first,
moreover, may be another indication of a tendency to carry on
the entire development of the corm within the seed.

* Engler. Monographiae Phanerogamarum, Araceae, 1 1 : 34.

TRIPHYLLUM AND ArISAEMA DrACONTIUM 53

Owing to the lack. of time and material the development of the
corm of Spatliycnia within the seed could not be traced but an ex-
amination of the literature of the subject brought out the fact
that many aroids manifest a tendency to carry on the development
of the plant as far as possible before actual germination, /. c,
emergence from the seed and development of assimilatory organs

occurs.*

Summary

The seeds of the two species present only a few unimportant
differences in shape. In histological structure they are similar.
The embryos are comparatively small and are imbedded in a co-
pious farinaceous endosperm. The only essential distinction be-
tween them consists in the slightly greater amount of food material
in the seed of A. Dracontiiim, the extra procambium strand of its
cotyledon and the weaker development of procambium in the
plumule of its resting embryo. The first .stages of the germina-
tion of the two seeds are of the same character and consist in the
emergence of the hypocotyl and stem- bud from the seed coats at
the micropyle, by means of the elongation of the cotyledon.

As the development of the seedlings proceeds, the production
of roots and a plumule takes precedence in A. triphvllmn while in
A. Dracontiiun the enlargement of the hypocotyl begins at once
and the growth of the root and plumule is retarded. This pre-
cocious development of the corm often takes place to such a de-
gree as to entirely inhibit the production of a functional plumule.
The same differences between the two species in regard to the
fibrovascular development is exhibited by the seedlings as is dis-
played by the resting embryo. The structure of the stem-bud
which develops upon the corm during the first season's growth
is absolutely the same in both species.

The seedling of A. Dracontium is diverging from what seems to
be the normal type of germination in Arisaema, i. c, the develop-
ment of an assimilatory plumule and the production of a corm by
means of the product of the photosynthetic activity of this organ,
and is tending to produce a corm without the aid of a plumule by
the direct transfer of the food material of the endosperm to the

* Engler. Monographiae Phanerogamarum. Araceae, ii : 34, 35-
Griffith. Trans. Linn. Soc. 20: 274-276. 1847.

54 Rennert : Seeds and Seedlings of Arisaema

hypocotyl. This type of germination does not seem to be due to
a diseased condition of the seed or embryo or to an unfavorable
environment nor does it appear to be advantageous to the young
seedling. It might be suggested therefore that it is an expression
of an inherent tendency in aroids to carry the young plantlet as
far as possible in its development before an assimilatory organ is
produced. If this conclusion be correct we have in A. Dracontimu
a transition stage between a seedling forming its corm outside of
the seed coats with the aid of a plumule as in A. triphyllum and a
corm formation within the seed coats at the expense of the endo-
sperm as is the case with SpatJiyema foetidus.

Explanation of Plate 3

Fig. I. Seedling of A. Dracontium showing hypocotyl just emerging from seed.

Fig. 2. Seedling of A. Dracoiitiuni. Hypocotyl has begun to enlarge, short root
has appeared.

Fig. 3. Seedling of A. Dracontitun. Root has elongated, root hairs have arisen.

Fig. 4. Seedling of A. Draconthtm. Hypocotyl has enlarged greatly, first root
has elongated, second root is just appearing.

Fig. 5. Seedling of A. Dracontium. Hypocotyl has attained full size of first
year's corm.

Fig. 6. Seedling oi A. Dracontium. Specimen in which the root system has been
only slightly developed.

Fig. 7. First-year corm of A. Dracontium. Formation of periderm at base of
cotyledon has separated corm from seed.

Fig. 8. Seedling of A. Dracontium which has produced plumule.

Fig. 9. Seedling of A. Dracontium. More advanced ; note single contractile root,
developed only in those cases where plumule is sent up.

Figs 10, 11, 12. Variations in form of plumule in those seedlings of A. Dra-
contium which send up a leaf.

Figs. 13, 14. Second-year leaves of A. Dracontium when a plumule had been
developed by seedling.

Figs. 15, 16, 17, 18, 19. First leaves produced by A. Dracontium corms which
have been developed without the aid of a plumule.

Fig. 20. Seedling of .4. triphyllum. Hypocotyl just emerging.

Fig. 21. Seedling of ^. triphyllum. Cotyledon lengthened, hypocotyl has begun
to enlarge, root has arisen.

Fig. 22. Seedling of ^. /';7)^/^)'//«w- Second root appearing.

Fig. 23. Seedling of ./. triphylhan. Second root has increased in length.

Fig. 24. Seedling of A. triphyllum. Plumule has emerged from cotyledonary
sheath.

Fig. 25. Seedling of A. triphyllum. Petiole of plumule has lengthened, a third
root has appeared. Hypocotyl perceptibly enlarged.

Fig. 26. Seedling of A. triphyllum. Contractile root formed, plumule fully ex-
panded.

Fig. 27. Seedling of A. hybrida. Plumule produced.

Figs. 28, 29, 30. Variations in form of leaf [jroduced second year by .4. hybrida.

Ik'i.u. ToKKEV Ci.rii, J9

Pi..

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 21

TWO NEW WESTERN MOSSES

BY R. S. WILLIAMS.

NEW YOEK
1902

[Reprinted from Bulletin Torrky Botanical Club, 29 : 66-68. 25 Feb., 1902].

[Reprinted from the Bulletin of the Torrey Botanical Club, 29: February, 1902.]

Two new Western Mosses

By R. S. Williams
(With Pi.atf.s 4 and 5)

Eurhynchium Taylorae

Evidently dioicous ; no ^ flowers found. In loose tufts with
stout, creeping stems (in places denuded of leaves and with dense
tufts of radicles) up to 20 cm. long and 0.44 mm. in diameter ;
cross sections show a well-defined central strand of numerous
small cells and outer wall of about three rows of small thick-walled
cells; branches rather short (i to 2 cm.), irregular and often
distant ; stem and larger branch leaves loosely spreading, ob-
tusely pointed, the branch leaves often truncate or praemorse at
tip, decurrent, serrulate all round to a little above the base, re-
flexed on margin above and near base and costate about ^ up,
the costa ending in spine and more or less papillose below; stem
leaves measure up to 2.5 x 1.5 mm., middle branch leaves 1.5
mm. and upper branch leaves 0.875 mm. long, the lowest branch
leaves very small, roundish and often ecostate ; inner perichaetial
leaves ecostate or very faintly costate, abruptly narrowed to a
slender flexuous serrulate point and irregularly toothed and
lacerate at base of point ; leaf-cells mostly linear-vermicular,
median .004 x .06 to .08 mm., apical very short and irregular,
alar forming an elongated not well-defined group of pale cells
about .008 mm. wide and two to four times longer, the basal cells
somewhat thickened and pitted, especially toward costa ; capsule
mostly horizontal, contracted more or less under the mouth when
dry, without lid 2.5 by 1.5 mm., lid with obliquely rostrate beak
2 mm. long, stout, smooth seta up to 15 mm. high.annulus of two
or three rows of cells; peristome teeth very slender-pointed, striate
below, papillose above, hyaline bordered, the segments more or
less split with mostly two somewhat appendiculate cilia between ;
stomata roundish, .025 mm. in diameter; spores nearly smooth,
up to .014 mm.

In size this plant most nearly approaches E. Orcganiun of any
of our North American species but is less regularly pinnate with
smooth seta, etc. It much more closely resembles E. striatuin of
Europe, but is a more creeping plant with less divided stems and
leaves never acutely pointed as in that, which also lacks the spine

at apex of costa.

66

67 Williams : Two new Western Mosses

Collected by J. B. Leiberg on decaying logs in streams, April,
1889, Traille River Basin, Idaho (172). Named for the artist who
has made the excellent drawing which accompanies this descrip-
tion, Miss Alexandrina Taylor.

Brachythecium Pringlei

Monoicous ; $ flowers abundant on fruiting stems. Growing
in compact tufts with ascending, more or less branching stems 4
—5 cm. long and 0.25 mm. in diameter; cross sections of stem
show a distinct central strand and 3 or 4 rows of thick-walled
outer cells ; branches mostly short, rather irregular, with curved
tips; stems and branch leaves very similar, up to 2.5 x 1.5 mm.,
secund, mostly broadly ovate with short acute point, long decur-
rent, concave, scarcely or not plicate, borders flat, entire or slightly
serrulate above ; leaf-cells somewhat thickened and near base,
pitted ; median cells linear, about .04 x .005 mm., alar cells mostly
about . 012 X. 016 mm., often forming a distinct convex cluster
packed with chlorophyll ; inner perichaetial leaves erect, grad-
ually lanceolate-pointed, the point somewhat variable in width,
nearly entire and ecostate ; seta distinctly papillose in upper part,
about I cm. high ; capsules (not quite mature in only specimens
seen) ovate-oblong, not quite symmetrical, nearly erect, with lid
2.5 mm. long; lid highly conical, .8 mm. high; annulus nar-
row, of I or 2 rows of cells ; teeth about .430 mm. high and
.065 mm. wide at base, rather broadly and irregularly pointed,
hyaline bordered, rather irregularly striate and somewhat papillose
below, and distinctly papillose above ; basilar membrane of en-
dostome extending about two fifth up, segments more or less split,
cilia mostly 2 and not appendiculate ; stomata slightly elongated,
about .025 mm. long; spores (immature) smooth, .015 mm.

This plant with its short-pointed, scarcely plicate and secund
leaves with vein sometimes forking, has much the appearance of a
Linuiobiiim [Hygroliypmuii), but the seta rough above and the
highly conical lid seem to relate it most closely to BracJiytlicciimi,
which also contains all the first-mentioned characters. Collected
by C. G. Pringle in the Huachuca Mts., Arizona, July, 1884 (22).

Explaiia>.ioii of Plates

Drawings made with a magnification twice the diameter reproduced on the plates.

Plate 4
Fig. I. Plant, natural size.
Fig. 2. Stem leaf, X I5-

Figs. 3, 4, 5. Lower, middle and upper branch leaf, X ^S-
Fig. 6. Alar region of stem leaf, X l-^o.

Williams : Two new Western Mosses

68

Fig. 7. Apex of same, X 120.

Fig. 8. Cross section of costa.

Fig. 9. Part of cross section of stem, X 87.

Fig. 10. Perichaetial leaf, X ^S-

Fig. II. Capsule, X 7-S'

Fig. 12. Stoma, X 210.

Fig. 13. Portion of peristome with annulus, X *20

Fig. 14. .Spores, X *20.

Plate 5

Fig. I. Plant, natural size.
Fig. 2, 3. Stem leaves, X ^5-

Fig. 4. Branch leaf, X ^S-

Fig. 5. Alar region of stem leaf, X ^20.

Fig. 6. Apex of same, X 120.

Fig. 7. Median cells, X 210.

Fig. 8. Cross section of leaf, X 120.

Fig. 9. Part of cross section of stem, X 120.

Fig. 10. Antheridial bud, X ^5-

Fig. II. Antheridia and paraphyses, X 45-

Fig. 12. Perichaetial leaf, X 17-

Fig. 13. Capsule with upper part of seta.

Fig. 14. Oblique view of seta showing papillae, X ^62.

Fig. 15. Stoma, X 210.

Fig. 16. Part of peristome and capsule, X ^20.
Fig. 17. Spores, X ^20.

Bull. Torrf.y Ci.i'b. 29

Pl.

W3-

EURHYNCHIUM TAYLORAE

15l'i.l. Torkev Cll'b, 29

PL. 5

BRACHYTHECIUM PRINGLEI

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 22

STUDIES ON THE ROCKY MOUNTAIN FLORA

VII.

BY PER AXEL RYDBERG.

NEW YORK
1902

[Reprinted from Bulletin Torrey Botanical Club, 29 : 1L5-160. 24 March, 1902].

Studies on the Rocky Mountain Flora— VII

By Per Axel Rydberg

Aquilegia Columbiana

Perennial with a short thick rootstock : stem 6-10 dm. high,
glabrous or sparingly and finely pubescent above; basal leaves
twice ternate ; petioles 1-2 dm. long ; segments rounded, obovate,
2-3 cm. long, more or less glaucous beneath, 2-3-cleft and deeply
crenate at the apex ; lower stem-leaves similar to the basal leaves,
but with short winged petioles, the upper similar but often with
larger segments : sepals and spurs very bright red ; laminae yel-
lowish : sepals ovate-lanceolate, acute, 15-25 mm. long, exceed-
ing the spur, widely spreading or reflexed : lamina very short,
scarcely 3 mm. long, truncate; spur 12-18 mm. long, thick, ab-
ruptly contracted about the middle : follicles erect, densely pubes-
cent ; styles about 1 2 mm. long.

This species is somewhat intermediate between A. formosa and
A. truncata and has been mistaken for both. It has the habit, the
spur and sepals of the former and the short truncate lamina of
the latter.

Washington: Montesano, 1898, A. A. & E. Gertrude Heller,
3936 (type in herb. N. Y, Bot. Garden) ; upper valley of the
Nesqually, 1896, 0. D.Allen, 2^g ; 1889, G. R. Vasey, 130;
Humptulips, 1895, F. H. Lamb, 1180; Puget Sound, Wilkes
Ex peel., 6^6.

British Columbia: Banff, 1897, Zoe W. Palmer.

Alaska: Short Bay, 1895, Gorman, 103; Disenchantment
Bay, i8g2, F. Fiinston, 121.

Idaho: Florida Mt., 1892, A. Isabel Mulford.

Aquilegia thalictrifolia

Perennial wath a short rootstock : basal leaves twice ternate ;
petioles 1-2 dm. long; segments broadly obovate-cuneate, 2-3-
cleft and crenate at the apex, 1-1.5 cm. long, bluish-green : stem
2-5 dm. high, glabrous below, finely viscid puberulent above :
stem-leaves subses'sile or very short-petioled, otherwise similar to
the basal ones : flowers yellow: sepals 10—18 mm. long, ovate to
ovate-lanceolate, acute: laminae of the petals 7-12 mm. long,
truncate or rarely rounded at the apex ; spur very slender from

145

146 Rydberg : Studies on the

the base, 4-5 cm. long : follicles without the styles, about 3 cm.
long and 5 mm. wide, strongly reticulate and pubescent, strongly
curved above.

This species is nearest related to A. cJirysantka with which it
has been confused, but differs in the smaller flowers, blunter
sepals, smaller and bluer segments of the leaves and the more
curved follicles. It grows at an altitude of i 500-3000 m.

Colorado: Canon City, 1873, E. L. Greene (type in herb.
Columbia University); Colorado Springs, 1893, De Alton Saun-
ders; Grand Canon of the Arkansas, 1873, Brandegee ; Bear
Creek Canon, 1892, C. S. Sheldon.

Texas: West Texas, 1888, Nealley.

Arizona: Mt. Humphrey, 1897, R. E. Kunze.

Aquilegia Eastwoodiae

Aquilegia ecalcarata Eastwood, Zoe, 2: 226. 1891. Not
Hortul. ; Steudel. 1841.

This has been regarded as a spurless form of A. micrantha
Eastw., but the form of the terminal segments of the leaves is dif-
ferent from any American species, being rhombic in outline and

acute.

Aquilegia oreophila

Aquilegia coernlea alpina A. Nelson, First Rep. Fl. Wyo. 78.
1896. Not A. alpina L. 1853 ; Aquilegia coendea \2iX. fiavescens
Jones, Cont. West. Bot. 8 : 2. 1898. '^olA.flavescens^dXs. 1871.

This is evidently what Jones regards as A. flavescens Wats.,
but is not Watson's species, for he describes the spur as being
curved. Specimens of both numbers cited by Watson, viz. j5 and
j(5, are in the Columbia University herbarium and these as well as
the plant cultivated under the name A. flavescens at Harvard have
curved spurs and can not be referred as a variety to A. coendea.
Therefore, I can not see why Mr. Jones' A. depauperata should
not pass into synonomy, being the same as the true A. flavescens.
The flowers of A. oreophila resemble most those of A. pubescens
Coville, but the habit is different and nearer A. coendea.

Delphinium alpestre

A low cespitose perennial : stems several, about i dm. high,
puberulent and viscid above, densely leafy ; leaves digitately di-

Rocky Mountain Flora 147

vided into about 5 divisions, finely and sparingly puberulent when
young, dark green ; petioles 4-5 cm. long ; divisions of the blades
1.5-2 cm. long, cuneate-obovate in outline, divided half-way into
oblong mucronate lobes : inflorescence short and few-flowered ;
pedicels ascending, i cm. or less long, viscid ; bractlets minute :
sepals dark blue, oblong, obtuse or the upper acute, viscid-puber-
ulent ; spur 8-9 mm. long : upper petals blue and yellowish, 2-
toothed at the apex ; lateral petals 2-cleft ; lobes lanceolate :
follicles not seen.

This is not closely related to any of the American species. It
has the cespitose habit of D. glaiicescens, but is a much smaller
plant. It grows among rocks at an altitude of 3300 m. or more.

Colorado: Mountains northwest of Como, 1895, Crandall &
Cozven, 184S (type in herb. State Agric. Coll., Colo.) ; West
Spanish Peak, 1900, Rydbcrg & Vreeland, 6218.

Delphinium multiflorum

A tall perennial with a woody caudex ; stem about i mm.
high or more, viscid above, glabrate below : leaves palmately
divided into 5-7 divisions, glabrous and glaucous, pubescent only
on the margins and veins ; petioles 1-2 dm. long, slightly dilated
at the base ; segments obovate-cuneate in outline, 5-7 cm. long,
first cleft about half-way and then again cleft into lanceolate, ovate
or oblong acute lobes : inflorescence long and dense, often
branched; bracts linear; pedicels ascending, 1-L5 cm. long,
densely viscid-pubescent : bractlets small, linear, close under the
calyx : sepals light blue with darker median lines or blotches
towards the tips, oblong-oval, obtuse or the upper acutish, finely
puberulent; spur thick, straight or slightly curved, 12-15 nini.
long, almost horizontal : petals of the same color as the sepals ;
the lateral ones cleft only at the apex, often wavy-toothed : follicles
erect, densely viscid-pubescent.

This is nearest related to D. aiadatum A. Nelson on one hand
and to D. occidentale on the other. From the former it differs in
the pubescence, which in D. cuadatmn is strigose, in D. nntlti-
florum viscid. D. occidoitalc has much darker flowers and more
acute sepals. D. niidtifloruui grows along streams and in damp
meadows or open woods at an altitude of 2000-2500 m.

Wyoming: Spread Creek, 1897, F. Tzueedy, ijg (type in herb.
N. Y. Bot. Gard.) ; Yellowstone Lake, 1888, R. S. Williams;
Laramie Mountains, 1899, Charles ScJmcJmt ; between Upper

148 Rydberg : Studies on the

Buffalo Fork and Du Noir River, 1899, C. C. Curtis ; Snake
River, 1899, Aven & Elias Nelson, 6^0 j.

Idaho : Henry's Lake, 1897, Rydberg & Bessej, ^oyp.

Colorado: North Park, 1896, C. F. Baker.

Delphinium Brownii

Perennial : stem about i m. high, glabrous and shining through-
out, simple : leaves palmately divided into about 5 divisions,
puberulent ; lower petioles 8-10 cm. long ; divisions 4-6 cm. long,
oblanceolate in outline, once to twice cleft into lanceolate, acute,
■ somewhat spreading lobes : inflorescence a rather lax raceme ;
bracts linear ; pedicels slender, 1—2 cm. long, almost erect, curved
and twisted ; upper portion sometimes slightly puberulent ; bract-
lets linear-subulate, small, 3-4 mm. long : sepals dark blue or
purple, oblong-oval, obtuse or the lower and upper acutish, 10-12
mm. long : lateral petals also blue or purplish ; blade cleft only
about one third its length ; upper petals of the same color or
paler, obliquely cuneate, acute ; spur short and rather thick, i cm.
or less long, straight and ascending : follicles erect, glabrous.

This is nearest related to D. glaiicnm, but differs in the puberu-
lent leaves with narrower segments, the lax raceme with more
erect pedicels and the darker flowers.

Alberta : Banff, 1893, Addison Brown (type in herb. Colum-
bia University).

Alaska: Porcupine River, 1891,/. H. Turner.

Delphinium elongatum

Tall perennial with a short woody caudex : stem glabrous and
glaucous up to the inflorescence, which is slightly strigose : leaves
digitately divided into about 7 divisions, glabrous and glaucous
beneath ; petioles fully i dm. long ; divisions of the blade oblance-
olate in outline, 5-7 cm. long, cleft into lanceolate, acute lobes :
inflorescence strict and elongated ; bracts small, linear, almost fili-
form ; pedicels 1—2 cm. long, at first ascending, in fruit almost
erect ; bractlets minute, subulate : sepals and petals dark blue, the
former oval, obtuse or acutish, about i cm. long ; spur short,
straight, about 1 2 mm. long : lateral petals cleft at the apex, more
or less toothed: follicles erect, 12-15 ^nm. long, strigose-puberulent.

In some respects nearest to D. glmicum, but differing in the
puberulent pod, strigose pubescence at least on the pedicels, and
the narrower divisions and lobes of the leaves. It grows at an alti-
tude of about 2000 m.

RocKv Mountain Flora 149

Colorado: Larimer county, 1895, Crandall &■ Coiven, /j
(type in herb. N. Y. Bot. Garden) ; Villa Grove, 1 896, F. Clements,
338 ; Middle Park, 1861, C. C. Parry, 83 ; Baxton's Ranch, 1890,
Craudal/, 13^^.

Montana: Lima, iSgi^, Rydberg.

Delphinium diversicolor

Perennial with a thick fleshy root, the branches of which are
often fusiform : stem about 4 dm. high, simple, strict, densely vis-
cid-pubescent above : leaves finely pubescent, palmatcly dissected
into linear lobes ; petioles of the lower leaves 3-5 cm. long, slightly
dilated below ; segments 1-3 cm. long, about 2 mm. wide : inflor-
escence a narrow raceme ; pedicels .5-1 cm. long, or in fruit 1.5
cm., densely viscid, almost erect: sepals dark blue, oblong-oval,
obtuse, 10-13 iTini- long, slightly pubescent ; spur almost straight,
horizontal : petals all white with blue veins and yellowish below ;
the upper about 5 mm. long, obliquely spatulate, obtuse ; the lat-
eral ones slightly longer, curved ; the blades deeply 2-cleft ; each
lobe of the form of one third of a circle : follicles densely hairy.

This species is perhaps nearest related to D. simplex and D.
Burkei. From the former, it differs in white lateral petals and the
pubescent leaves, and the latter is described as subscapose. The
type was collected in a bog, at an altitude of 2000 m.

Montana: Rattlesnake Creek, Beaver Head county, 1887,
Frank Tweedy, 3^ (type in herb, Columbia University).

Aconitum tenue

Perennial with small tubers about 8 mm. long and 4 mm.
thick : stem very slender, about 3 dm. high : leaves few ; the lower
with petioles 5-8 cm. long, the uppermost subsessile : blades
glabrous, thin, shining, almost pentagonal in outline, 5-7-divided to
near the base ; divisions rhombic in outline, 3-cIeft and the lobes
oft.en again cleft, the ultimate divisions oblong-linear to linear-lan-
ceolate, acute : inflorescence racemose, 2-6-flowered : pedicels
erect and bent near the glabrous blue calyx : hood 15-18 mm.
long, boat-shaped, sHghtly saccate, almost semicircular in outline ;
lateral sepals rounded ovate, only slightly oblique, 10-12 mm.
long; lower sepals oblong, obtuse at the apex, 8-10 mm. long :
fruit unknown.

The type was collected in a damp canon among rocks at an
altitude of about 2000 m. It is perhaps nearest related to the
Alaskan A. delphinifolmm.

150 Rydberg : Studies on the

South Dakota : Sylvan Lake, Black Hills, 1892, P. A. Ryd-
berg, soy (type in herb. Columbia University).

Aconitum atrocyaneum

Perennial with thickened-fusiform roots : stem stout, 4-6 dm.
high, leafy, glabrous below, densely glandular viscid and purplish
above: lower leaves with petioles 5-10 cm. long; blades about
6 cm. wide, glabrous, pentagonal in outline, 5-7-divided to near
the base ; divisions rhombic-cuneate to oblanceolate, usually twice
cleft ; ultimate segments lanceolate acute ; upper stem-leaves sim-
ilar but with short petioles or those of the inflorescence lanceolate,
toothed and often with recurved basal lobes : inflorescence race-
mose, many-flowered, leafy : pedicels 1-3 cm. long, erect, very
viscid : flowers dark indigo blue, more or less pubescent : hood
deeply saccate, helmet-shaped, about 18 mm. long, with the front
line more or less concave : beak somewhat porrect, acuminate, the
lower margins nearlyhorizontal : lateral sepals broadly rounded-
ovate, somewhat oblique, 12-15 mm. long : lower sepals 9-12 mm.
long, oblong, obtuse.

This species is perhaps closest related to A. rainosum A. Nel-
son, but the divisions of the leaves are broader and the hoods are
different, having a porrect beak, while in A. ramosiivi the beak is
short and directed downward. The flowers are also darker and
the inflorescence more viscid. It grows at an attitude of 2,500
-3,400 m.

Colorado: Boreas, 1897, C. S. Craiuiall (ty^e in herb. N. Y.
Bot. Garden ; cotypes in herb. State Agric. College, Colo.) ;
mountains about Steamboat Springs, 1891, C. S. Crandall ; near
Ironton, 1899, C. C. Curtis ; Cameron Pass, 1896, C. F. Baker.

Utah: Uinta Mountains, 1873, T. C. Porter.

Aconitum porrectum

Perennial with thick fusiform roots : stem stout, 6-7 dm. high,
leafy, glabrous below, densely viscid-pubescent above ; basal leaves
glabrous, with petioles about i dm. long ; blades reniform-pen-
tagonal in outline, 6-10 cm. wide, 5-7 divided to near the base;
divisions rhombic or rhombic-oblanceolate, variously cleft above,
cuneate at the base : inflorescence racemose or somewhat panicu-
late, not leafy; pedicels ascending, 1-3 cm. long: sepals dark
blue, ciliate on the margins ; hood very saccate, helmet-shaped ;
beak acuminate, porrect, horizontal or even slightly ascending,
15-18 mm. long ; lateral sepals broadly obovate or semi-reniform,

RocKv Mountain Flora 151

about 12-14 nini- ^ong, very oblique; lower sepals lanceolate or
oblong, acute, 7—10 mm. long.

This is closely related to the preceding, but differs in the
broader and less deeply divided divisions of the leaves, the shorter
lower sepals, the more open and less leafy inflorescence, and the
more oblique lateral sepals. It grows at an altitude of 2400-
2800 m.

Colorado: Coffee Pot Spring, 1894, Cranda// (type in herb.
N. Y. Bot. Garden ; cotypes in herb. State Agric. College, Colo.) ;
Graymount, 1892, Crandall ; foothills, Larimer county, 1895,
Q'andall & Cowe 11 ; Avsip^hoQ Pass, 1891, Crandall, 36 ; Lake
City, 1 88 1,/. 5. Newberry ; North Park, near Teller, 1884, C. S.
Sheldon.

Aconitum glaberrimum

Perennial with a tap root : stem almost i m. high, perfectly
glabrous : leaves 5-divided, thin, perfectly glabrous, the lower long-
petioled ; the uppermost subsessile ; divisions oblanceolate to
rhombic-oblanceolate in outline, 5—10 cm. long, cuneate and entire
at the base, irregularly doubly cleft above ; lobes or teeth lanceo-
late, acute or acuminate : inflorescence compound ; branches and
pedicels spreading or divergent, glabrous : flowers blue : lower
sepals lanceolate, 12-15 n^"^- ^o^g ; lateral ones very oblique, as
broad as long, about i 5 mm. each way, rounded, slightly reniform
on the upper side ; hood about 2 cm. long, deeply saccate, elong-
ated helmet-shaped ; beak long-attenuate, very porrect or even
ascending.

The type specimen was included in A. Cobtvibiamnn by Dr.
Gray, his " Syn. Fl. N. Amer." label being on the sheet ; but it
is so unlike all other material of that species that I can not see
the reason why it should be included therein. The perfectly
glabrous stem, the branched inflorescence, the peculiar, deeply
saccate hood and the slender porrect beak are characters not found
in any other American aconite.

Southern Utah, Northern Arizona: 1877, ^''- ^- Palfnc^',
II ( type in herb. Columbia University).

Anemone tuberosa

Anemone sphenopJiylla Britton, Ann. N. Y. Acad. 6 : 224, in
part. Not Poepp. 1827-29.

Perennial with a thick tuberous root, basal leaves with petioles

152 Rydberg : Studies on the

about 5 cm. long, twice ternate ; divisions rhombic-cuneate in out-
line, ternately cleft and toothed ; teeth oblong-ovate : scape 1-3
dm. high, perfectly glabrous below the involucre ; peduncles finely
appressed silky ; involucral leaves similar to the basal ones, but
short-petioled and with longer lobes and teeth ; peduncle solitary
or a second one with a smaller involucre : sepals white or purplish,
8-10, oblong-linear, 1-2 cm. long, finely silky : head of fruit
ellipsoid, about i cm. thick and 2 cm. long : achenes densely
woolly: style filiform, about 1.5 mm. long.

This is closely related to A. splicnophylla of Chili, but in that
species the whole scape is decidedly pubescent with spreading
hairs and the segments of the involucral leaves are narrower. The
following species belong here :

Arizona: Sierra Tuscon, 1884, C. G. Pri ngl e {ty^Q in herb.
Columbia University) ; Fort Huachuca, 1892, 5. E. Wilcox.

New Mexico: Mountain near Las Cruces, 1895, E. 0.
Wooton.

Utah : St. George, 1880, M. E. Jones, i6oj.

California: Panamint Mountains, 1891, Coville & Funston,
500 ; 1849 (locality not given), Fre^noiit.

Anemone lithophila

Perennial with a short thick rootstock : basal leaves several,
glabrous or nearly so, thickish, shining, somewhat glaucous ;
petioles 5-8 cm. long ; blades ternate ; divisions obovate-cuneate,
about 3 cm. long, strongly veined, deeply 3-cleft, again cleft and
toothed ; the ultimate segments short, oblong-oblanceolate : scape
1-2 dm. high, sparingly pubescent with long silky hairs : in-
volucral leaves subsessile or short-petioled ; divisions similar to
those of the basal leaves : sepals silky, ochroleucous, tinged with
blue, 12-15 mm. long, broadly obovate or oval : achenes densely
villous all over : style filiform, about 2 mm. long.

The type was growing with A. parviflora at an altitude of 1800
m. As none of the specimens there had well-developed fruit and
the plant is intermediate between A. parviflora and A. glohosa
Nutt., which also grew in the neighborhood, this species may be
a hybrid. The strong veins and shiny glaucous leaves suggest
the former but the general leaf-form the latter, although the
segments are broader and shorter and the leaves more glabrate.
Watson's specimen, which resembles the type perfectly has fully
developed fruit, however. The type was labelled A. Tetonensis,

Rocky Mountain Flora 153

which it resembles most in habit ; but the achenes are wooly all
over, not merely strigose on the back as in that species and the
flowers are larger and lighter. The following specimens are to be
referred here :

Montana : Little Belt Mountains, nine miles from Barker,
1896, /. H. Flodnian, ^jg (type in herb. N. Y. Bot. Garden),
and 46J ; Lake Stanton, 1894, R. S. Williains.

Utah: Uinta Mts., 1869, i\ Watson, 10.

Anemone Piperi Britton

Perennial with a rather slender rootstock : stem 1.5—3 ^"^•
high, slender, very sparingly appressed silky ; basal leaves ternate,
minutely appressed-puberulent when young, in age glabrous ; peti-
oles 1-2 dm. long ; middle lobe rhombic-obovate or rhombic-cune-
ate, coarsely toothed above the middle, 2-7 cm. long ; the lateral
ones broader, obliquely ovate, 2-cleft to about the middle, coarsely
toothed : involucral leaves similar, but the lobes usually narrower :
petioles 1-3 mm. long : pedicel 2-5 cm. long, erect in fruit, spar-
ingly appressed silky : sepals elliptic-obovate to oval, about 1 5
mm. long, 6-8 mm. wide, white, glabrous : achenes about 4 mm.
long, densely short-pubescent, ellipsoid, slightly compressed, taper-
ing to both ends : beak very short.

This has been included in A. qiiinquefolia and all specimens
cited for that species from the Northwest may belong here. It
differs, however, from that species in the form of the basal leaves,
the erect pedicels and the short, almost straight beaks of the
achenes.

Idaho : Latah county, 1893, C. V. Piper, r^6g (type in herb.
Columbia University); Craig Mountain, 1892, Saudbcrg, Mac-
Doiigal & Heller, ig^.

Washington : Kamiac Butte, 1896, A. D. E. Elmer, jgj.

Clematis Jonesii (Kuntze)

Clematis Douglasii var. Jonesii Kuntze, Verh. Bot. Ver. Bran-
denburg, 26 : 180. 1886.

Perennial with a thick rootstock or caudex : stems 3-6 dm.
high, simple or later in the season branched, more or less woolly
when young: leaves twice pinnately divided, i— 1.5 dm. long, vil-
lous when young, in age glabrate, rather firm, segments lanceolate
to linear-lanceo'ate, often cleft, 1-5 cm. long, acutish : flowers nod-

154 RvDBERG : Studies on the

ding : calyx campanulate, brown, more or less villous, especially
near the margins; sepals 2-2.5 cm. long, ovate, acute or acumi-
nate, upper half with a wavy dilated margin : achenes about 5 mm.
long and 4 mm. wide, flattish, densely silky; tails of the achenes
4—5 cm. long, beautifully plumose throughout.

This is nearest related to C. Dovglasii, with which it has been
confused. It differs from that species in the dilated margins of
the sepals, the distinctly petioled and less compound leaves. In
the true C. Doiiglasii the upper and middle leaves are twice pin-
nately divided and subsessile, so that they with the first pair of
primary divisions look as if verticillate. Kuntze's description of
C. Doiiglasii var. Jonesii is so meager that nobody could know
from it what he meant, but fortunately we have one of Jones'
specimens. The following specimens belong to C. Jonesii.

Colorado: Howe's Gulch, 1899, W. F. M.; Dolores (7300
ft.), 1892, Crandall ; lat. 39°-4i°, 1862, Hall & Harbour, 2;
Howe's Gulch, 1893, C. F. Baker; near Boulder, 1892, H. M.
Patterson, 168 ; Dixon Canon, 1891,^. H. Cozuen, j68.

Utah : Uinta Mountains, 1869, 5. Watson, i ; American Fork,
1880, M. E.Jones, 135 1.

Wyoming : Headwater of Tongue River, Big Horn Mountains,
1898, Frank Tweedy, lyi.

Clematis eriophora

Perennial, from a woody caudex : stems and leaves prominently
white-villous, the former 3-5 dm. high, simple : leaves 5-10
cm. long, distinctly petioled, twice pinnately divided ; ultimate
segments narrowly linear, 1—3 cm. long, 1—2 mm. wide: flowers
nodding ; calyx villous, campanulate, about 3 cm. long ; sepals
oblong, obtuse, the upper third spreading, with a dilated margin :
achenes oblong, about 6 mm. long and 3 mm. wide, silky, with a
blunt ridge on each side ; tails about 4 cm., beautifully plumose.

This is closely related to the preceding and to C. Bakeri, but
differs from the former in the narrower leaf-segments, the obtuse
and thicker sepals and the denser and more persistent pubescence,
and from the latter in the longer leaf segments and the obtuse
dilated sepals. It grows at an altitude of 1500—2000 m.

Colorado: Vicinity of Horsetooth, 1896,/. H. Coiven (type
in herb. N. Y. Bot. Garden ; cotypes in herb. State Agric. College,
Colo.); Foothills, Larimer county, 1893, C. S. Crandall ; Colorado,

RocKv Mountain Flora 155

1870, Dr. G. W. Hulsc ; 1844, Fremont, 36 j ; Pagosa, 1883,
Brandegee ; Golden City, 1871, E. L. Greene ; Clear Creek, 1873,
John Wolf, g2 ; Colorado Springs, \(^oo, Rydberg & JVeeland,
62J2 ; Rist Canon, 1890, Crandall, 2^2 ; 1893, ijyg ; Horse-
tooth Gulch, 1898, Crandall.

Clematis Wyethii Nutt. Journ. Acad. Phila. 7: 6 1834

I think that this species is distinct from C. Doi/glasii. Hook., the
leaves being never twice pinnately divided, having much broader
segments and never being falsely verticillate as in that species. The
specimens from Beaver Cafion, Idaho, cited in the synoptical flora
under C. Doiiglasii var. Scottii and in Howell's flora of the North-
west under C. Scottii belongs undoubtedly here. A broad-leaved
C. Wyethii and a narrow-leaved C. Scottii resemble each other very
much and can be distinguished only by the flower, which in the
former is campanulate with oblong-lanceolate sepals and in the
latter rounded urn-shaped with broadly ovate sepals. C. Wyethii
is common in Montana and northern Idaho. It is represented by
the specimens cited under C. Scottii and some under C. Douglasii
in my catalogue of the flora of Montana.

Atragene occiDENTALis Homem. Hort. Hafn. 1813 : 520

Atragene Cohunbiana Nutt. Journ. Acad. 7:7. 1834 ; Clematis
CohimbianaToYv. & Gray, Fl. N. Am. i : 10. 1838 ; Clematis vcr-
ticillaris var. Cohunbiana Gray. Syn. Fl. 1:8. 1895 ; Clematis
Pseudoatragene a. normalis Kuntze, Verh. Bot. Ver. Brandenburg,
26 : 160, in part, 1884 ; C. alpina a occidentalis 3 typica Kuntze,
I. c. 161.

Dr. Gray adopted the name occidentalis as a varietal name for
another species, and cites Homeman's plant as a synonym. It is
very evident that the latter is the common plant of the Columbia
Valley with simply ternate leaves and subentire leaflets, for Home-
man expressedly describes them as entire. Dr. Kuntze was so
far right that he placed Atragene occidentalis Homeman with A.
verticillaj-is, although he made both forms of Clematis alpina. His
revision of Clematis is far from good especially as to the West
American species. The only distmction he makes between C.
pseudoatragene and C. alpina is the absence or presence of stami-

156 Rydberg : Studies on the

nodia, a very unfortunate distinction, for in most of the West
American forms both conditions exist. The filaments of the outer
stamens are flattened and more or less petaloid. In some cases
the outermost have no anthers (true staminodia), but often all are
antheriferous. The present species is therefore placed in both
species by Kuntze. Atragene occidcntalis differs, however, from the
eastern A. Americana in the fact that the staminodia as far as I
knovv always are linear, while they are in A. Americana decidedly
spatulate. A. occidcntalis is common from British Columbia and
the Canadian Rockies south to California and Colorado.

Atragene grosseserrata

Trailing or climbing over bushes : leaves ternate ; petioles
5-8 cm. long; petiolules 1-2 cm. long; blades broadly ovate,
oblique, somewhat cordate at the base, 3-4 cm. long, coarsely
toothed and often somewhat lobed : peduncles 5-10 cm. long:
sepals lanceolate, about 4 cm. long, 10-13 mm. wide, acute : stami-
nodia about 18 mm. long, decidedly spatulate, a little exceeding
the stamens : achenes small, sparingly hirsute : tails about 3 cm.
long, plumose throughout.

This is closest related to the eastern A. Americana; but the leaves
are deeper serrate and the sepals are longer, lanceolate, and resem-
ble more those of A. teniiiloba. From A. occidcntalis it differs in
the spatulate staminodia, the shorter leaflets, which are coarsely
serrate except the very base. In A. occidcntalis the leaflets are en-
tire or merely crenate above the middle.

Idaho : Palouse county and about Lake Coeur d'Alene, 1892,
G. B. Alton (type in Herb. N. Y. Bot. Garden, flower).

Washington : Cascade Mountains, 1882, F. Tweedy (fruit).

Atragene repens (Kuntze)

Clematis alpiiia a occidcntalis 2 rcpcns Kuntze, Verh. Bot.
Ver. Brandenberg, 27, 161, in part. 1884.

Plant trailing : leaves biternate, glabrous, firmer than in the
two preceding : petioles 5-8 cm. long : secondary leaflets sub-
sessile except the terminal, which is short-stalked, broadly ovate,
coarsely toothed with broadly ovate teeth, 3-4 cm. long, abruptly
short acuminate at the apex : peduncles over i dm. long : sepals
ovate-lanceolate, about 4 cm. long, acute : staminodia linear :
achenes about 5 mm. long, pubescent : tails about 4 cm. long,
plumose throughout.

RocKV Mountain Flora 157

Dr. Kuntze's description of the subvariety repens is just as
meaningless as the rest of his descriptions. He cites specimens,
however, v\z., Jones no. 1SS2 and iSSy. We have two sheets of
each and they evidently do not belong together. No. 1882 in
both the Columbia University and New York Botanical Garden
herbaria are in fruit, have simply ternate leaves with subentire
leaflets and must be referred to A. occidcntalis. No. 1887 in
flowers, on the contrary, has biternate leaves, and the leaflets are
coarsely toothed. Dr. Kuntze's subvariety is, therefore, " eine
zusammengesetzte " and according to some Germans, should be
ruled out. After the part that belongs to A. occidcntalis has been
removed, I have adopted the name for the residue.

Utah: American Fork, 1880, M. E. Jones, iSSj (type in
herb. N. Y. Bot. Gard.).

Wyoming: Wolf Creek Caiion, 1896, Aven AWson, 22()^, in
part, the larger part belonging to A. tomiloba.

Atragene pseudoalpina (Kuntze)

Atragene Ochotensis Gray, PI. Fendl. 4. 1849. ^^t Pall.
1874; A. alpina var. Ochotensis A. Gray, Am. Jour. Sci. II. 33 :
241 ; Clematis alpina var. Ochotensis S. Wats. Bot. King Exp. 5 :
4 ; C. alpina var. occidcntalis A. Gray; Powell, Geol. Surv. Dakota,
531. 1880. ^o\. Atragene occidcntalis Wova^Ta. 181 3; Clematis
pseiidoatragene y psendoalpina Kuntze, Verb. Bot. Ver. Branden-
burg, 26 : 160. 1884.

As stated above, the name occidcntalis should be used for A.
Colnmbiana Nutt. We are therefore obliged to adopt Kuntze's
varietal name. In the Synoptical Flora Clematis pseiidoatragene
Kuntze is given as a synonym, but from the short description
neither of the varieties «, ;9 or 0 can belong here. The var. a
nonnalis is evidently partly A. Americana and partly A. occidcn-
talis as here understood. The var. o subtriternata is A. tenniloba
(A. Gray) Britton. What Clematis pseiidoatragene var. /9 Wender-
othioides* is I do not know. In the Synoptical Flora the range
is given as extending from New Mexico to Dakota and Wash-
ington. I have not seen any specimens from any station north of

* What a name ! Translated it would mean : A variety of the false Atragene
Virgin's Bower, resembling Mr. Wenderoth.

158 Rydberg : Studies on the

Colorado and Utah. All specimens referred here from higher lati-
tudes belong to A. tcmiiloba.

Ranunculus Utahensis

Perennial with a cluster of fibrous fleshy roots, glabrous : stems
1-2 dm. high, branched above; basal leaves petioled ; petioles 2-
5 cm. long ; blades almost orbicular, rather thick, crenate or round-
lobed : stem-leaves sessile, deeply cleft or divided into 4-5 oblong
or elliptic, obtuse segments : sepals somewhat hairy, elliptic to
obovate, at last reflexed : petals 4-5 mm. long, exceeding the
sepals: headof fruit oblong ; achenes glabrous ; style rather short.

This species resembles most a depauperate R. abortknis in the
leaves, but the petals are much larger and the head of fruit is ob-
long instead of spherical. It has gone under the name of R. affinis
var. leiocarpns, but it is not that of Trautvetter, and is distinguished
from all of those species which have been included in R. affinis by
its thicker leaves and their lobing.

Utah: Alta, Wasatch Mts., 1879, M. E. Jones, 11 jo (type in
herb. Columbia University); Uinta Mts., 1873, T. C. Porter.

Ranunculus micropetalus (Greene)
Ranunculus affinis \'ar. micropetalus Greene, Pittonia, 2 : 1 10.
1890.

An excellent species, very distinct from R. affinis Hook., and still
more from R. cardwpliyllus Hook., with which Davis associated it.

Ranunculus Helleri

Perennial with a short rootstock, perfectly glabrous : stem
slender, 1—2 dm. high : basal leaves and lower cauline with slen-
der petioles 2-5 cm. long ; blades of the earlier basal leaves reni-
form, 3-cleft 7^ or 3^ the distance to the base ; middle division
oblong, entire, the lateral ones with 3 ovate lobes ; blades of the
other basal leaves and the lower cauline divided to near the "base
into 3 cuneate divisions, 15-20 mm. long; the middle division 3-
lobed, the lateral ones 4-lobed ; stipules of lower cauline leaves
conspicuous, rounded : upper cauline leaves sessile, divided unto
the base into 2—4 linear-oblong divisions : flowers solitary : sepals
oblong, obtuse, about 5 mm. long : petals oblong to obovate,
golden yellow, 6-7 mm. long : head of fruit short-oblong or
spherical : achenes turgid, almost spherical, 1.5 mm. long, gla-
brous : style slender and curved.

This species is nearest related \.o' R. EschscJiotzii and R. alpco-

RocKv Mountain Flora 159

philns. In the former, which is very rare in the Rockies, the
upper stem-leaves have broad and short lobes, which are elliptical
to obovate ; and in the latter the basal leaves are not reniform but
almost orbicular in outline. A'. //it'/Zr/v grows in bogs at high altitudes.

Idaho : Near Lake Pend d'Oreille, 1892, Sandbtrg, MacDougal
& Heller, 842 (type in herb. N. Y. Bot. Card.).

Wyoming : Big Horn Mountain, 1899, F. Tiveedy, 2j8^.

Montana: Sperry Glacier, 1901, F. K. Vreelami, mo.

Ranunculus stenolobus

Perennial with a cluster of fibrous roots, perfectly glabrous :
stems 1-3 from the same clump, 1—2 dm. high, usually branched :
basal leaves and lower cauline petioled ; petioles 4— 10 cm. long ;
blades three times ternately dissected into almost linear segments,
which are 6—15 mm. long and 1—3 mm. wide, usually somewhat
narrowed at the base ; the primary divisions distinctly pctiolulate :
sepals oval, 4-5 mm. long : petals obovate, 7-8 (rarely 9-1 1)
mm. long : head of achene spherical or slightly oblong, about 6
mm. broad : achenes turgid, glabrous : style slender.

This species is somewhat intermediate between R. adoneus and
R. triternatus, but the stem is more elongated and branched.
Some of the specimens cited below have been referred to the
former, but in that plant the stem is usually simple, the petals
over I cm. long, cuneate-flabelliform and overlapping each other,
the lobes of the leaves narrower and not narrowed downward. In
the leaf-form and flowers it resembles more R. triternatus, but that
grows in big clumps with numerous stems, and the segments are
more decidedly petiolulate.

Wyoming : Headwaters of Cliff Creek, 1900, C. C. Curtis
(type in herb. N. Y. Bot. Card.).

Utah: Alta, Wahsatch Mts., 1879, ^^- ^- Jones, 1104.;
Wahsatch Mt., 1869, 5. Watson, ^o.

Papaver pygmaeum

A cespitose and scapose perennial ; leaves all basal, numerous
2—3 cm. long ; blade less than i cm. long, broadly ovate in out-
line, deeply cleft into rounded-oblong or obovate segments, spar-
ingly hispid or glabrous : scape 4-6 cm. high, sparingly hirsute ;
so also the calyx, which is obovate in bud : petals yellow, i cm. or
less long : pod obovoid, about i cm. long, densely bristly.

This species is nearly related to P. radicatum Rottb., but is a
still smaller plant, with shorter, broader, less divided, less bristly

160 RvDBERG : Studies on the Rocky Mountain Flora

leaf-blades and smaller flowers. It resembles still more the Euro-
pean P. Pyrenaiciun, which has larger petals, 1—2 cm. long, sta-
mens much exceeding the ovary and spherical flower-buds.

Montana : Mountain above Stanton Lake, 1894, R. S. Williams,
pp2 (type in herb. N. Y. Bot. Garden).

Alberta: Sheep Mountain, i8g^, /oli;i Maco?f;i, io26p ; top
of Rocky Mountains, 1881, G. M. Dazvson.

Argemone rotundata

Stem stout, densely and strongly bristly, but otherwise gla-
brous : leaves oval in outline ; the cauline ones sessile and broadly
auricled, bristly, especially on the veins and margins, round-lobed :
flowers short-pedicelled or subsessile : calyx very bristly ; its horns
erect or slightly spreading, bristly : petals white, fully 4 cm. long :
pod ovoid, very bristly.

This has been mistaken for A. liispida Gray but is perhaps
nearer related to A. intenncdia and A. platyceras. From A. hispida
it is easily distinguished by the absence of a finer indumentum and
by the round-lobed leaves. The latter character also distinguishes
it from the other two species mentioned. It is also much more
bristly than the two. It grows at an altitude of 1500— 1800 m.

Nevada : Diamond Mountains, July, 1868, 5. Watson, ^7
(type in herb. Columbia University).

Utah : Utah Valley, i86g, S. Watson, ^.g.

BicucuUa occidentalis

Perennial with a very short rootstock bearing numerous tubers :
scape and leaves perfectly glabrous; petioles of about 1.5 dm.
long ; blades twice ternate ; the divisions twice pinnately divided
or cleft into linear-oblong lobes, 1—2 cm. long, 3-5 mm. wide :
scape about 3 dm. high : raceme simple : sepals oval, about 5 mm.
long : petals pink ; the outer about i cm. long ; their spurs about
12 mm., divergent, forming with each other an angle of 90° or
more ; crest of the inner petals prominent ; capsule fusiform, with
the style about 1.5 cm. long.

This is nearest related to B. cucnllaria, and has been confused
with it, but has coarser foliage, more diverging and longer spurs,
more prominent crest on the inner petals and the underground parts
more gruinose and not scaly. Its range includes parts of Oregon,
Washington and eastern Idaho, and it grows on shaded hillsides.

Washington: West Klickitat county, 1892, ]]\ A\ Suksdorf
(type in herb. Columbia University).

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 23

STUDIES OiN THE ROCKY MOUNTAIN FLORA

YIII.

BY PER AXEL RYDBERG.

NEW YORK
1902

[Reprinted from Bulletin Toreey Botanical Clue, 29 : 2S3-2i6, April 24, 1902.]

Studies on the Rocky Mountain Flora- Vlll

By Per Axkl Rydberg.

Stanleya arcuata sp. nov.
Perennial with a ligneous caudex, 4—10 dm. high, perfectly
glabrous ; stem light green and shining, in age straw-colored : leaves
entire or the lower with short and broad lobes, lanceolate, long-
petioled, thick, 5-12 cm. long ; pedicels rather short, in fruit, about
I cm. long : sepals yellow, linear about i 5 mm. long ; petals yel-
low ; blades 7-8 mm. long, broadly elliptic, almost as long as or at
least two thirds as long as the claw : pods 7-10 cm. long, strongly
arcuate, and spreading.

This species is nearest related to 5. integrifolia, but is taller^
more shrubby, with narrower leaves and broader petals. .S. pi}i-
nata (Pursh) Rritton, with which both have been confused, has.
hairy leaves, most of which are deeply pinnatifid, and the blades
of its petals are linear oblong and only one third or one half as
long as the claws. vS. arcuata grows in dry regions from Wyoming
to Nevada and south to Colorado and Arizona ; probably also in
California.

Nevada : Unionville Valley, 1868, S. Watson, io§ (type in
herb. Columbia University).

Stanleya canescens sp. nov.

Stout perennial .5-1 m. high; stem light green, finely puberu-
lent or glabrate, terete : leaves grayish puberulent, the upper entire
and oblanceolate, 5-7 cm. long; the lower pinnatifid with oblong
lateral lobes and a large oval or elliptic end-lobe : raceme dense,
pedicels short, in fruit i cm. or less long, divergent : sepals linear,,
about 1 2 mm. long ; blades of the yellow petals linear-oblong, about
half as long as the claw : pods arcuate, spreading, 5-7 cm. long.

This is nearest related to the preceding, differing in the puberu-
lent leaves, which are shorter and broader and more inclined to be
pinnatifid.

Utah : Frisco, 1880, M. E. Jones, i8og (type in herb. N. Y.
Bot. Garden).

Arizona: Fort Verde, 1887, E. A. Mearns ; Verde River,
1883, H. H. Rushy ; Mexican Boundary Survey.

232

233 Rydberg : Studies on the Rocky Mountain Flora

Thelypodium macropetalum sp. nov.

A glabrous perennial, 4-6 dm. high : basal leaves oblanceolate
or spatulate, short-petioled, 2—5 cm. long : lower stem-leaves ob-
long, obtuse, sessile, with rounded auricles at the base, 5-6 cm.
long ; upper leaves lanceolate-sagittate, acute ; pedicels in fruit
about I cm. long, divaricate : sepals about 5 mm, long, purplish
with white margins ; petals purple, spatulate, long-clawed, nearly
three times as long as the sepals : pods 4—5 cm. long, ascending
or erect and somewhat incurved.

This is related to T. sagittat2un (Nutt.) Heller and T. paniada-
tuvi A. Nelson. From the former, it differs in the smaller leaves,
the rounder auricles, the erect, not reflexed nor spreading pod ; and
from the latter by the much longer petals. It grows at an altitude
of 1,000-1,200 m.

Utah: Farmington, 188 1, J'/ E. Jones, 18^1 (type in herb.
N. Y. Bot. Garden).

Idaho: Soda Springs, 1892, Isabel Miilford.

Thelypodium Utahense sp. nov.

Annual or biennial : stem 4-6 dm. high, sparingly pubescent
below; leaves all lanceolate in outline, 5-10 cm. long, lanceolate,
glabrous ; segments broad and crowded, oblong-lanceolate to tri-
angular : pedicels very short, in fruit scarcely more than i mm.
long : sepals oblong, about 2 mm. long : petals spatulate, almost
twice as long, white : pod about 3 cm. long, strongly reflexed.

This is probably nearest related to 5. reflexuni Nutt., which is
decidedly hispid on the stem and the veins of the leaves and has
much narrower leaf-segments. It grows at an altitude of about
600 m.

Utah : St. George, 1880, M. E. Jones (type in herb. N. Y.
Bot. Garden); 1877, E. Palmer, 28.

Lepidium Jonesii sp. nov.

Lepidmuimontaiinniv-A.x. alf ssoides ] ones, Zoe, 4: 266. 1893.
Not L. alyssoides A. Gray. 1849.

Perennial with a short woody caudex : stems 2-3 dm. high,
glabrous, branched, glabrous : basal leaves pinnatifid with linear
acute divisions ; stem-leaves similar or the upper entire, narrowly
linear : racemes rather short and dense ; pedicels in fruit 6-8 mm.
long, spreading : sepals oblanceolate, obtuse ; petals white, much

Rydberg : Studies on the Rocky Mountain Flora 234

exceeding the sepals ; blade broadly obovate : pod acutish, oval in
outline, about 4 mm. long and slightly over 2 mm. wide, glabrous,
reticulate, only slightly wing-margined above, retuse; st}'le about
twice as long as the width of the wing margins.

This species is in some respects intermediate between L. alys-
soidcs and L. montamun. In the former only the lowest leaves are
pinnatifid, the pod is more rounded and the style is shorter, scarcely
longer than the width of the wing-margin. It is closer related to
L. montajium, differing mainly in the longer and narrower leaf-
segments, having a more persistent base and being less pubescent.

Utah: St. George, 1880, M. E. Jones, i6j6 (type in herb.
N. Y. Bot. Garden); Price, 1900, vS". G. Stokes ; southern Utah,
1874, C. C. Parry, 18.

Nevada: Trinity Mountains, 1868, 5. Watson, 120.

Lepidium elongatum sp. nov.

Annual : stem branched near the base, about 3 dm. high,
glandular pruinose above : basal leaves spatulate or oblanceolate,
about 4 cm. long, coarsely toothed or somewhat pinnatifid with
short lobes ; stem leaves oblanceolate, tapering into a short petiole,
entire or with a few small teeth, 3-4 cm. long, acutish : racemes
elongated, at the ends of the branches, in fruit i dm. or more
long : sepals oblong or oblanceolate, acute, thin and whitish ; petals
minute, linear-oblanceolate, one half to two thirds as long as the
sepals or lacking : pods nearly orbicular, about 3 mm. broad, usually
broadest a little above the middle, wing-margined and deeply retuse
at the apex, glabrous or slightly pruinose : stigma sessile.

This species is nearest related to L. apetahini and L. ramosis-
siuiuin. From the former it differs in the branching near the base,
the more entire leaves, the somewhat larger pod and the petals
which are generally present ; and from the latter in the longer
racemes, in the lack of the small axillary racemes characteristic to
that species and a different pod. In L. ramosissinmvi this is
broadest below the middle. L. clongatinn grows on rocky hilltops.

Washington: Almata, 1896, A. D. E. Elmer, 21 (type in
herb. N. Y. Bot. Card.).

Idaho: About Lewiston, 1896, A. A. & E. Gertrude Heller,
3008.

235 Rydberg : Studies on the Rocky Mounjain Flora

Thlaspi Nuttallii

TJilaspi cocJilcariforuie Nutt. Journ. Acad. Phila. 7 : 13. 1834.
Not DC. 1 82 1.

This species has been merged in T. alpestre, but none of our
American forms belong to that species. T. Nuttallii is inter-
mediate between T. payvifloruin A. Nelson and T. glauciivi A.
Nelson, having larger flowers than the former and smaller than
the latter. The pod is obovate, rounded at the apex with a narrow
sinus. In T. glaucmn the sinus is broad and open.

Roripa clavata sp. nov.

Biennial, tall, perfectly glabrous : stem 6-IO dm. high, simple
up to the inflorescence: leaves lyrate-pinnatifid, 5-10 cm. long,
more or less auricled at the base ; lobes 4-8 pairs, oblong to ovate,
obtuse or acute, broadly dentate : inflorescence branched ; petioles
in fruit 5-10 mm. long, divaricate: petals spatulate, about 3 mm.
long, exceeding the sepals : pod 10-12 mm. long, 2.5-3 mm. thick,
decidedly clavate, obtuse, strongly curved ; style nearly i mm. long.

This may have been included in Watson's Nasturtium terrestre
var. occidentale ; but the name occidcntale cannot be used as there
is already another older Roripa occidcutalis Greene. It is not,
however, Roripa Pacifica Howell, which is supposed to be a syno-
nym of Watson's variety, for that species is described as being
slightly pubescent and having pods which are acute at both ends,
while in R. clavata the pod is almost truncate at the apex.

Washington: Hogeman, Chehalis county, 1897, Frank H.
Lamb, 122 1 (type in herb. N. Y. Bot. Gard.) ; West Klickitat
county, 1885, W. M. Suksdorf.

Oregon : Sauvie's Island, 1888, T. Hoivell, ijjj.

British Columbia : Port Henry, i88g,/o/m Macoun.

Idaho: Near Hope, Kootenai county, 1892, Sandberg, Mac-
Dougal & Heller, 102^, at least in part.

Roripa Underwoodii sp. nov.

Biennial, divaricately branched, even near the ground, perfectly
glabrous ; stems 2 dm. or more high : leaves 3-5 cm. long, lyrately
pinnatifid, thin ; lobes oblong, obtuse ; the terminal one elliptic
to broadly oval, sinuately toothed ; petioles short, winged and

Rydberg : Studies on the Rocky Mountain Flora 236

conspicuously auricled at the base : racemes numerous : pedicels
short, in fruit scarcely more than 4 mm. long, diverging or even
reflexed : sepals bright orange, obtuse ; petals light yellow,
scarcely equalling the sepals and not quite i mm. long : fruit (im-
mature) 5-6 mm. long, slightly thicker at the base, abruptly con-
tracted into a minute style, not .5 mm. long.

In general appearance, this species resembles most R. pahistris,
but it is more branched with divaricate branches, has much shorter
pedicels, rather longer and more tapering pod and not half as long
style. From R. obtusa, it differs in the shape of the leaves, which
are less dissected, the shorter style and general habit. The type
grew at an altitude of 3300 m.

Colorado: Red Mountain, south of Ouray, 1901, L. M. Un-
derwood, 2gga.

Roripa Integra sp. nov.

Biennial, glabrous : stems few, strict, branched above ; basal
leaves not seen as they are fallen off in the type specimen ; stem-
leaves spatulate or rhombic spatulate, 1-2 cm. long, not auricled
at the base, entire or slightly sinuate : racemes slender, lax : petals
1.5 mm. long, scarcely exceeding the sepals; pedicels 1-3 mm.
long, ascending : pod about 8 mm. long, somewhat curved : style
about 5 mm. long.

The type specimen is labelled Nasttirtmm pahistre var., but it
is not closely related to this species. It is nearer R. obtusa, but
differs in the entire leaves ; the more tapering and more curved
pod, the erect habit and the short style. It grows at an altitude
of about 2700 m.

Utah: Wasatch Mountains, 1869, 5. Watson, 64, in part
(type in herb. Columbia University).

Lesquerella arenosa (Richards).

Vesicaria arenosa Richardson, Frankl. Journ. App. 743.

Lesqncrella Lndoviciana var. arenosa S. Wats. Proc. Am. Acad.
23: 252.

This is amply distinct from L. Lndoviciana (DC.) S. Wats, or
L. argentca (Pursh) MacM., by the more slender ascending stems,
the shorter leaves, the oblanceolate, instead of linear stem-leaves
and the comparatively larger and often somewhat ellipsoid pod.
The range given in the Synoptical Flora should be extended to
the Black Hills of South Dakota and northeastern Wyoming.

237 Rydberg : Studies on the Rocky Mountain Flora

Lesquerella Shearis sp. nov.

Perennial with a large rosette of basal leaves ; stems several,
ascending, 1-2 dm. high, densely stellate : basal leaves 3-4 cm.
long ; blades broadly oval or ovate, entire or slightly angular den-
ticulate ; stem leaves oblanceolate, about 2 cm. long, subsessile :
raceme many-flowered ; pedicels in fruit bent S-shaped : pods erect,
densely stellate, 7-8 mm. long, acute, compressed above ; septum
elliptic in outline : style about as long as the pod.

This species differs from L. curvipes A. Nelson in the broad
basal leaves, the elliptic instead of ovate outline of the septum and
the longer style. It grows along creek banks at an altitude of
about 2000 m.

Colorado: Idaho Springs, 1895, C. L. Shear, 326^ (type in
herb. N. Y. Bot. Gard.); also at the same locality, P. A. Rydberg.

Physaria brassicoides sp. nov.

A tufted perennial with a large and dense rosette of basal leaves
and a thick taproot : basal leaves large, with winged petioles ;
blades almost orbicular in outline, sinuate-dentate, 2.5-5 cm. in
diameter : stem leaves small and few, spatulate : flowering stems
about I dm. high, ascending : flowers numerous : fruit almost
obcordate, acute at the base, deeply divided above ; cells inflated,
rounded pyriform, 8- 10 mm. long and 5-8 mm. in diameter: style
about 5 mm. long.

This species resembles most P. didymocarpa in habit, but differs
in the smaller fruit, which is not at all cordate at the base but on
the contrary usually acute, and in the larger basal leaves which
form a large rosette, 7-10 cm. in diameter and having some resem-
blance to a young cabbage-head. It grows in crevices of magnesian
rocks in canons and badlands.

Nebraska : Canon south of Scott's Bluffs, 1891, Rydberg, 24.

Cardamine acuminata (Nutt.)

Cardaminc Idrsiita [i acuminata Nutt.; T. & G. Fl. N. Am. i :
85. 1838.

This species has been variously referred to C. Jdrsiita, C. obhgo-
sperma, and C. Pennsylvanica. It is probably nearest related to the
last, but the lower part of the stem is hairy and the pod has an
evident style over i mm. long. It ranges from the Mackenzie
River to British Columbia, south to California and Wyoming.

RvDBERG : Studies on the Rocky Mountain Flora 238

Cardamine multifolia sp. nov.

Stem about 2 dm. high, very leafy, lower part hirsute : leaves
4-6 cm. long glabrous; leaflets 7-13. oblong to oblong-obovate
or oval, entire or sinuate-dentate ; the terminal one larger and
obovate : flowers numerous : petals spatulate, about 2.5 mm. long ;
pedicels ascending, about 8 mm. long : pod erect, about 2 cm.
long and .75 mm. wide.

This is perhaps closest related to A. parvifolia L., but is more
leafy, has shorter and broader leaf-segments and erect pods, and
the plant is less diffusely branched.

Idaho: Beaver Caiion, 1895, P. A. Rydberg.

Utah : Wasatch Mts., 1869, .9. Watson, 80, in part.

Sophia brevipes (Nutt.)

Sisyinbrmm canescois c {brevipes Nutt. MSS.) Torr. & Gray,
Fl. N. Am. I : 92. 1838.

Sisymbrium Hartivegianiim Fourn. Rech. Auct. & Tax. Fam.
Cruc. & Gen. Sisymb. 66, in part. 1865.

Sisyinbritim incisum var. Hartwegiamim S. Wats. Bot. Calif, i :
41.

The plant described by Dr. Watson in the Botany of California
and by Dr. Robinson in the Synoptical Flora (p. 139) is I think
without doubt the var. brevipes of Torrey and Gray's flora. Prob-
ably also Bourgeau's plant, cited by Fournier, belongs here ; but I
think that it is very doubtful that Hartweg's plant, which must be
regarded as the type of 5. Harttvegianiim Fourn., could be the same.

Sophia Calif ornica (T. & G.)

Sisymbrium canescens e (Ca/iforuieum) T. & G. Fl. N. Am. i :
92. 1838.

This species is characterized by the large flowers, the sparingly
stellate and often glandular-puberulent stem, the uniserial linear
pod and once or twice pinnatifid leaves. The segments of the
lower leaves are broad and obtuse, those of the upper linear to
oblong. Its range extends from Montana to Washington south to
California.

Sophia viscosa sp. nov.

Biennial, 5-10 dm. high : stem, at least the upper part, decidedly
glandular or viscid-villous : leaves obovate or oblanceolate in out-

239 Rydberg : Studies on the Rocky Mountain Flora

line, 3-10 cm. long, twice pinnatifid, densely pubescent; primary
divisions of the lower leaves ovate and obtuse ; those of the upper
lanceolate and acute ; secondary divisions oblong acute or obtuse,
often few-toothed : inflorescence usually much branched ; pedicels
in fruit 8-12 mm. long : sepals yellowish, 1-15 mm. long, oblong ;
petals bright yellow, spatulate, about 2 mm. long : pod (in the
specimens seen not fully developed) over i cm. long and less than
I mm. wide : seeds uniserial.

This is perhaps nearest related to 5. incisa (Engelm.) Greene ;
but differs in the dense viscid pubescence of the stem, the pubescent
leaves and deeper divided primary segments. It grows among
rocks and on grassy slopes.

Idaho : Beaver Caii on, 1895, C. L. 5//r^r, jo.?p (type in Herb.
N. Y. Bot. Garden) and joj/.

Wyoming: Trout Lake, 1899, Aven & Elias Nelson, 584.8 ;
Head of Pole Creek, 1895, Avcn Nelson, 13 21.

Sophia leptophylla sp. nov.

Stems 5-7 dm. high, sparingly glandular : leaves very thin,
dark green, only once pinnate ; segments of the lowest leaves
rounded ; those of the middle lanceolate and somewhat incised ;
the upper segments entire : pedicels slender, in fruit 5-8 mm. long,
spreading: petals bright yellow, about 1.5 mm. long : pod linear,
8-10 mm. long, much less than i mm. wide, torulose, spreading,
arcuate, curved up, uniserial.

This is nearest related to 5. incisa (Engelm.) Greene, but differs
in the less incised leaves and horizontal curved pod. It grows at
an altitude of about 2000 m.

Colorado: Foothills, Larimer county, 1895, Crandall& Coiven,
S5 (type in herb. N. Y. Bot. Card.).

Wyoming: Antelope Basin, Albany county, 1900, Aven Nel-
son, 7466 ; Centennial Hills, 1895, 16S6.

Smelowskia Americana

Hutcliinsia calycina Hook. Fl. Bor. Am. i: 58. 1834. Not
Uesv. 1 8 14.

Smelowskia calycina Gray, Proc. Acad. Philad. 1863 : 58. Not
C. A. Mey. 1831.

While in Europe last summer, I looked up the Asiatic type Qf
S. calycina, and this differs considerably from ours especially in the
long villous pubescence.

Rydberg : Studies on the Rocky Mountain Flora 240

Draba brachystylis sp. nov.

Perennial with a taproot : stems several, decumbent or ascend-
ing with divergent branches, 1-1.5 dm. high, hirsute : basal leaves
oblanccolatc or spatulate, 1.5-2 cm. long, hirsute with branched
hairs ; stem leaves few, oblong or lanceolate, sessile : pedicels
short, in fruit 2-5 mm. long, spreading : pod oblong, finely stellate,
8-12 mm. long, ascending: style very short, almost obsolete.

The type sheet of this species is labeled Draba Carolina var.
micrantha, but the plant evidently is a perennial and nearest related
to D. streptocarpa ; but differs in the divergently branched stem,
stellate pod and the almost obsolete style. It grows at an altitude
of about 3000 m.

Utah: Alta, Wasatch . Mts. , 1879, M. E. Jones, ijjy (type
in herb. N. Y. Bot. Card.).

Draba decumbens sp. nov.

Perennial with a taproot, a short cespitose caudex and a large
rosette of basal leaves : stems decumbent, 1-2 dm. high, sparingly
pubescent, few-leaved ; basal leaves broadly .spatulate, entire, 2-4
cm. long, finely grayish stellate, rather thick ; stem leaves ovate
to ovate-lanceolate, sessile, entire, 1-2 cm. long : calyx sparingly
villous ; petals pale yellow, about 4 mm. long : pedicels very
short, in fruit 2-4 mm. long : pod oblong, 8-15 mm. long, densely
pubescent, usually twisted : style about i mm. long.

This is a member of the D. aiirca group, and distinguished
from all the North American species by its decumbent stems, large,
thickish entire basal leaves.

Colorado: Gray's Peak, 1895, Rydberg {ty^Q in herb. N. Y.
Bot. Gard.).

Draba Macouniana sp. nov.

Annual, almost scapose : stems ascending or decumbent, less
than I dm. high, glabrous and shining ; basal leaves numerous,
broadly spatulate or obovate, 5-10 mm. long, finely stellate, rather
thick ; stem leaf usually one, oval, obtuse, 5 mm. long : petioles in
fruit spreading, 6-8 mm. long: petals white, less than 3 mm. long :
pod oblong-linear, erect, about i cm. long and 2 mm. wide : style
obsolete.

This species is nearest related to D. crassifolia, but differs in
the whiter flowers, and the broad and short basal leaves, which
are finely stellate, not hirsute ciliate. The type grew at an alti-
tude of about 2000 m.

241 Rydberg : Studies ox the Rocky Mountain Flora

British Columbia: Summit of Rocky Mountains, \?,go, Jolui
Macoiin (type in herb. Columbia University).

Draba Parryi sp. nov.

Annual, perfectly glabrous, except a few cilia on the petioles :
stems several, usually less than i dm. high, scapiform or rarely
with a stem leaf: basal leaves numerous, linear or narrowly linear-
oblanceolate, 1.5-2.5 cm. long; pedicels spreading, in fruit 5-8
mm. long : flowers small ; petals scarcely 2 mm. long, white or
light yellow : pods erect, oblong, 5-8 mm. long, 1.5-2 mm. wide,
glabrous : style obsolete.

This is closely related to D. crassifolia but the latter has
broader leaves, which are more or less hirsute on the surface and
distinctly ciliate on the margins.

Colorado : Foot of Gray Peak, 1872, E. L. Greene (type in
herb. Columbia University); 1872, C. C. Parry; Cameron Pass,
1896, C. F. Baker.

Wyoming : Telephone Mines, 1900, Aven Nelson, jSyg.

Draba cana sp. nov.

Perennial with a taproot and short cespitose caudex, whole
plant densely grayish stellate ; stem 1-2 dm. high, often branched :
basal leaves numerous, oblanceolate or spatulate, 1-1.5 cm. long,
entire or minutely but sharply toothed, densely stellate ; stem
leaves lanceolate to ovate, about i cm. long : racemes many-flow-
ered ; pedicels short, nearly erect, in fruit 2-3 mm. long : flowers
small ; petals white, about 3 mm. long : pod Hnear-oblong, 6-8
mm. long, densely pubescent : style about 5 mm. long.

This has gone under the name of D. incana, but the latter is
much less densely stellate and has a glabrous, much broader pod.
I have seen only one specimen of true D. incana from this conti-
nent, and it was collected in Labrador. Most of those under this
name belong to D. cana, which ranges from Labrador to the
Yukon Territory, south in the mountains to Colorado. The fol-
lowing is regarded as the type.

Alberta : Morley, foothills of the Rocky Mountains, 1887,
John Macoun (herb. Columbia University).

Draba McCallae sp. nov.

Perennial with a cespitose base, whole plant sparingly stellate
but green : stems several, simple, about 3 dm. high : basal leaves

Rydberg : Studies on the Rocky Mountain Flora 242

spatulate, 10-18 mm. long, entire or nearly so ; stem leaves ovate,
about I cm. long, sharply dentate or entire : raceme elongated :
pedicels slender, in fruit about i cm. long, erect : petals white, 4-5
mm. long : pod (in the specimen not fully developed) ovate, densely
but finely pubescent, much shorter than the pedicels : style 0.75
mm. long.

This species belongs also to the D. incana group, but differs
from that species in the elongated peduncle, long pedicels, short
pubescent pod and large petals.

Alberta: Valley below Mt. Aylmer, 1899, W. C. McCalla,
226^ (type in herb. N. Y. Bot. Gard.).

Draba Columbiana sp. nov.

Perennial ; with a cespitose base, sparingly and finely stellate
and ciliate on the margin of the leaves : stems slender, about 2
dm. high, simple: basal leaves many, oblanceolate, 10-15 mm.
long, denticulate or entire ; stem leaves ovate to lanceolate, about
I cm. long denticulate : raceme elongated, pedicels slender, ascend-
ing, 5-7 mm. long in fruit : flowers small ; petals slightly over 2
mm. long, white : pods linear-oblong, finely pubescent, over i cm.
long, style very short.

This species resembles much the preceding in habit, but the
flowers are much smaller and the pods longer.

British Columbia: Revelstoke, i^^o, John Macowi {iypQ in
Herb. Columbia University).

Geranium Pattersonii sp. nov.

Cespitose perennial with a thick root and short caudex : stems
ascending or diffuse, 2-4 dm. long, densely glandular villous with
long spreading hairs, petioles of the basal leaves 5-10 cm. long,
also glandular villous ; blades reniform in outline, about 3 cm.
broad, glandular villous on both sides, deeply 5-7-cleft ; lobes
broadly obovate-cuneate, 3 -toothed ; teeth broadly ovate, abruptly
short-acuminate : stem leaves similar, but short-petioled or the
uppermost subsessile : pedicels and calyx glandular long -villous ;
the former in fruit 1.5-2 cm. long: sepals in fruit about 8 mm.
long, oval with a short bristle-tip about i mm. long : petals obo-
vate about i cm. long, light purple, or rose color, with darker
veins : style in fruit about 2 cm. long, glandular-pubescent ; beak
about 4 mm. long : carpels pubescent : seeds finely reticulate.

This species is perhaps nearest related to G. Parryi (Engelm.)
Heller, but is lower, more densely glandular, has broader and

243 Rydberg : Studies on the Rocky Mountain Flora

shorter lobes to the leaves and their teeth are broadly ovate, ab-
ruptly acuminate, instead of lanceolate. The general habit is per-
haps more like G. cacspitosian James, but in that species the lower
part of the stem is grayish-pubescent with short reflexed hairs.
G. Pattersonii is a subalpine plant growing at an altitude of 2500-
3000 m.

Colorado: Gray Peak, 1895, P. A. Rydberg (type in herb.
N. Y. Bot. Gard.); foot of Pikes Peak, 1896, C. L. Shear, ^702 ;
mountains, near Empire, 1892, H. N. Patterson, lyy and ij6 in
part ; North Cheyenne Canon, 1896, Ernest A. Bessey ; also 1845.
Free 1)10 lit, 61.

Geranium strigosum sp. nov.

Gcranuim Riehardsonii Wats. King's Rep. 5 : 49, in part.
1 87 1. Not Fisch & May.

Erect perennial with a short thick rootstock : stem 4-8 dm.
high, finely reflexed strigose ; petioles of the basal leaves 2-5 dm.
long, also sparingly strigose ; blades reniform in outline, 7-12 cm,
in diameter, finely strigose on both sides, deeply 7-cleft ; divisions
rhombic in outline, 3-cleft and toothed; teeth lanceolate, acute;
stem leaves similar, but short-petioled ; stipules linear-lanceolate,
long-attenuate : inflorescence open, glandular pubescent : sepals
densely glandular, oval, about 8 mm. long ; bristle tips 1.5-2 mm.
long ; petals broadly obovate, rose color or light violet, strongly
purple-veined : style in fruit about 3 cm. long, beak about 6 mm.:
seeds reticulate.

This is nearest related to G. viscosissiviuni, but the corolla is
lighter in color and the pubescence is different, being decidedly
reflexed, strigose, not at all glandular, except on the inflorescence.
It grows in the valleys to an altitude of perhaps 27CO m.

Wyoming: Copperton, 1901, F. Ttveedy, 759/ (type in herb.
N. Y. Bot. Gard.); Big Horn Mountains, 1900,^^5^7.

Utah: City Creek Canon, 1880, M. E.Jones. i8yi ; Parley's
Cafion, 1901, S. G. Stokes; Black Rock and Uintahs, 1869, S.
Watson, 204.

Oxalis Coloradensis sp. nov.

Perennial with a slender rootstock : stem slender, erect, 1-2
dm. high, sparingly villous ; leaves digitately ternate ; petioles
2-4 cm. long, sparingly villous : leaflets broadly obcordate, gla-

Rydberg : Studies on the Rocky Mountain Flora 244

brous, 8-12 mm. long: peduncles very slender, about 5 cm. long :
inflorescence cymose, but often reduced to 2-3 flowers ; pedicels
sparingly strigose : sepals 4-5 mm. long, oblong : petals yellow,
6-7 mm. long: pod cylindric with a conical apex, 12—15 mm.
long, 2.5-3 ^^- wide, glabrous.

This is nearest related to 0. cymosa Small and 0. Brittoniae,
but differs from the former in the blunter sepals, the appressed
pubescence of the pedicels and the glabrous pod, and from the
latter in the simpler habit, the longer glabrous pod and paler and
larger leaves. It grows in grassy valleys at an altitude of 1800-
2500 m.

Colorado: Sangre de Christo Creek, 1900, Rydberg & Vrce-
land jg20 (type in herb. N. Y. Bot. Gard.); Foothills, Larimer
county, 1895,/. H. Coiven (mixed with 0. stricta L.).

Lupinus Scheuberae sp. nov.

Perennial ; stem 4—6 dm. high, sparingly strigose or almost
glabrous, slender : petioles of basal leaves about i dm. long ; leaf-
lets 7-8, oblanceolate, 5—7 cm. long, acute and mucronate, green,
glabrous above, sparingly appressed pubescent beneath ; stipules
lanceolate, attenuate ; stem leaves similar but shorter petioled :
racemes 1-1.5 dm. long, not dense; bracts lanceolate, 7-8 mm.
long, deciduous : flowers 10-12 mm. long: calyx densely white
silvery puberulent, strongly gibbous above ; lower lip lanceolate,
fully twice as long as the triangular-ovate upper Up : corolla dark
purple but lighter and almost white towards the base ; banner
slightly longer than the wings, silky puberulent on the middle of the
back : pod densely white silky with appressed hairs, 2— 4-seeded.

This is nearest related to L. pscudoparviflonts , but differs in the
somewhat larger flowers, the strictly appressed white pubescence
on the calyx and pedicels, the gibbous, but not spurred calyx, and
the broad and short upper lip of the latter. It grows in woods at
an altitude of 2,000-2,400 mm.

Montana : Garnet county, 1901, Mrs. Emma Ware Sheiiber^
/J5 (type in herb. N. Y. Bot. Gard.).

Wyoming: Copperton, 1901, Frank Tweedy, 4.216.

Lupinus spathulatus sp. nov.
Lupinus parviflorus S. Wats. King's Rep. 5: 1871. Not
Nutt.

245 RvDBERG : Studies ox the Rocky Mountain Flora

Perennial : stem 8-10 dm. high, glabrous or sparingly strigose
above, leafy ; stipules narrowly lanceolate, attenuate : petioles 3—5
cm. long ; leaflets about 7, spatulate or broadly oblanceolate, about
4 cm. long, mucronate, those of the lower leaves obtuse, those of
the upper more acute, glabrous above, finely puberulent beneath ;
raceme elongated, about 2 dm. long, many-flowered : flowers sub-
verticillate ; pedicels about .5 cm. long, as well as the calyx,
pubescent with short spreading hairs ; bracts lanceolate, attenuate,
deciduous, longer than the buds ; calyx gibbous at the base ; lower
lip lanceolate, about twice as long as the short ovate upper one :
corolla dark blue, with lighter keel ; banner glabrous, rounded and
shorter than the wings.

This has been confused with L. parviflorns, which it closely
resembles, differing mostly in the shape of the calyx ; the lower
lip in that species is only slightly longer than the upper and the
banner is larger and less rounded. The type grew at an altitude
of 2400 m.

Utah: Wasatch Mountains, 1869, 5. Watson, 22 j (type in
herb. Columbia University).

Lupinus flavescens sp. nov.

Perennial : stem about 3 dm. high, appressed pubescent with
rather long hairs: stipules lanceolate, densely hairy: petioles often
over I dm. long, appressed silky ; leaflets about 7, oblanceolate,
3-4 cm. long, appressed silky on both sides : inflorescence short;
bracts lanceolate, early deciduous ; pedicels 8-10 mm. long, as
well as the calyx densely pubescent with short, spreading hairs:
calyx gibbous at the base ; its lips almost equal in length, both
about 8 mm. long: corolla 12-14 mm. long, pale yellow; banner
with a darker spot, glabrous.

The type has been known as L. sulplmreus 2ir\d v/diS included
therein questionably by Watson ; but differs in the larger, paler
flowers, and the longer lips of the calyx, and the longer pubes-
cence.

Idaho or Montana : Medicine Clay Prairies, Wyeth (type in
herb. Columbia University).

Lupinus lucidulus sp. nov.

Perennial : stem 5-6 dm. high, branched above, finely silky
strigose, especially the upper portions, leafy ; stipules minute, ovate,
acuminate : petioles 3-5 cm. long ; leaflets y-d), linear-oblanceo-

Rydberg : Studies on the Rocky Mountain Flora 246

late, silky and silvery on both sides, about 3 cm. long : raceme
1-2 dm. long, lax: flowers scattered ; pedicels and calyx densely
pubescent with short strictly appressed silky hairs ; lower lip of
the calyx about 5 mm. long, lanceolate, almost twice as long as
the short ovate upper one : corolla yellow or light pink ; banner
slightly exceeding the wings, with a few short hairs on the back,
with a darker spot, about 8 mm. long : pods densely silky pubescent.

This species is nearest related to L. sulpJnircus, but differs in
the more silvery pubescence, the smaller flowers and the appressed
pubescence of the calyx and pedicels. The type grew at an alti-
tude of 1900 m.

Wyoming: Spread Creek, 1897, F. Tweedy, zyi (type in
herb. N. Y. Bot. Garden).

CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 24

THE NIDULARIACEAE OF NORTH AMERICA

BY V. S. WHITE.

NEW YORK
1902

[Reprinted from Bulletin Torkey Botanical Club, 29 : 251-280, May 26, 1902.]

The Nidulariaceae of North America =■'

By V. S. White

(With Plates 14-18)

The members of the family Nidulariaceae are small fungi,
seldom attaining a height of 15 mm., the average height being
from 5-7 mm. The ripe sporophores are usually more or less regu-
larly cup-shaped, containing from 10-20, or in some species, more,
lenticular seed-like bodies, which are analogous to the chambers
of the gleba in the other chief groups of the gastromycetes ;
these bodies are usually known as sporangioles or peridiola,
while the cup is known as the peridium.

They are widely distributed, as will be shown in the following
pages, and several of the species are fairly common, appearing
principally in wet weather, in late summer and autumn, often per-

* This revision of the family was made possible by the large amount of material
found at the New York Botanical Garden, principally forming part of the Ellis collec-
tion, and by additional material furnished by Professor L. M. Underwood, from his own
private collection. Professor F. S. Earle and Professor C. H. Peck also kindly
loaned specimens. Thanks are due to Professor G. F. Atkinson, of Cornell Univer-
sity, for the loan of three drawings of N'idtilaria Alabamemis made by Mr. H.
Hasselbring, and for a specimen of this same species ; to Professor Farlow, of Harvard
University, for kindly allowing specimens of Cyathtis pallidus Berk. & Curt, and Cya-
thus IVrightii Berk., to be examined and sketched ; to Mr. Stewardson Brown, of the
Philadelphia Academy of Science, for his courtesy in facilitating the examination of
Schweinitz's specimens nos. 2211-2220 ; to Miss Anna Murray Vail, Librarian of the
New York Botanical Garden, for help in tracing references ; to Mr. E. S. Salmon, for
looking up some data at Kew, England, and most especially to Professor L. M. Under-
wood, under whose direction the work was undertaken.

[Issued 26 May] 251

252 White : The Nidulariaceae of North America

sisting through the winter months. They grow on manure, sticks,
shavings, dried herbaceous plants, and Cntcibiilnvi cnicibidiforme is
frequently found on old pieces of sacking. On a certain old
board walk at Lakevvood, N. J., this last species was found by the
hundred growing in regular series between the joints of the boards.
A further study of the plants will probably show that they are far
more common and widely distributed than is at present thought,
for they are likely to escape detection owing to their small size
and inconspicuous coloring,.

The order Nidulariales comprises but one family, the Nidula-
riaceae. Until quite recently this family was considered by De
Toni * and others to include the genera Cyatlms, Crucibiihtin,
Nididaria, Thdcbolus, Dacryobohis, Sphaerobolus, Polyangmm and
Atractobolus, the last two included as doubtful genera. Fries t in-
cluded besides the three first mentioned genera, /^;-^,f/i!;/z(?;/Schwein.
Myriococciim Fries, and Polyangmm Link. The two latter genera
are certainly out of place here, Myriococciim belonging under the
Perisporiales,! and Polyangium is not mentioned by either Sac-
cardo or Fischer. In Fischer's revision of the family in 1899,
he places Arachnion Schwein.§ under "Doubtful Genera" in the
Sclerodermataceae ; for Sphaerobolus Tode he establishes a sepa-
rate family ; Thclcbolus is placed under the Ascobolaceae, and
Dacryobohis and Atractobolus are included in the same family but
as doubtful genera.

This leaves only the three genera, Cyathus (for which there is an
earlier form of the name, Cyathia) Crucibulum, and Nididaria, in this
family. Concerning the last genus a question already raised by Otto
Kuntze presents itself in regard to the correct name, since Roth's
GranulariaW antedates Fries and ^ox^\\o\m s> Kididaria\ by over
a quarter of a century. Roth's description and figure of Granu-
laria pisiformis place this genus on a certain footing, and there can
be no hesitation in answering the above question. Recently some

*Saccardo, Syll. Fung. 7 : 2S. 1887.

fSyst. Myc. 2 : 296. 1822.

+ Saccardo, Syll. Fung, i : 29. 18S2.

\ Engler & Prantl, Nat. Pflanzenzenfam. i'-** : 339. 1899.

|| Ust. Ann. Bot. 1 : 6. //. /./. i. I79I.

\ Syn. Gast. 2. 1817-18.

White : The Nidulariaceae of North America 253

plants resembling Crucibiihan in habit * have been described under
Nidiilaria which differ so much in the peridial structure and char-
acter from the type of the genus, with the exception of not having
the sporangioles attached to the inner wall of the peridium, that it
seems best to establish another genus in order to simplify as much
as possible the arrangement of the family. It was ardently desired
to retain the name Nidiilaria, but unfortunately it has been found
impossible to do this as all seven of the species which Fries f gives
under his section Nidiilaria conform to the type of Granularia, so
the name Nidiilaria must pass into synonymy.

It is interesting to note that Corda % included only Nidiilaria
and Cyathiis under the Nidulariaceae. Nees noted a resemblance
between this family and Polysacciini, but the latter genus is in-
cluded by Fischer § under the Sclerodermataceae as Pisolithiis
Alb. & Schwein.

Although the members of this family have been mentioned in
nearly all works treating of fungi from the time of Clusius, i6oi^
little was known of their structure and life habits until 1842,
when Schmitz || wrote a short paper on CyatJuis, and two years
later the brothers Tulasne,! instigated by this first effort to con-
tinue the study of these interesting plants, prepared their treatise
on the organization and fructification of the Nidulariaceae, which,
with the supplementary work of Sachs,** Eidamff and Brefeld,J|
forms the foundation of our exact knowledge of this group of
fungi.

The American species have never been monographed, and even
the mention of plants belonging to this family has been compara-
tively infrequent.

* Under Crucibulum De Bary (Comp. Morph. 321) calls attention to the fact that
he was unable to find a funiculus in some peridioles but says " the coil always showed
a small point which answers to it." In the specimens referred to this genus there was
absolutely no trace or rudiment and it would be impossible even in young sporangioles
to determine to which side they had been attached.

t Anleit. 103. 1842.

X Syst. Myc. 2 : 300. 1823.

\ L. c. 338.

II Ueber Cyathus. Linnaea, 16: \\l. pi. 6, y. 1842.

^Ann. Sci. Nat. III. i : 41-107. //. 1844.

**Bot. Zeitung, 833-845, 849-861. 1855.

tfCohn's Beitr. z. Biol. 2 : 221-245. 1876.

f J Bot. Mitt, iiber Schimmelpilze, 3 : 176-180. 1877.

254 White : The Nidulariaceae of North America

For a long time the Nidulariaceae seemed to be a bone of con-
tention among various botanical writers, in the fact of their appar-
ently differing from other fungi in having true and visible seeds.*
In 1688 Camerarius published a dissertation " de Fungo calyci-
formi seminifero," setting forth his ideas on the subject and claim-
ing that these fungi had seeds. Marsigli in 171 3, on the contrary,
tried to exclude what he calls "these simple plants" from the
fungi, no fungi having true seeds as these plants seemed to* have.
In 1714 Tournefort was also inclined to exclude them from the
fungi, but Ant. de Jussieu in 1728, denied these propositions, and
claimed emphatically that fungi have seeds like true plants. Forty
years later fungi were still being classed outside of the vegetable
kingdom, particularly so because the sporangioles of CyaiJms were
never seen to germinate. Necker in 1783 wanted to make an
intermediate kingdom to consist of the fungi, and Pico five years
later, said that he had absolutely proved that the nature of these
productions was purely animal. In 1791, BuUiard incontestibly
proved that all fungi have seed, but he called the sporangioles of
Cyatlius seeds, and added "that they seem somewhat out of pro-
portion to the size of the fungus." Hoffman says " potius capsa
seminalis quam semen ipsum " — " they are less seeds than capsules
filled with seeds," while Micheli said that the spores are hard to
see with a good magnifying glass.

These fungi have had several popular names, probably owing
to their quaint and attractive appearance. Besides the well-known
name of " bird's-nest fungi," as far back as 1724 they have been
"called in Worchestershire cornbells, where they grow plenti-
fully,"t and in Lincolnshire " we find that a kind of fungus like a
cup or old-fashioned purse with small objects inside is called a
' fairy purse ' and we presume that the small objects represent the
fairies' cash. "J

Synopsis of the Genera of tlie Nidulariaceae

Sporangioles attached to the inner wall of the peridium.

Peridium composed of three layers ; spores mixed with filaments. I. Cyathia.

*Most of what follows in this paragraph is translated and adapted from Tulasne's
monograph, pages 54-56, as several of the works referred to were unfortunately unob-
tainable.

t Ray, Syn. Ed. 3, 2 : 20. 1724.

X Friend, Flower Lore, 34. 1889.

White : The Nidulariaceae of North America 255

Peridium composed of one homogeneous layer ; spores not mixed with filaments.

II. Crucibulum.
Sporangioles not attached to the inner wall of the peridium.

Peridium thick, opening by a regular definite mouth. III. NiDULA.

Peridium thin, rupturing irregularly. IV. Granularia.

I. Cyathia p. Br., Civ. and Nat. Hist. Jamaica, 78. 1756
Cyathus Hall. Stirp. Helvet. 3: 127. 1768

Peridium composed of three distinct but closely connected
layers. Mouth at first closed by a membrane (epiphragm), usu-
ally white, opening at maturity; sporangioles flattened, umbilicate
beneath, attached to the inner wall of the peridium by a complex
elastic cord (funiculus), and with thick horny filaments intermixed
with the spores.

Clusius,"^ in 1601, made the first mention of a Cyathia under
the name of fungus minimus avMVjao^, and described it in Latin,
the following being a free translation of the entire paragraph :
" Moreover this fungus, which I will call anonymous, is very dif-
ferent from the preceding ones, and I consider it to be the smallest
of all, for it is barely half an inch high. In the fall a great many
grow, without petiole, on wooden boards away from dust and sand.
They have the color of cinders or are of a lifeless color. The shape*
which is so small, appears to be undeveloped, scarcely as large
and as thick as the top of a little finger. Sometimes they grow
alone or when in numbers two, three or four adhering together,
and when ripe, they throw off the top part and appear full of a
viscous juice, and of seeds which are about the size of the seeds of
cyclamen, but have the outline of small fungi and are apparently
cinder-colored. There is no doubt that this fungus grows in
Italy, for I remember that a friend of mine sent me, once upon a
time from there, some of these seeds, dried and with a certain
strange name, asking me if I could find out what they were. For
there are certain characters who endeavor to catch and buy praise
from the ignorance of others."

There seem to be no intermediate references to this plant until
1 67 1, when Bauhin f mentioned " Fungus minimus lignelis tabellis
areolarum hortorum adnascens." Mentzel,t in 1682 first figured

* Rar. Plant. Hist, cclxxxvii. i6oi.

t Pinax, Lib. X., Sec. 5, no. 39, 374. 1671.

% Ind. Nom. Plant. Univ. //. 6. 1682.

256 White : The Nidulariaceae of North America

a plant which was, without doubt, Cyathia Icntifera. Loeselius,*
and Marsilius,t made passing references to this same species, and
in 1729 MicheH I described and figured two species, citing one
from the above-mentioned writers, and the second from Ray,§
Dillenius |1 and Vaillant, 1 the latter species being unquestionably
Cyathia striata. . Linnaeus,** in 1753, gives under his section
"Acaules" of the genus Peziza — Peziza campamdata lenfifera,
first referring to Hortus Cliffortianus (1737) where we find that
the only species of Peziza which he gives, and to which he refers
what is evidently a Cyathia is not a Peziza in the modern sense,
but the first known type of Cyathia; and the only proof that he
knew what, a Peziza really was, is his reference to Dillenius' tt
work published in 1719, in which a true Peziza is figured. Dil-
lenius divided \X\q Pezizae into two sections: {i) Membranacei et
tenuiores — the true Pezizae, and (2) Dunores calyciformes, seinini-
feri pHtati, under which head he gives Peziza calyciformis lentifera
laevis (our Cyathia lentifera) and Peziza calyciformis lentifera hir-
sutd' (our Cyathia hirsuta).

The genus Cyathus was founded in 1768 by Haller, twelve
years after Cyathia P. Br., and since that time many species have
been described from nearly all parts of the world. Fischer ||
states that there are forty-two known species.

From our own country not many species have been described,
Schweinitz was the first to mention these plants ; in 1 818 he named
ten species, but not all of them are to be referred to the genus

Cyathia. In 1869 Berkeley & Curtis§§ described Cyatlms pallidus
from Cuba, also reporting Cyathus intermedins, Cyathus Montagnei,

Cyathus limbatus, Cyathus Poeppigii, Cyathus inicrosporus, and

Cyathus Lesueurii collected by Wright from the same island.

Berkeley, II II in 1873, described from Connecticut, Cyathus Wrightii,

*Flor. Prus. 98. 1703.

t Dissert. Gen. Fung. 17. //. a. 1714.

jNov. Plant. Gen. 222.pl. 102. f. i, 2. 1729.

§Syn. 3: 21. 1704.

||Giss. 196. 1719.

•jBot. Paris. 57. //. //. / 4, J. , 1727-

**Sp. PI. 2: 1 180. 1753.

tt/. ^.

Jt Engler & Prantl, Nat. Pflanzenfam. i^** : 328. 1899.

g§ Jour. Linn. Soc. 10: 346. 1869.

II II Grevillea, 2 : 34. 1873.

White : The Nidulariaceae of North America 257

which has not since been reported. CyatJuts rufipcs was described
in 1897 by Ellis & Everhart * from Kansas, and the next year
Hennings, || described CyatJins nivcotomentosns from California.

The following summary will show the distribution of the
known species of Cynthia :

Europe, 7 species : C. complanata, C. deforviis, C. fimetaria, C. hirsuta, C.
lentifera, C. suhkulosa, C. umbrina.

Asia, 6 species : C. emodensis, C. Hookeri, C. intermedia, C. minima, C. Alon-
tagnei, C. sulcata.

Africa, 5 species : C. affinis, C. dasypus, C. pallida, C. hirstita, C. lentifera.

Australia, g species : C. Baileyi, C. descrtorum, C. fimicola, C. intermedia, C.
stercorea, C. Montagnei, C. pezizoides, C. pusio, C. lentifera.

New Zealand, 3 species : C. Colensoi, C. N'ovae-Zelandiae, C. similis.

North America, ii species: C. intermedia, C. lentifera, C. melanosperma,
C. Berkleyana, C. riigisperma, C. rufipes, C. hirsuta, C. stercorea, C. pallida, C.
Wrii^litii, C. fragilis.

Cuba, 7 species : C. intermedia, C. Montagnei, C. pallida, C. limbata, C. Poep-
pigii, C. Berkleyana, C. stercorea.

South America, 12 species : C. ambigua, C. dasypus, C. limbata, C. micro-
spora, C. Montagnei, C. plicata, C. Poeppigii, C. Puiggarii, C. scutellaris, C. hirsuta,
C. lentifera, C. Gayana.

i$>ii4>psis of tbe Species of Cyatliia

Peridium sulcate-striate within.

Spores large, more than 35// long. I. C. Poeppigii.

Spores small, less than 8 m long. 2. C. Berkleyana.

Spores of medium size, 12-20/i long.

Peridium sparingly clothed with short fasciculate hairs without.

3. C. intermedia.

Peridium thin, shaggy-tomentose without. 4. C. hirsuta.

Peridium very tough and brittle, rough tomentose. 5- C. dura.

Peridium faintly striate near the mouth, not sulcate within. 6. C. Montagnei.

Peridium smooth within, not sulcate.

Spores large, more than 40 /i long. 7- C. melanosperma.

Spores small, less than 15 /^ long.

Sporangioles darkish ; margin of peridium straight.

Peridium smooth or nearly so. 8. C. pallida.

Peridium strigose-tomentose. 9. C. rugisperma.

Sporangioles light colored ; margin of peridium often recurved.

10. C. lentifera.
Spores of medium size, 15-30 /< long.

Peridium with a red felt-like mycelial bulb at base. II. C. rufipes.

Peridium with a thick bulb of whitish mycelium. 12. C. Wrightii.

Peridium scarcely thickened at the base. 13. C. stercorea.

\ Bull. Torrey Club, 24 : 125. 1897.
f Hedwigia, 37 : 274. 1898.

258 White : The Nidulariaceae of North America

I. Cyathia Poeppigii (Tul.)
Cyathiis Poeppigii Tul. Ann. Sci. Nat. III. i : 'jj. pL ^. f.

23-25 ; pl- 5- /• 3-4- 1 844.

Peridia slender, goblet-shaped, 7-12 mm. high, 5-7 mm. wide
at the top, about i mm. at the base of the stem, dark chocolate-
brown, the outer surface somewhat shaggy, peeling, leaving a
comparatively smooth, deeply and closely striated surface, corre-
sponding with the deep inner striations, the inner surface brown,
somewhat lighter colored than the outer surface, somewhat shiny,
deeply striate ; mouth minutely fimbriate ; sporangioles blackish,
flattened, dull, 2-2.5 mni- in diameter; funiculus long; spores
38—45 fi long, 18—22 fi in diameter, very thick-walled, granular
within, oblong and often slightly curved. {PL i^. f. 1-4..)

Growing singly and in groups on manure and wet ground.

St. Croix (Danish West Indies) : A. E. Rickscckcr.

This species was originally described from Cuba and French
Guiana ; it is easily distinguished from the other striated species
of CyatJiia by its unusually large spores, and by its deeply and
closely striated outer surface.

2. Cyathia Berkleyana (Tul.)

Cyatlms uiicrosporus ^ Berkley anus Tul. Ann. Sci. Nat. III. i :
74. 1 844.

Peridia small, cup-shaped, 5—8 mm. high, 5-7 mm. wide at
the top, very much narrowed and rounded at the base, brownish
outside and somewhat shaggy ; mouth finely fimbriate, the inner
surface of the peridium shiny, grayish-brown, lighter colored than
the outer, coarsely striate ; sporangioles small, 2 mm. in diameter,
shiny, nearly black, flattened; spores small, 6-8 /i long, 4-6 /i
wide, somewhat thick-walled, hyaline. (/-*/. ^-Z,/- 5-7.)

On decayed stems of coarse herbaceous plants.

Jamaica : Cockerell.

This species was originally described from Brazil ; it some-
what resembles small specimens of Cyathia hirsnta, but it differs
from that species in the texture of the outer coat, which is much
less shaggy and thick, and in the much smaller spores.

3. Cyathia intermedia (IMont.)

Nidularia i)iteniieiiia^lor\\.. ; Sagra, Hist. Phy. Pol. Cuba, 321.
1838-42.

White : The Nidulariaceae of North America 259

Cyathiis intermedins (Mont.) Tul. Ann. Sci. Nat. III. i : 72.
//. 4.f. 4-7. 1844.

Peridia cup-shaped, not rounded at the base, 7-9 mm. high,
6-8 mm. wide at the top, 1-2 mm. at the base, hghtish brown,
clothed with fine hairs which grow in clusters, the tips of several
of which join together in little outwardly curved tufts, thin and
showing the inner sulcate markings through in the older peridia;
inner surface brown, shiny, sulcate-striate ; the mouth straight, not
flaring, clothed with a circle of short stiff bristles ; the sporangi-
oles about 2 mm. in diameter, flattened, blackish, depressed be-
neath, somewhat angular with a short attachment ; spores hya-
line, 12-18/Z long, 6-9// wide, thick-walled. {PI. 14. f. 8-10;
pi. 18. f. 18)

Growing in loose soil and on decaying wood.

[Cuba : Ramon de la Sagra^

Delaware: Faulkland, Commons.

This species was originally described from Cuba ; it differs
from CyatJiia hirsuta in the character of the tomentose outer cov-
ering of the peridium, and in the shorter and more rudimentary
attachment of the sporangioles to the inner peridial wall.

4. Cyathia hirsuta (Schaeff.)

Peziza lentifera ^ Linn. Sp. PI. 2 : 1180. 1753.

Peziza sessilis campaimlata villosa Scop. Fl. Carn. 57. 1760.

Peziza prima Schaefif. Fung. Bav. et Palat. Icon. 2 : //. iy8.
1763.

Cyathushirsiitusintus striates Hall. Stirp. Helvet. 3:127. 1 768.

Peziza cyathiformis Scop. /. />.* Fl. Carn. 2 : 486. 1772.
[Ed. 2.]

Peziza hirsuta Schaefif. Fung. Bav. et Palat. 4 : 1 24. 1 774,

Peziza striata Huds. Flor. Ang. 634. 1778. [Ed. 2.]

Peziza hirsuta ')^?!Xsz\\, Elench. Fung. 127. 1783.

Nidtilaria striata With. Boi. Arrang. 3 : 446. 1792 [Ed. 2] ;
Sibth. Fl, Ox. 393. 1794; Fries. Syst. Mycol. 3: 298. 1822;
Schwein. Trans. Amer. Phil. Soc. 4: 252. 1834.

Cyathus striates Willd. Fl. Berol. 399. 1787; Hoffm. Veg.
Crypt, fasc. 2 : 33. //. 8. f. j. 1790; Pers. Syn. Meth. Fung. 237.
1801; Nees, Syst. 140. 1817; DC. Fl. Fr. 2: 269. 1805; Tul.
Ann. Sci. Nat. III. i : 6y. 1844.

*As the first cited reference under the description of this species belongs to C. len-
tifera this name cannot be taken up for this species.

260 White : The Nidulariaceae of North America

Peridia 10-15 mm. high, 8-10 mm. wide at the top, 2-4 mm.
at the base, usually straight trumpet-shaped, very gradually spread-
ing toward the top ; outer surface of the peridium dark brown,
shaggy fibrillose ; inner surface brown or grayish-brown, shiny,
striate-sulcate for about one half the height of the peridium, be-
coming smooth at the base ; mouth densely clothed with a circle
of stiff regular bristles ; sporangioles darkish, 2 mm. in diameter,
shiny, depressed beneath, and somewhat angular from the pressure
of one upon the other; spores 12-18 ji long, 6-9 11 wide, thick-
walled, hyaline, somewhat crescent-shaped. [PI. 14./. 11-14 ; pi.
18. f. 1-6.)

Growing singly and in clusters, on sticks, bark, etc.

Exsicc. : ElHs, North American Fungi, 729.

New York : Underivood, Tooke, Clinton, Brown, Mnrrill ;
Maine: Harvey; Connecticut: White; Pennsylvania: Gentry,
Lloyd; Ohio: Lloyd, James ; Georgia: Undei"wood ; Alabama:
Earle ; Iowa: Langlois ; Wisconsin: Williams; Indiana: Un-
dcriuood ; Montana: Anderson, no. 601; Colorado: Bethel;
Washington : Suksdorf ; Canada : Dearncss ; Puerto Rico : Un-
derzvood & Griggs ; Mexico : Egeling.

Tulasne * has given an account of the development of this
species from which the following is freely translated :

The young plants arise on the thick felt-like mycelium in the
shape of small cylindrical jelly-like masses. They are at first
white and smooth, gradually changing to a brown color and their
surface becomes somewhat scaly. The three coats which com-
pose the peridium are indicated at this stage, when the outer
brownish scaly portion begins to form itself into the outer coat,
while the walls of the white glutinous inner substance become
differentiated from the fruiting portion into a very thm membrane
which encloses these parts, a thin portion of the glutinous mem-
brane remaining between these two coats welding them together
and so forming the triple peridial wall. It is now that the
sporangioles begin their growth, gradually absorbing the sur-
rounding substance into themselves, so that when the plant is
mature and gradually opens, the sporangioles alone remain within
the cup attached to the sides of the peridium by means of a funic-
ulus of a complicated structure. The sporangioles are in most

*Ann. Sci. Nat. III. i : 50-54.//. 3 ; pi. 4./. 1-3 : pi. 8. f. 1-12. 1844.

White : The Nidulariaceae of North America 261

cases composed of three parts, namely, of a thick and usually
dark colored coat, a light hyaline thickly interwoven portion
which bears the basidia, which in their turn bear the spores. The
center part is composed of innumerable spores mixed with branch-
ing filaments. The walls of the spores are very thick, these
spores having long been separated from their basidia. The funic-
ulus is composed of three parts ; the lower portion is variable
as to length and thickness, its lower extremity is expanded and
attached to the peridium ; the upper portion is more regular in
shape and is attached by its upper extremity to the sporangiole ;
it is rather swollen in the center, in which portion there is enclosed
a long and slender filet composed of innumerable hyaline, inter-
woven filaments, which can be drawn out if great care is used in
the process and forcibly distended without breaking, according to
Tulasne to a length of 12 cm. These two parts are joined by a
much narrower and shorter portion to which Tulasne gives the
name of " filet median."

4a. Cyathia hirsuta infundibuliformis var. no v.

Peridia 1.5-18 cm. high, funnel-shaped, 1-2 cm. wide at the
top, 1.5-2 mm. at the base, 4-5 mm. in the center where the
peridium is contracted ; outer surface of the peridium brownish,
coarsely shaggy-tomentose ; inner surface dull brownish, clearly
striate, but not below the contraction ; mouth minutely fimbriated ;
sporangioles dull brownish, 2 mm. in diameter, flattened ; spores
8-12 /J. wide, 14-17// long, thick-walled, hyaline. (P/. i-f-.f. ij.)

Growing singly on leaves and moss.

Montana : Sheridan, Fitch.

This variety differs from the type in the definite funnel-shape
of the peridium, and in the nature of the tomentum, and to a less
degree in the size of the spores. Further material may necessitate
a different treatment.

5. Cyathia dura sp. nov.

Peridia flaring trumpet-shaped, 8- 13 mm. high, 8-12 mm.
wide at the top, 2—4 mm. at the base, brownish-gray outside,
rough, shaggy, and very tough and brittle ; inner surface whitish,
dull, and marked with rather wide shallow striations ; mouth
recurved, ragged and much split, but not fimbriate ; bases of the
peridium contracted and in some specimens there is a thick felty

262 White : The Nidulariaceae of North America

brown mycelial band adhering ; sporangioles dull grayish, oval
or rounded, 2.5 mm. long, 1.5 mm. wide; spores 14-18 // long,
8-10 n. wide, hyaline and thick-walled. {PL 14. f. i6-ig.)

Colorado : Denver, Bethel, 8.

This species differs from all others seen in the character of the

peridium, which is very thick, hard and brittle.

6. Cyathia Montagnei (Tul.)

Cyathns Montagnei Tul. Ann. Sci. Nat. III. i : 70. pi. 4, f.
g-ii. 1844.

Peridia 8-10 mm. high, 6-9 mm. wide at the top, 2-3 mm.
at the base, spreading, cup-shaped, reddish-brown outside, closely
woolly-tomentose, but not shaggy, rather thin ; inner surface gray-
ish, dull, faintly striate close to the top, not sulcate ; mouth at
first curved inwards, then straight, but not recurved, closely and
very minutely fimbriate ; sporangioles 2 mm. in diameter, grayish,
black, somewhat lighter beneath, dull, flattened, angular ; spores
I 5-18 // long, 8-12 /i wide, thick-walled, granular, h)'aline. {PI.
14./. 20-22.)

On the ground and on decaying wood and chips.

Yucatan: C. F. Mill span gh, 7^7; [Cuba].

This species was originally described from Brazil ; it is readily
distinguished from the preceding species: (i) By the faint and
only partial striae of the upper part of the inner surface of the
peridium, and (2) By the woolly tomentose, and not shaggy nature
of the outer coat.

7. Cyathia melanosperma (Schwein.)
Nidiilaria vtelanospcruia Schwein. Trans. Am. Phil. Soc. 4 :

253- 1834.

Cyathus inelanospennus (Schwein.) De Toni ; Saccardo, Syll.
Fung. 7 : 42. 1887.

Peridia cup-shaped, clustered, 5-7 mm. high, 5-6 mm. wide
at the top ; the outer surface brownish and ver>^ shaggy ; the
inner surface smooth, shiny, dark gray ; mouth straight and
slightly fimbriate ; sporangioles black, flattened, about 2 mm.
wide ; spores large, 45-55 « long, 18-45 !'• wide, very thick-walled,
granular within, hyaline. {PI. ij.f. /-■/•)

Gregarious on the earth.

Penxsylvania : Bethlehem, Schiveinitz.

The above description is based on Schweinitz's original speci-

White : The Nidulariaceae of North America 263

mens, which are still in good condition at the Philadelphia Acad-
emy of Sciences. This appears to be a rare species, as it is known
only from its original collection. It differs from all the following
species of 6j'(?///zrt', which have a smooth inner surface, in having
a fimbriate mouth, though Schweinitz in his original description
notes that it has a smooth mouth, but he writes that Xididaria
stcrcorea has a fimbriate mouth, which is contradicted by his own
specimens.

8. Cyathia pallida (B. & C.)

CyatJms pallidiis B. & C. Jour. Linn. Soc. lO : 346. 1869.

Peridia 4-8 mm. high, 5-7 mm. wide at the top, about 2 mm.
at the base, slender urn-shaped, slightly expanded at the top and
contracted at the base, thin and membranous, fawn-colored out-
side, and minutely shaggy ; inner surface light lead-colored,
smooth, shiny ; mouth entire or torn but not fimbriate, sometimes
slightly striated ; sporangioles 2 mm. in diameter, flattened, the
edges very thin, and thicker in the middle, darker above than be-
low, shiny as if frosted, and under high magnification showing a
network of delicate rugose wrinkles ; spores 5-9 fi long, 5—7 n
wide, hyaline, thick-walled, oval-pointed or nearly round, pale
yellow in the mass. {PI. ij.f. 5-7.)

Growing on decayed wood.

Cuba : Wright ; Puerto Rico : Coamo Springs, Underzvood &■
Griggs.

The above description is based on the co-types of this species
in the Curtis collection at Harvard University.

9. Cyathia rugisperma (Schwein.)

Nididaria rugisperma Schwein. Trans. Am. Phil. Soc. 4: 253.

1834.

Cyathus rugispenmis (Schwein.) De Toni ; Saccardo, Syll.
Fung. 7: 42. 1888.

Peridia somewhat obconic, sharply contracted at the base, 5-8
mm. high, 4 mm. wide at the top ; outer surface lightish fawn-
colored, strigose-tomentose, becoming nearly smooth when
mature ; inner surface darkish brown, smooth, and shiny ; mouth
incurved in the young peridia, and completely covered by the
tomentose outer covering ; sporangioles black, flattened, roundish,
i-r.5 mm. in diameter ; spores 10-14 P- long, 8-10 fi wide, some-
what granular and thick-walled, hyaline. {PL 16. f. j-6.)

Growing singly on bits of shaving and fibrous matter.

Pennsylvania : Bethlehem, Schzveinitz.

264 White: The Nidulariaceae of North America

The above description is based on the original specimens.
Schweinitz, in his description of this species, says : " Sporangiis
non nitentibus, sed superficie reticulato-rugosa," but no such
markings were noticeable.

This is a pretty little species, and is known only from its
original collection.

lo. Cyathia lentifera (L.)

Peziza lentifera a Linn. Sp. PI. 2 : i i8o. 1753.
Peziza sessilis canipamdata laevis Scop. Fl. Carn. 57. 1760.
Peziza tertia Schaeff. Fung. Bav. et Palat. Icon. 2 : pi. 180.
1763.

CyatJms sericeiis intus laeins Hall. Stirp. Helvet. 3:127. 1 768.
Peziza cyathiformis Scop. /. p. Fl. Carn. 2 : 486. 1772 [Ed. 2].
Peziza sericea Schaeff. Fung. Bav. et Palat. 4: 125. 1774.
Peziza lentifera Huds. Fl. Ang. 2: 633. 1778 [Ed. 2].
Peziza Olla Batsch, Elench. Fung. 127. 1783.
Nidiilaria vernicosa Bull. Champ, i : 164. pi. 488. f 1 . 1791.
Cyathus laevis Hoffm. Veg. Crypt, fasc. 2 : -i^i. pi. 8.f. 2. 1792.
Coccigrne a lentilles Paulet. Tr. des Champ. 2 : 406. //. 18 j.

f 7-12- •1793-

Nidtdaria campamilata\\!\\ki. Bot. Arrang. 3 : 445. 1792 [Ed.

2] ; Sibth. Fl. Ox. 393. 1794 ; Fries, Syst. Myc. 2 : 298. 1822.

CyatJms Olla Pers. Syn. Meth. Fung. 237. 1801.

CyatJins vernicosiis DC. Fl. Fr. 2: 270. 1805; Tul. Ann.
Sci. Nat. III. I : 81. //. 5./. 14-23. 1844.

Nidularia pliimbea Pers. Champ. Comest. iio. 18 18.

Nidularia fascicularis Schwein. Trans. Am. Phil. Soc. 4: 253.

1834.

Cyathus campamdatus Corda, Anleit. Ixxx. //. D. f. 42 {iQ-

23). 1842.

Peridia 8-12 mm. high, 5-15 mm. wide at the top, 2-7 mm.
at the base, campanulate, the edge often recurved and expanded ;
outer surface lightish, brown or grayish-yellow, almost smooth,
sometimes having a few tow-like fibers attached which give it a
slightly roughened appearance ; inner surface smooth, whitish or
lead-colored, shiny ; mouth entire, often ragged and split ; spor-
angioles light-colored, 2-3 mm. in diameter, flattened beneath ;
spores 8-15 n long, 6-iO« wide, thick-walled, hvaline. {PI. 15.
/• 8-13:)

White : The Nidulariaceae of North America 265

Plants sessile, or sometimes stalked, growing singly or in clusters
on the earth, on dry chips, twigs, etc.

New York : Undenvood; Maine: Harvey, Bartle ; Connec-
ticut: White; Pennsylvania: Martin; Alabama: Earle ; Indi-
ana: Undenvood; Kansas: BartJiolomeiv, ioj2 ; Colorado:
Ravenel ; Utah : Harkness ; California: Orciitt, Baker ; Texas:
Young; Puerto Rico: Goll.

II. Cyathia lufipes (Ell. & Ev.)

Cyathus riifipes Ell. & Ev. Bull. Torrey Club, 24: 125. 1897.

Peridia very slender, obconic, .8-1.5 cm. high, 3-6 mm. wide
at the top, 1-1.5 mm. at the base, outer surface lightish brown or
tow- colored, very shaggy tomentose, and with a quantity of red-
dish-brown mycelium adhering at the base, forming a felt-like tuft ;
inner surface smooth, dark gray, shiny ; sporangioles 2 mm. in
diameter, dark, almost black, shiny, depressed beneath ; spores
25—29 n long, 18-22 fx wide, thick- walled, granular within and
hyaline. {PL 15./. 14-16.)

On old sods, "growing head downward,"

Kansas : BartJiolome'w ; Nebraska : Bates.

This species is readily distinguished from the other species of
Cyathus with a smooth inner surface by its very slender elongated
shape and its light densely tomentose peridium, besides the char-
acteristic feature of the red felt-like mycelial tuft at the base, from
which the species derives its name.

12. Cyathia Wrightii (Berk.)

Cyathus IVrightii Berk. Grevillea, 2: 34. 1873.

Peridia rather large, cup-shaped, 1.2-1.8 cm. high, .5-1.2 cm.
wide at the top, contracted somewhat at the base, then bulbose,
the bulb composed of mycelium and adhering earth ; outer surface
darkish brown, very rough, shaggy and encrusted with earth ;
inner surface smooth, shiny, darkish ; mouth entire, not fimbriate
though the fine hairs circle up about it; sporangioles dark, 1.5-2
mm. in diameter, shiny, flattened; spores 22-25 /" lo"g> 15-18 fx
wide. {P/. 16. f. 7-p.)

Connecticut : Growing singly on earth, C. WrigJit.

This species is not known to have been reported since the
original collection. The above description of external characters
is based on the cotypes at Harvard University. The spores were

266 White : The Nidulariaceae of North America

not seen ; the measurement being taken from some mss. notes on

Cyatlius by Massee, made from the type of C. Wrightii at Kew.

De Toni gives the measurement in Saccardo as " i 5 fx long by 10

p. wide."

13. Cyathia stercorea (Schwein.)

Nididaria stercorea Schvveinitz, Trans. Am. Phil. Soc. 4: 253.

1834.

Cyatlius LesiieiLrii Tul. Ann. Sci. Nat. III. i : 79. //. 5. /.

S-13. 1844.

Cyatlius Lcsueurii, var. minor Tul. /. c. 80.

Cyatlius stercoreiis (Schwein.) De Toni ; Saccardo, Syll. Fung.
7 : 40. 1888.

Peridia slender, campanulate, sessile or with an elongated
slender base, .5-1.5 cm. high, 4-8 mm. wide at the top, 1-3 mm.
at the base ; outer surface brownish fawn-colored, the young plants
being strigose with a copious covering of shaggy hairs, which
gradually disappear, leaving the mature peridium almost smooth,
or marked with circular depressed zones ; inner surface smooth,
shiny, lead-colored, somewhat darker toward the base; the mouth
entire or at first appearing fimbriate from the surrounding hairy
covering, but not ciliate-fimbriate ; sporangioles blackish-lead-
colored, smooth, shiny, 2 mm. in diameter, depressed beneath ;
spores 25-30// long, 20-25 a wide, thick-walled, hyaline, granu-
lar within. {PI. ij.f. 17-20 ; //. 16. f. i, 2.)

Growing on manure, bits of wood, etc.

Exsicc. : E. & E. Fungi Columbiani, 644 (as Cyatlius verni-
cosus) ; Ravenel, Fungi Amer. Exsic. 473 (as Cyatlius vernicosus\
474 (as Cyatlius Lesueurii) ; Ravenel, Fungi Car. Exsic. 3 : 73 (as
Cyatlius campanulatus)\ Ellis, N. A. Fungi, 1308 {Cyatlius ver7ii-
cosus) ; Shear, N. Y. Fungi, 316.

Pennsylvania : Schweiuitz ; New York : Underwood, Shear ^
Brozun, Vail; Massachusetts: Ellis, Knight ; Maine: Harvey;
New Jersey : Ellis ; Delaware : 0. S. ; South Carolina : Ravenel;
Virginia : Murrill ; Iowa : Langlois ; Indiana : Underwood, Arthur ;
Ohio : Morgan, Lloyd ; Alabama : F. S. Earle & C. F. Baker,
Tuskegee, Carver; Kansas: Cragin, 27 j ; Colorado : Baker ;
Nebraska : Williams ; New Mexico : F. S. Earle & E. S. Earle
Cock ere II ; Canada : Dearness, 114^.

This species varies very much as to size, shape and smooth-
ness of the peridium, which facts have been rather misleading, but

White : The Nidulariaceae of North America 267

such different types have been found in specimens from one locality
from the short, strigose, sessile or bulbose form to the slender,
almost smooth, elongated form — that there can be no question as
to their being in reality but one species. Tulasne does not mention
Schweinitz's Nidularia stercorea, and his species Cyathiis Lcsiicurii
from New Orleans was presumably described without knowledge
of the preceding species. His variety Cyatlins Lesiieiirii \2S . minor
from Carolina, closely resembles small forms of Cyathia stercorea.

Cyathia (?) sp.

Peridia cup-shaped, clustered, 4-6 mm. high, 3-5 mm. wide
at the top, 2 mm. at the base ; outer surface lightish brown,
nearly sriiooth, or minutely fibrous ; inner surface smooth, very
dark brown, almost black, shiny ; mouth entire or slightly
ragged, but not fimbriate ; sporangioles black, shiny, angular
beneath, rounded above, i mm. in diameter, showing no trace of
any attachment, numerous; spores 20—30/^ long, 18— 24/i wide,
thick-walled, granular within, hyaline. {PL 16. f. 16, 21, 22.^

No mention as to habitat.

The specimens on which the above description is based were

sent to Mr. Ellis by Professor E. Bethel (no. 9) from Denver,

Colorado, and were named Cyathus vcrnicosus. They consist

of four or five groups of from two to five plants in each. The

peridial wall is triple and the structure of the sporangioles is

much as in Cyathia, the spores being also intermixed with thick

horny filaments as in that genus ; but there is no trace whatever of

any attachment either on the under surface of the sporangioles or

on the inner surface of the peridium. The specimens are all

mature, so it is impossible to know what the nature of the young

peridia was, but the sporangioles are still in the peridia, filling them

completely, even with the margin quite unlike anything seen

belonging to the genus Cyathia. It is very possible that a genus

standing in the same relation to Cyathia, as Nidula does to Crjici-

buhini, will eventually have to be established to accommodate these

forms, but the material and data are not sufficient to justify taking

such a step at present.

Species inquirenda

Cyathus niveo-tomentosus P. Henn. Hedwigia, 37: 274. 1898

Sparsis : peridio obconico-cyathiformi, crassiusculo papyraceo

primo operculo albo tecto, sessili vel substipitato, extus niveo

268 White : The Niuulariaceae of North America

sericeo, dense tomentoso, margine integro, crasso, 3-5 mm. alto,
4-5 mm. lato, intus albo-flavescente ; sporangiolis innumeris,
lentiformibus, planis, tenue tunicatis, cinnamomeis, sericeis, 1-1.2
mm. diametro ; sporis ellipsoideis, obtusis, intus granulatis, hyalinis,
^-8.5 X 4- 5-5. 5 /^, episporio hyaline, tenui, levi.

California: Potter Valley. On decaying wood (Sept. 1894).
Purpus.

The original description is quoted, as no specimens of this
species were seen, and until more material has been collected it is
difficult to decide with any degree of certainty just where it
belongs. The author adds that this species appears to be related
to Cyathia pallida.

II. Crucibulum Tul. Ann. Sci. Nat. III. i: 89. 1844

Peridium composed of a single, homogeneous, though some-
times layered, felt-like membrane, which is at first continuous over
the mouth, forming a sort of epiphragm ; sporangioles more nu-
merous than in the preceding genus, and the funiculus is much less
complex, consisting of a little bundle of elastic filaments gathered
closely together in a very thin, hardly noticeable outer covering, the
upper end of which is attached to a nipple-like protuberance on
the under surface of the sporangiole, and the lower end is attached
to the inner wall of the peridium ; there are no filaments intermixed
with the spores.

Although the genus Cnicibtilum \Nd.s not founded until 1844,
for nearly a century and a half plants belonging to it have been
known and unmistakably figured. Ray* in 1696, in a list of
plants collected by D. Samuel Doody, mentions " Fungus seminifer
minor," which he figured in 1724.! Micheli, five years later, gave
a good figure of Crucibuluui cnicibulifornie. As early as 1697
Boccone % figured this plant, under the name of " Fungus ar.epiiaxtac,
calyculatus," but does not seem to give it more than this passing
notice. Gleditsch,§ Schaeffer,]] Scopoli,! Hudson,** Hoffmanft
and others made mention of this plant, under various names, but

*Syn. 333. 1696. [Ed. 2.]

t Ray, Syn. 20. //. /./. 2 b, c. 1724. [Ed. 3.]

% Mus. Fis. //. 301. f. I. 1697.

gMeth. Fung. 138..//. 4. 1753.

II Fung. Bav. et Palat. Icon. 2 : pi. i-jg. 1761 ; 4 : 125. 1774.

\y\ox. Carn. 2: 486. 1772. [Ed. 2.]

**Flor. Ang. 2 : 634. 1778. [Ed. 2.]

ft Vegt. Crypt, fasc. 2: 29.//. 8. f. i. 1790.

White : The Nidulariaceae of North America 269

not until 1801, was the present generic name employed by Per-
soon * — even then only to denote a species — Cyatlms Crucibulum.

Schweinitz was the first to report Crucibulum crucibuliformc from
this country under the name of Nidularia Crucibulum. He also
described another species, Nidularia juglandicola which though
retained as a good species of Crucilmlum by De Toni,t proves to
be a form of Crucibulum crucibuliforme, slightly larger and more
flaring, but otherwise similar to the type species. Though the
genus as represented by Crucibidum crucibuliforme is comparatively
common and widely distributed, it is interesting to note in contrast
with the genus Cyathia, which is very prolific of species, that only
one other species has been described, viz., Crucibulum simile
Massee % from New Zealand and Australia, and even this is said
to resemble Crucibtdutn crucibidiforme closely.

Bulliard § says that there are two varieties of Crucibulum
both of which he figures, one glabrous, almost as smooth inside
as out, the other smooth inside but somewhat tomentose on
the outside, and always larger and more yellow than the first
It has been noticed in examining quite a number of specimens
from different localities that those collected in one place often vary
greatly in size and color, and in the nature of the tomentose outer
surface of the peridium. Cooke, || in 1 879, described a variety which
he calls lanosum, and which De Toni says might possibly be Bui-
Hard's second variety (figured on//. 4.0) but as Cooke described it as
"paler than in the usual form" this disposition cannot be correct.

I. Crucibulum crucibuliforme (Scop.)
Peziza sessilis campanulata (b) Gled. Meth. Fung. 138.//. 4.

1753-

Peziza secunda Schaeff, Fung. Bav. et. Palat. Icon. 2 : //. //p.

1763-

Peziza lentifera Oeder, Fl. Dan. 2 : 9. //. /05. 1763 ; not

Linnaeus.

Peziza crucibidiformis Scop. Flor. Carn. 2 : 486. 1772 [Ed.

2]; Schaeff. Fung. Bav. et Palat. 4: 125. 1774.

*Syn. Meth. Fung. 239. 1801.
fSaccardo, Syll. Fung. 7 : 44. 1888.
JGrevillea, ig : 93. 1891.
^ Champ. I : 165. pi. 40 and//. 4S8. 1809.
II Grevillea, 8 : 58. 1879.

270 White : The Nidulariaceae of North America

Pezizalaevis Huds. Fl. Ang. 634. 1778. [Ed. 2.]

Cynt/iHs cjlindnais YJ'xWd. Fl. Berol. 399. 1787.

Cyathus cnicibidiformis Hoffm. Veg. Crypt, fasc. 2, 29. pi. 8.
f. I. 1790.

Nididaria lacvis Bull. Champ, i: 164. //. 4^8. f. 2. pi. 40.
1791 ; With. Bot. Arrang. 3: 446. 1792 [Ed. 2] ; Sibth. Fl.
Ox. 393. 1794.

Cyathus Crucibulum Pers. Syn. Fung. 238. 1801 ; Nees, Syst.
1^0. pi. ij.f. 133. 18 17.

Cyathus laevis DC. Fl. Fr. 2 : 269. 1805.

Nididaria Crucibulum SQcret Mycogr. Suisse, 3: 378. 1833;
Schwein. Trans. Am. Phil. Soc. 4: 253. 1834.

Nididaria jiiglandicola Schwein. Trans. Am. Phil. Soc. 4 : 253,

1834.

Crucibidum vidgare Tul. Ann. Sci. Nat. III. i: go.pl. 6.f.

g-24. pi. 7.f. I. pi. 8.f. 13-17. 1844.

Crucibulum juglandicolum De Toni ; Sacc. Syll. Fung. 7: 44.
1888.

Peridia 5-1 o mm. high, 5-10 mm. across at the top, 4-7 mm.
at the base, cylindrical-campanulate, base truncate or but slightly
contracted ; outer surface dirty cinnamon or grayish fawn-color,
young plant minutely velvety tomentose, becoming smoother with
age ; the inner surface smooth, shiny and whitish ; mouth en-
tire, firm and even, sometimes slightly contracted, sporangioles
pale ochraceous, becoming whitish, 1.5-2 mm. in diameter, very
numerous and crowded, flattened beneath ; spores hyaline, elliptic,
Z-\o IX long, 4-6// wide, smooth, some thick-walled. {PI. 16. f.
10-13 ; pi. 18. f. 7-13 and 16.)

Plants gregarious or single on twigs, old bagging, chips, etc.;
common and variable.

Exsic: Ellis, N. A. Fungi, no. 728; Ravenel, Fungi Amer.
Exsic. no. 139.

New York: Underzvood, Clinton, Jclliffe ; Maine: Harvey;
Connecticut: Underivood, White; Massachusetts: Underwood,
Underwood & Seymour, g 4.1 ; New ]ersey : Bllis, White; Pennsyl-
vania : Haines, Everhart ; Alabama : Underivood, Earle ; Virginia :
Underwood ; Kentucky : Underwood ; Indiana, Underivood ; Kan-
sas: Cragin, 4go ; Oregon: Carpenter ; Montana: Reynolds ; Qq-l-
OKXHo: Bethel, Underivood & Selby ; Washington: Suksdoif.

White: The Nidulariaceae of North America 271

The followincr is extracted and translated from Tulasne's more
extended description of this species : The inner white evanescent
pelHcle is a remnant of the mucilaginous matter which fills the cup
in its early stages. The sporangioles are accumulated in large
numbers in a space apparently too small to contain them. They
are covered with a thick fibrous coat which can be peeled off, and
when deprived of this the sporangiole is black and of a horny con-
sistency. It is composed of two parts, a thick and much inter-
woven darker outer part, and an inner hyaline portion com-
posed of irregularly shaped filaments taking the place of basidia
and forming a hymenium of which the surface is nearly uniform.
The rest of the inner substance is composed of the spores
without any filaments, which fact distinguishes these sporangioles
from some of those belonging to species of Cyathia. The funic-
ulus of Cnicibulum is much more simple than that of Cyathia
hirsuta, being composed of a little bunch of elastic filaments
gathered closely together in a very thin, transparent outer cover-
ing, which is hardly noticeable, and of an equal filet, about 2-3
mm. long, which expands itself at the base into the inner wall of

the peridium.

III. Nidula gen. nov.

Peridium composed of a single homogeneous, but layered mem-
brane which is at first continuous over the mouth much as in
Crucilnihivi ; sporangioles very numerous, at first immersed in a
glutinous substance, very closely packed, entirely filling the central
cavity and in no way attached to the peridium wall ; no filaments
intermixed with the spores.

Peridium shaggy-tomentose; sporangioles light-colored, 2 ram. wide.

1. N. Candida.

Peridium minutely tomentose ; sporangioles dark-colored, barely I mm. wide.

2. iV'. microcarfa.

I . Nidula Candida (Peck)

Nididaria Candida Peck, Reg. Rep. 45: 24. 1891.

Peridia 6-15 mm. high, 6-15 mm. wide at the top, 5-10 mm.
at the base, cylindrically cup-shaped, somewhat truncate at the
base ; outer surface white, becoming dingy with age, thick, felt-
like, shaggy-tomentose ; inner surface smooth, at first snowy white,
becoming brownish with age, somewhat shiny ; mouth entire, firm,
somewhat spreading but not recurved; sporangioles 1.5-2 mm. in
diameter, light grayish fawn-colored, very thin, flattened, both sur-

272 White : The Nidulariaceae of North America

faces perfectly even, angular, but not depressed; spores 6-10 /i
long, 4-8 n wide, globose to elliptic, hyaline, somewhat granular,
thin- walled. {PL 16. f. 17-20.)

Growing singly on twigs, moss, etc.

Washington : Olympia, Henderson, Suksdorf.

British Columbia : Macoun, loy.

2. Nidula microcarpa Peck, sp. nov.

Peridia 4-6 mm. high, 4-5 mm. wide at the top, 3-5 at the
base, subcylindrical ; outer surface whitish, or pale grayish-yellow,
minutely tomentose, firm ; inner surface smooth, with a thin
brownish shiny layer formed by the drying of the glutinous inner
substance ; mouth entire, straight and firm ; sporangioles very
numerous, lenticular, covered with a thick fibrous outer coat, which
peels off, as in Cntcibidum crucibnlifonne, when dry rugosely
wrinkled, becoming smooth when moistened, reddish-brown, .5-1
mm. wide ; spores 6-9 ijl long, 4-6 // wide, broadly elliptic or sub-
globose, hyaline, thick-walled. {PL 17. f. 1-6 ; pL 18. f. 14, 13,

Growing on wood, and on the ground.

California : W. R. Dudley.

Montana : Columbia Falls, R. S. Williams.

2a. Nidula microcarpa rugispora var. nov.

Cnicibidiini riigispovnm E. & E. in herb.

Peridia 3-5 mm. high, 3-6 mm. wide at the top ; outer surface
yellowish-white, w^ith a rather thin close tomentum ; inner surface
smooth, whitish or brownish, somewhat shiny ; mouth recurved,
and spreading, and minutely lacerate, fimbriate ; sporangioles very
numerous, reddish-browai, rugose when dry, barely I mm. w^ide,
flattened, subrotund ; spores broadly elliptic, hyaline, slightly
thick-walled, 6-9 /i long, 4-7 n wide, slightly larger than in the
foregoing species. {PL ij.f. 7-9-)

Growing on dead twigs.

Washington : Skamania county, alt. 3,000 ft., Suksdorf, 510.

Though this variety closely resembles the species it differs in
the mouth characters and in the somewhat larger spores.

IV. GRANULARIA Roth, Ust. Ann. Bot. i : 6. pi. i.f. i. 1791
Nidularia Fries & Nord. Symb. Gaster. 2. 181 7-1 8.
Peridium composed of a single homogeneous rather thin closed

membrane, or a thin, closely interwoven layer of branched fila-

White : The Nidulakiaceae of North America 273

ments, opening more or less irregularly by the breaking away or
falling to pieces of the walls at maturity, having no true epi-
phragm : sporangioles enveloped in mucus and not attached in any
way to the inner surface of the peridium.

The history of the genus Gramdaria, though not as long as
that of the first two genera belonging to this family, appears to have
become more involved and complicated, very probably owing to
the fact that the species of Graiiiilaria are as a rule, rarer, and less
conspicuous than the species of Cyathia and Cmcibulum. Micheli *
in 1729 figured " Cyathoides sciitcllatiim^ which is unquestionably
a true Gramdaria, and the reason why this species has since been
placed under Cyadius by Roth t and later by Tulasne, it hard
to understand. Micheli, in writing of this genus, says " [fructus]
vel prope centrum, vel ad circumferentiam brevissimo pediculo
seu umbilicali funicolo firmantur," but of the figure of three spor-
angioles of " Cyadioidcs scutedatian" only one has a short lateral
attachment, and that much shorter than in the figures of the spor-
angioles of the three other species represented. Fries J says of
his second division, which contains true Grannlaria, that the spor-
angioles have no umbilicus or umbilical thread, but are attached by
the margin. Tulasne writes that notwithstanding these authorities,
the lateral position of a funiculus seems very problematical in the
genus Cyathia, and that the sporangioles of the specimens of Gran-
ularia which he has studied do not adhere any more by the edge
than by any other point, either to the mucilage in which they are
immersed or to the peridium. This idea must have arisen from
the fact that the mucus contiguous to the walls of the peridium
dries up a little quicker than that in the center of the cup and for
this reason, several sporangioles appear to be fixed by their outer
margin, while they are still loose in the center of the cup. But
this does not explain why " scutellaris " has been made a species
of Cyadua.

No other writer seems to have mentioned this plant after
MicheH until 1 791, when Roth described a new genus Gramdaria,
as follows : " Fungus subrotundus, granis mucilagine immersis

*Nov. PI. Gen. 222. pi. 102. f. 4. 1729.
t Roth, Cat. Bot. I : 237. 1797.
J Syst. Myc. 2 : 300. 1822.

274 White: The Nidulariaceae of North America

farctus," with one species Granitlaria pisiformis, which he fully
describes and figures. This is unquestionably a Nidularia, as
this genus was called until Otto Kuntze restored Gramilaria to its
proper place in 1891. Roth* himself described and figured two
species six years later under the name of Cyatlius farctus, and
Cyatlins scutcllaris ; it is possible that these belong to our genus
NiJula, but with the imperfect descriptions it is difficult to deter-
mine. It is very evident that they are not species of Cyatliia.

Bulliard,t in 1780 made the first mention of the name Nidu-
laria, but he referred it to figures which have been unquestionably
proved to be CyatJna Iciitifcra and Criicibuhnn criicibidiforme.
Sibthorp % also employed this name for Cyathia kntifera, Cyat/iia
hirsni a dind Criicibuhuii cnicibidiforinc. In 1817 NeesJ described
and figured Nidularia granulifcra, but although he gives a colored
plate, it is difficult to form a correct idea of his plant. Two years
later Ehrenberg || described and figured Nidularia globosa which
he says resembles Cyathus farctus Roth and Cyatlius deformis
Willd.^ but agrees with neither. About this time, 1 8 1 8, Schweinitz
described from North Carolina Cvathus pidviiiatus, which, though
unfortunately none of the original material remains except some
of the wood on which it grew, is evidently a true Grauularia and is
a fairly common species in our own country, and the only clear and
definite one of the whole tangled thread of the present history.

The family Nidulariaceae was established by Fries in 1822.
Under the section ''Nidularia,'' he gives seven species, five of
which are the above-quoted ones of Nees, Schweinitz, Roth, Ehren-
berg and Willdenow ; the last one, Nidularia denudata Fr. &
Nordh. ** appears from the description to be a true Grauularia,
while the third species which he gives as Nidularia farcta he refers
to Ray's ft "O- 21, which is the first mention of Crucibidum cruci-
bidiforme.

*Cat. Bot. I : ly]. pi. 7. f. 2. 1797.

t Herb. Fr. //. 4SS.

JFlor. Ox. 393. 1794.

^. Syst. Pilze. 139 note. //. 13. f- 133c. 1S17.

II Sylv. Myc. Berol. 16, 2S./. S. 1819.

\ Ust. Bot. Mag. 2 : 14. p!. 3. f. S. 1788.

** Symb. Gaster. 4. 1817-18.

ft Syn. 20. />/. /. / 2.

White : The Nidulariaceae of North America 275

Fischer * in writing of Nidularia says that there are sixteen
known species, but though De Toni f gives eighteen from all parts
of the world, it is more than probable that several of these are
either synonyms of other species or belong to other genera.
Tulasne gives eleven species, to nine of which he adds " non
vidimus" ! the remaining two being new species — Nididaria aus-
tralis from Chili, South America, and Nidularia Diiriacana from
LaCalle, Algeria (Mauritania), From America several additional
species have been described, namely, Nididaria Alabauiensis Atk. %
from Alabama, which we have reduced to Granularia pidvinata,
and Nidularia rubella E. & E. § from New Jersey. This last
species is very uncertain, in fact it may not belong to this family at
all, as the sporangioles are quite different in texture from those of all
the other species, and have never been found to contain any spores,
being filled with small pieces of amorphous hyaline matter. It is
hard to believe that the plants are in an immature condition as the
peridial walls have in some specimens almost entirely disappeared.
The only remaining species of Nididaria which has been described
in this country is Nidularia Candida Peck which has been referred
to the preceding genus. Two heretofore unpublished species of
Granularia complete our present list : Granularia castanea, a
herbarium species of Ellis & Everhart, under the name of Nidularia
castanea, from New Jersey found in 1883, and Granularia rudis
Peck, from California.

Synopsis of the Species of Granularia

Sporangioles numerous, small (l mm. or less in diameter).

Peridium tubercular, pulverulent. I. G. pulvinata.

Peridium only slightly tubercular, not pulverulent. 2. G. castanea.

Sporangioles few, larger (2 mm. or more in diameter). 3. G. rudis.

I. Granularia pulvinata (Schwein.) Kuntze, Rev. Gen. PI.

2:855. 1891
Cjathus pidvinatus Sch-wtm. Fung. Car. Sup. 51. 18 18.
Nidularia pulvinata Fries, Syst. Myc. 2 : 301. 1822 ; Schwein.
Trans. Am. Phil. Soc. 4: 253. 1834.

*Engler & Prantl. Nat. Pflanzenfara. i^** : 326. 1899.

t Saccardo. Syll. Fung. 7 : 28. 1888 ;— 9 : 265. 1891 ;— 11 : 156. 1895 ;— 14 :
256. 1899.

I Bull. Cornell Univ. (Science) 3 : 23. 1897.
I Bull. Torrey Club, 11 : 18. 1884.

276 White : The Nidulariaceae of North America

Nidiilaria Alabamcnsis Atk. Bull. Cornell Univ. (Science) 3:23
1897.

Peridia subspherical, sessile, 2-10 mm. wide, 2-9 mm. high,
reddish-brown or dirty cinnamon-colored, at first very floccose-pul-
verulent, gradually becoming smoother with age, dehiscing irregu-
larly or not at all ; peridium thin and rather brittle, tuberculose ;
inner surface is shiny, smooth, brownish ; sporangioles very numer-
ous, darkish brown, barely i mm. in diameter, somewhat angular
and depressed; spores hyaline, thick-walled, 6-10 « long, 4-7 ix
wide. {PI. 17. f. 10-16, 20; PL 18. f. ig.)

Plants gregarious, rarely singly on wood.

New York : Ellis, Faivman, Kiipfer ; Louisiana: Langlois,
2666 and 182 1 ; Connecticut: White; Maine: Harvey; Ala-
bama ; Auburn, Atkinson.

The floccose pulverulent appearance of the surface of the peri-
dium is caused by the innumerable ends of the filaments which
compose the peridium protruding above the main structure, soon
becoming broken or eroded. This is a very characteristic feature
which renders this species easily recognizable.

Atkinson (/. c.) describes Nidularia Alabamensis, which un-
doubtedly belongs here substantially as follows : Peridia spherical,
sessile, reddish-brown, roughened, 4-5 mm. in diameter, irregularly
dehiscing by the breaking into fragments of the upper portion ex-
posing the sporangioles which completely fill it; no "rooting"
threads ; sporangioles lenticular, shiny, dark or blackish-brown,
barely i mm. in diameter, corrugated, hard, filled with a whitish
pulpy material which is composed of stout irregular flexous or
blanched knotty strongly tuberculose threads with which the
spores are mixed ; the spores are obovately hyaline, 4-6 x 3-4 /^.
On decaying wood.
Alabama : Auburn, July, 1890.

2. Granularia castanea (Ell. St Ev.) sp. no v.

Nidularia castanea Ellis & Everhart in herb.

Peridia small, elongate-globose, or subspherical, sessile, some-
times confluent, .5-2 mm. long, 0.5-1 mm. high ; peridial walls
very thin, and brittle at first, pale yellowish fawn-colored, grad-
ually as the plant matures becoming grayish, and somewhat tuber-
culose from the inner pressure of the sporangioles, dehiscing
irregularly, the outer covering often breaking away completely,

White : The Nidulariaceae of North America 277

leaving the sporangioles seated alone on the wood, barely visible
without magnification ; sporangioles yellowish-brown, very numer-
ous, crowded, circular, flattened above and below, barely 0.25 mm.
in diameter, even or slightly creased under high magnification ;
spores subglobose, hyaline, thick-walled, 4-7 ix long, 3-6 jj. wide.
{PL 17. f. 17-19; pi. iS.f. 20.)

Gregarious on wood.

New Jersey: Newfield, July, 1883. Ellis.

3. Granularia rudis Peck sp. nov.

Peridia hemispherical or subglobose, sessile ; outer covering
thin, smooth, fragile when dry, cellular, rather gelatinous when
moist, yellowish-brown with a faint greenish tint ; indehiscent or
the peridial walls gradually breaking away ; sporangioles few, irreg-
ular, angular or compressed, 2-2.3 ^^- i" diameter, blackish or
greenish tinted ; spores broadly elliptic or subglobose, hyaline,
7-9 ijL long, 6-8 fji broad. {PL 17./. 21-2J.)

On decaying wood.

California : W. R. Dudley.

The material from which this species is described is very scanty,
but the specific features are so individual, and they differ so much
from any known species of Granularia that it has been decided to
include it in the present revision. It differs from all known species
in the gelatinous cellular and greenish texture of the peridium, and
in the large angular sporangioles, only a few being contained in
each peridium.

Species inquirenda

NiDULARiA rubella Ell. & Everh. Bull. Torrey Club, 11 :
18. 1884.

The following table will show the present known distribution
of the family in North America :

278 White : The Nidulariaceae of North America

•ooixaj^j

•BOIBUIBf 1

+

+

•XI0J3 -js

+

■oDi^ ojjan<j 1

+

+

■Bqnn

+

++++

++

+

■Biqiuniof) qsi^ug |

+

•BIUJOJilB^ 1

+

+

+

■uoSaJQ

+

•aojSuiqsBAV 1

+

+++

■

•BUBiUOJ^

+

+

+

•q^jri

+

•OpBJOI03

++

++

+

•00!X3I\[ AV3^ 1

+

•sBxax

+

•SBSUB-^

+

++

+

■•B5tSBjq3>I

+++

•BAVOJ

+ +

•UISU03SI^\\

•X5{on5U3^

+

•BUBipUI

-+ +

•OHO

+•+

•BUBismoq

••BUiEqBiv

+

•eiSjo3Q

+

••EU110JB3 qjnog

+

■BlUl3jly\^

+

+

■3JEA\-B[3Q;

+

■B!UBA]XSUU3J

+ +

+ +-

+

•XsSJPf M3^

+ +

•5[JOj^ AV3^

+

++ 4-

•jnoiioauuoQ

+

•SJ)3Snq3BSSBJ\[

•SUtBJ^

-+ +

■BpBUBf)

+ +

+ + + + .

+

+

+ + + + +

+ +

+

ulifor
ata.

:S K

~ -S <u 5; '*^ -^ K

ii.

n cruc

idida.

arpa.

a pulvt

a.

•§ -S -^ -S i ^ ^ <^ ^

1

IVriqht
icibulut
I Ilia cat

mi croc
anulari
casiane
rudis.

§.

G'G U U 'o ^ 'o 'o ^O to U ^

o'

U G ^ ^ ^ ^ ^

White : The Nidulariaceae of North America 279

Rxplanation of nates

The figures on Plates 14, 15, 16 and 17 were drawn one quarter larger than
indicated. Those on Plate 18 were made twice the size. The microscopic details were
drawn with a camera lucida and reduced on the same scale, as above noted.

Pl.\te 14
Fig. I. Cyathia Poeppigii, nat. size. 2. The same, enlarged about three times
after moistening. 3. Single peridium in dry condition. 4. Spores, X 335- 5- Cyathia
Berkleyma, nat. size. 6. Peridium of same, enlarged three times. 7. Spores, X 335-
8. Cyathia intermedia, nat. size. 9. Peridia, enlarged twice, one in dry condition,
the other after expanding when moistened. 10. Spores, X 335- ^^- Cyathia hirsuta
nat. size. 12. Group, enlarged about three times. 13, Spores, X 335- ^4) ^5-
Cyathia hirsuta inftindibuliformis, enlarged about three times. 16. Cyathia dura,
nat. size. 17, 18. Peridia, enlarged twice. 19. Spores, X 335- 20. Cyathia Mon-
tagnei, nat. size. 21. Peridium, enlarged twice. 22. Spores, X 335-

Plate 15
Fig. I. Cyathia tnelanospernta, nat. size. 2. Single peridium, nat. size (after
moistening). 3. Single peridium, enlarged about three times. 4. Spores, X 335- 5-
Cyathia pallida, nat. size. 6. Mature and young peridia, enlarged twice. 7.
Spores, X 335- 8. Cyathia lentifera, nat. size. 9, 10, 11, 12. Different specimens
showing character of growth and shape, enlarged twice. 13. Spores, X 335- '4
Cyathia rujipes, nat. size. 15. Peridia, enlarged twice. 16. Spores, X 335-
17- Cyathia stercorea, nat. size. 18, 19. Different specimens showing character of
growth and shape, enlarged about three times. 20. Spores, X 335-

Plate 16

Figs, i, 2. Cyathia stercorea, showing additional types and characteristics, en-
larged twice. 3. Cyathia rugispcrma, natural size. 4. Peridia enlarged twice. 5.
Spores, X 335- 6. Spores, X 5^5 ■ 7- Cyathia Wrightii, natural size. 8,9. Peridia,
enlarged twice. 10. Crucibulum crucibuli/orme,x\2ii\xra\s\zt. 1 1. Section of sporangiole,
X 50- 12. Group of peridia, enlarged twice. 13. Spores, X 335- I4- Group of
peridia from the Schweinitz specimens of Crucibidiun juglandicolum. 15. Spores
from the same, X 335- ^6. Cyathia (?) sp., natural size. 17. N^idula Candida,
natural size. 18, 19. Peridia, somewhat enlarged. 20. Spores and filament, X33S-
21, 22. Spores of Cyathia (?) sp., X 335-

Plate 17

Fig. I. Nidida microcarpa, nat. size. 2. Peridia, enlarged twice. 3. Section-
enlarged about five times. 4. Sporangiole showing torn outer covering, enlarged about
twenty-five times. 5. Sporangiole still in its enter covering, enlarged twelve times. 6.
Spores, X 335- 7- Nidula microcarpa rugispora,w2i\.i\zt.. 8. Peridium, enlarged three
times. 9. Spores, X 335- ^O- Gramdaria piilvinata, nat. size, reduced from draw,
ing by Mr. Hasselbring of iVidularia A laba mensis Ai'k.. 11. Gramdaria pidvinata,
nat. size. 12. Peridia, enlarged three times. 13. Spores and filaments, X 335- '^^•
Very old peridia, the walls of which have almost completely fallen away. 15. Spores.
X 585. 16. Spores, X 335- ^7- Gramd iria castanea,n2ii.sizQ. 18. Group of peri-
dia, enlarged twice. 19. Irregular peridia, having grown together, enlarged twice,
20. Spores of Gramdaria pulvinata, yC, t^t,^ [Nidtdaria Alahaniensis Atk. ). 21.
. (7/'a««/«r?rt rwaVj, enlarged twice. 22. Spores, X 335- 23. Sporangioles, X ^S-

280 White : The Nidulariaceae of North America

Plate i8
Figs. 1-6. Cyathia hirsuta. I. Sporangiole, showing attachment, enlarged about
fifteen times. 2. Spore, X 39°- 3- Spores and filaments, X 215. 4. Threads of
outer coating of sporangiole, X 215. 5. Nodose filaments of the funiculus, X 390.
6. Portionof the funiculus, X I lO- 7-^3- Cnuibulum criicibuliforme. 7. Sporangiole,
X 15- 8- Sporangiole, with the outer coat peeling off". 9. Small portion of a section
of a sporangiole, X 215. 10, 1 1. Spores of young specimen, X 39°- *2. Spores
of fully mature specimen, showing contraction of inner substance, X 390- ^3- Small
portion of the ends of the filaments of the thin membrane covering the mouth of the
immature specimen, X Z^S- 14- Spores of Nidula microcarpa, X 390. 15. Lateral
portion of outer coat of the sporangioles of same, X 215. 16. Small portion of the
outer coat of the sporangiole of Crucibulum crucibtiliforme, X 215. 1 7. Section of a
sporangiole oi Nidula microcarpa, X ^^O- ^8. Fascicular hairs from the outer cover-
ing of the peridium of Cyathia intertiiedia, X ^S- ^9- Small portion of the peridial
wall of Granidaria pulvinata, X 215. 20. Sporangioles of Granularia castanea,
when devoid of peridial covering, X ^ ^^-

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CONTRIBUTIONS FROM THE NEW YORK BOTANICAL

GARDEN— No. 25

NOTES ON AMERICAN HEPATICAE

BY MARSHALL A. HOWE.

NEW YOEK
1902

[Reprinted from Bulletin Torrey Botanical Club, 29 : 281-289, May 26, 1902.]

Notes on American Hepaticae

By Marshall A. Howe

I. Cephalozia CONNIVENS

Cephalozia CONNIVENS (Dicks.) Lindb. Proc. Linn. Soc. 13 :

190. 1872. Kritisk Gransk. Mossorna Dill. Hist. Muse. 38.

1883. Spruce, On Cephalozia, 46. 1882. Lindb. & Arn. Sv.

Vet.-Akad. Handl. 23': 20. 1889. Kaalaas, Nyt Mag. 165.

1893-

Jmigennannia coniiivcns Dicks. Fasc. PI. Crypt. Brit. 4: 19.

//. //./. /J". 1 80 1.

Ceplialozin nmltiflora Lindb. Act. Soc. Sci. Fenn. 10 : 501.
1875. Muse. Scand. 4. 1879.

This species has been much confused with its near relative
Cephalozia lunulaefolia Dumort. {^C. media Lindb., C. vmltiflora
Spruce) both in Europe and America, but is very distinct as was
first well pointed out by Spruce {On Cephalozia, 39. 1882). The
plant may be readily distinguished from C hinulaefolia by the large
leaf-cells (3 5-90 y., while only 25-50 /i in C. hinulaefolia), by the
long-ciliate perianth-mouth, by the perianth- wall being unistratose
throughout, and by being autoicous. The specimens distributed
by Sullivant under the name oi Jungcrmaiinia connivens in Musci
Alleghanienses (no. 246) belong with C. liimilaefolia so far as we
have been able to see them, and the same is true of no. 57 of
Austin's Hepaticae Boreali-Americanae, issued as Cephalozia con-
nivens. In 1896, Professor Underwood, in his review of "The
Genus Cephalozia in North America" (Bull. Torrey Club, 23 :
381-394), placed Cephalozia connivens under "Species dubiae et
inquirendae," remarking that it was likely to occur northward. In
October, 1898, the writer of the present notes announced* at a
meeting of the Torrey Botanical Club the discovery of the genuine
Cephalozia connivens on the grounds of the New York Botanical
Garden, supposing this to be the first collection of the real C. con-
nivens in America. The species was soon found in several other
places in the neighborhood of New York, and this has led to an

*Bull. Torrey Club, 26 : 24. 1899.

281

282 Howe : Notes on American Hepaticae

examination of the specimens of Ccplialozia in the accessible her-
baria with the result that we are now able to cite seventeen North
American specimens of the true CcpJialozia connivcfts, showing a
coastwise range from Prince Edward Island to Florida, and an
extension inland as far as London, Ontario.

The specimens cited may be arranged as follows :
Prince Edward Island : /. Macoiin, June 27, 1888, no. 139 (a
little only, mixed with C. bimilaefolia and Kantia tricJiomanis). On-
tario: London,/. Dcarncss (in herb. Underwood, comm. in 1889
as no. i). Rhode Island : Providence, from Olney collection
in Chapman herbarium ; also, in same herbarium, a second speci-
men from Rhode Island, without locality. Connecticut : on the
ground in wet woods, Westville, A. W. Evans, November 29,
1890. New York: in a tamarack swamp, Syracuse, L. M.
Underivood, April, 1888 ; Staten Island, Mrs. E. G. Britton, Febru-
ary 14, 1892 ; on the top of a decayed hemlock stump near Bronx
River, New York Botanical Garden, M. A. Hozvc, September 21,
1898, with Odontoschisma spliagni ; on bank of a stream in a boggy
wood, Freeport, Long Island, M. A. Hoive, October 17, 1898; on
margin of a swamp, Richmond, Staten Island, M. A. Hozve, Octo-
ber 23, 1898 ; Sayville, Long Island, F. E. Lloyd, March 5, 1899.
Delaware : on ground in woods, Wilmington, A. Commons,
March 12, 1890; in herb. Columbia University. North Caro-
lina : Salem, Schzveiuitz, in herb. Torrey (labelled Jungermannia
bicornis by Schweinitz). South Carolina : Summerville, C. G.
Dubois, April, 1889. Florida: Lisbon, L. M. Underivood, Janu-
ary, 1 89 1 (distributed from U. S. National Herbarium as no.
1 217); Port Orange, F. C. Stranb, February 23, 1895 (no. 207
in herb. Underwood) — also in same locality, March, 1895, mixed
with Telaranea and a minute Lcpidozia in Hepaticae Americanae,
no. 1 80. Bermuda : Devonshire Marsh, M. A. Hozvc, July 4,
1900.

Lindberg appears to have been the first to have combined the
specific name r(^7///777/jr (Dicks.) with the generic name Cephalozia,
although at the outset he attributed the binomial to Dumortier.
Dumortier however persisted even in his " Hepaticae Europae " in
keeping the species under Blcpharostoma and seems never to have
written C'pli.d.K.ia conn'v:ns. The synonymy of the species is

Howe : Notes on American Hepaticae 283

confused, owing partly to the fact that is was not until 1881 and
1882 that Lindberg and Spruce began to distinguish, in their pub-
lished works at least, between the true Cephaloda connivms and
C. Inmdaefolia, and owing furthermore to some uncertainties at-
tending the determination of Jiingcrmannia multiflora of Hudson
(Fl. Angl. 431. 1762), The question of interpreting correctly
Hudson's /. multiflora, difficult at best, is needlessly complicated
by Dr. Spruce (/. c. 40) by quoting the 1778 edition of Hud-
son's Flora Anglica, where a supposed synonym from Linne's
Mantissa is introduced, though the original 1762 edition with-
out the Linnaean synonym is all that needs to be considered.
The original Jtingermannia multiflora seems to have been founded
wholly upon a plant described and figured by Dillenius (Hist.
Muse. 481. //. 6g. f. 4. 1 741). As noted already by Hooker,
Spruce, and Lindberg, the figures given by Dillenius point strongly
to Ccphalozia bicuspidata. It is quite impossible to believe that
they were drawn from specimens of C. connivens and Dillen's
words ''per letitem vero pcrangicsta" in describing the leaves
could not well apply to this species, yet Lindberg in 1875
(Act. Soc. Sci. Fenn. 10 : 501), after determining the specimen in
the Dillenean herbarium bearing the corresponding name and
number from the Historia Muscorum to be the Jungermannia con-
nivens'^ of Dickson, brought forward for this species Hudson's
name multflora. Finally,! however, interpreting this " Liche-
nastrum multiflorum exile foliis angustissimis " of Dillenius by the
figure and description, Lindberg considers it a synonym of
Ccphalozia bicuspidata (L.) Dumort., which seems to us the most
reasonable course in the matter. Lindberg at the same time most
unwarrantably interprets Jiingermannia multiflora Huds. in the
light of additions made by Linnaeus nine years after its original pub-
lication and shifts the specific name multiflora to Lepidozia setacea
(Web.) Mitt. It is clear that if Jungermannia midtiflora Huds.
(1762) can be justly treated as a synonym of/, bicuspidata L.
(1753) it should always remain a " dead name."

*This determination, it may be noted, was made several years before the distmc-
tions between the true C. connivens and C. lunulaefolia ( C. media Lindb. ) were rec-
ognized in print, at least, yet the statement that the Dillenian specimen is " C. conni-
vens (Dicks.) Lindb.," was repeated by Lindberg in 1883, two years after his publica-
tion of C. media.

I Kritisk Gransk. Mossorna Dill. Hist. Muse. 38. 1883.

284 Howe : Notes on American Hepaticae

II. Telaranea

Telaranea Spruce, Trans, and Proc. Bot. Soc. [Edinb.] 15:
365. 1885. [As a synonym.] Schiffn.; Engl. & Prantl, Nat.
Pflanzenfam. i'^ : 103. 1895.

The genus Telaranea has points of contact with Lcpidozia,
Blepharostoma, and Arachniopsis, and if generic rank is denied to
it, might be incorporated with any one of the three with almost
equal justice. To the Miero-Lepidozia section of the genus Lepi-
dozia its affinity is perhaps especially close, yet, according to pre-
vailing conceptions of genera among the Hepaticae, it seems a
rather violent proceeding to include it in a generic group the his-
torical type of which is Jiingermanida reptans L. Telaranea di-
verges from Lepidozia in its delicate, filmy, conferva-like habit and
in the division of the leaves very nearly or quite to the base, the
segments being never more highly connate than through half the
altitude of the basal cell and usually for a considerably less dis-
tance ; the leaf-segments are capillary, of a single row of cells
unless at the very base, the place of the basal cell being often oc-
cupied by a pair of cells side by side. From Blepharostoma, Tel-
aranea is probably best distinguished by the disparity in size between
leaves and underleaves and by the more pinnate branching. The
archegonia, though usually borne on short postical branches as in
Lepidozia and Cepkalozia, are sometimes terminal on the main stem
or lateral branches as in Blepharostovia.

In its filmy habit and delicacy of structure Telaranea is sug-
gestive of Arachniopsis Spruce, but the latter genus represents a
still further departure along the same lines from the Lepidozia
type. In Arachniopsis the branches are all postical in origin, the
underleaves are entirely wanting or are now and then to be recog-
nized only in a most rudimentary form, and the leaves never have
more than two segments which are wholly free at the base and
consist of a single series of cells throughout.

The species and varieties to be arranged under the genus are,
so far as we know them, as follows :

I. Telaranea nematodes (Gottsche)
Jiingcnnainra )n!natodLS Gottsche, Hepaticae Cubenses Wright-
ianae.

Howe : Notes on American Hepaticae 285

Cephalozia nematodes Aust. Bull. Torrey Club, 6 : 302, 1879.

Lepidozia chaetopliylla Spruce, Trans, and Proc. Bot. Soc.
[Edinb.] 15: 365. 1885.

Lepidozia nematodes Spruce, /. c. 366.

Telaranea chaetopliylla Spruce, " Mst. nov. gen." /. c. [as syno-
nym]. Schiffn. ; Engl. & Prantl, Nat. Pflanzenfam. i^ : 103. 1895.

Lepidozia chaetopliylla tenids Pearson, Christ. Vid.-Selsk. For-
handl. i886'^: 9. 1886. Evans, Bull. Torrey Club, 20: 308.
1893.

Blepharostoma nematodes Underw. Bull. Torrey Club, 23 : 383
(footnote). 1896.

Spruce in comparing his Lepidozia chaetopliylla with Lepidozia
nematodes (Gottsche) emphasizes the flaccidity of the latter as a
distinguishing character and this indeed appears to be about the
only difference of importance to be detected between Wright's
Cuban plant and those from South America distributed by Spruce
as Telaranea chaetophylla (Hepaticae Spruceanae : Amazonicae et
Andinae). Wright's specimens revive quite imperfectly on being
soaked out with water, but there is such a range of variability in
this respect in different parts of a single tuft and even of different
parts of a single plant, both in the Cuban and South American
specimens that we are not inclined to give much weight to this
character. We therefore agree with our friend Mr. Pearson (/. <;.)
in believing that the two forms represent a single species. We
would, however, differ from Pearson in retaming for this species
the oldest published name nematodes. The specific name nemoides
given earlier by Taylor to another plant now recognized as a Lepi-
dozia seems sufficiently different both to the eye and ear to prevent
confusion.*

In both the Cuban and South American plants the archegonia
are usually borne on a short postical branch, rarely at the end of
the main axis or of an elongated branch. In 1886, Pearson (/. f.)
identified a specimen from Natal, South Africa, with Gottsche's
Cuban species, which he renamed Lepidozia chaetophylla tenuis.

*The remark is attributed to Spruce (Pearson, /. c.') that nemoides and nematodes
differ only in case-ending. It seems to us that the two words are equally nominative
in form but that Taylor's name nemoides is an etymological monstrosity on account of
his failure to use the true stem of the Greek noun in constructing the adjective.

286 Howe : Notes on American Hepaticae

Through the courtesy of Mr. Pearson we have been able to exam-
ine this South African plant, and like him, can find no reasonable
grounds for distinguishing it from the American specimens alluded
to. Telaranea nematodes, then, may be said to be found in a fairly
typical condition in South America and South Africa as well as in
Cuba, and to this range is now to be added Bermuda, where sterile
plants were collected by the writer in Devonshire Marsh, July 4,
1900, growing in company with CepJialozia comiivens 2iX\di C. divar-
icata. In addition to this more or less typical form, illustrated by
the specimens and descriptions cited, the species presents itself also
in two forms which we think are sufficiently marked to receive
varietal names. These are :

I a. Telaranea nematodes Antillanum (Besch. & Spruce)

Blepharostoma Antillamwi Besch. & Spruce, Bull. Soc. Bot.
France, 36 : clxxxiii. 1889.

Archegonia terminal on main stem, a lateral branch, or elong-
ated postical branch, rarely on a short postical branch ; leaves and
perianth rather rigid.

Le Gommier, Gaudeloupe, Ed. Marie. In a specimen of the
Guadeloupe plant kindly communicated by M. Bescherelle we find
on a single individual all the various modes of bearing archegonia
described above. The leaves, described by the authors as 4-parted,
we find much more frequently 3- and 2 -parted. It seems as im-
possible to separate this plant specifically from Lepidozia chaeto-
phylla Spruce as it is to distinguish satisfactorily between the latter
and Gottsche's Jiingennannia nematodes. So far as we know, the
identification of Blepharostoma Antillanum with Jungermannia
nematodes was first made by Professor Schiffner (Engl. & Prantl,
Nat. Pflanzenfam. I^ : 105. 1895).

lb. Telaranea nematodes longifolia var. no v.

Leaves more rigid than in type and more widely spaced, 0.4-
0.8 (rarely 0.9 mm.) long, leaf-cells 2-4^ times as long as broad.

Collected by the writer on humus in a swampy wood in
company with Sphagmim, Pallavicinia Lyellii, and Ceplialozia
catemdata, Freeport,* Long Island, New York, October 17, 1898

*This Freeport plant was listed by Dr. Smith Ely Jelliffe in\his Flora of Long
Island (48. 1899) under the name Blepharosfoma nematodes.

Howe : Notes on American Hepaticae 287

(type specimen in herb. N. Y. Botanical Garden). Also, Florida,
John Donnell Smith, 1877; C. F. Austin, March, 1878; F. C.
Stranb, March, 1895 ; Brunswick, Georgia, C. F. Austin, April,
1878.

The Long Island specimens agree essentially with those from
Georgia and Florida, though the latter make a slightly closer ap-
proach to the Cuban plant. The Long Island station is therefore
a noteworthy northward extension of the known range of a plant
which has heretofore figured in papers on North American Hepati-
cae as coming only from a limited region of the South. The arche-
gonia in these United States specimens, so far as we have observed,
always occur on a short postical branch. The leaves are 5-8 cells
long ; the underleaves are 3- or 2-parted, their prongs of 2 or 3
cells each, incurved at the apices. All are autoicous. In the orig-
inal Jungcnnannia nematodes, collected in Cuba by Wright, the
leaves are 0.25-0.5 mm. long, 4-6 cells high, the cells being 2-3
times longer than broad.

The variety longifolia often gives the impression of being two
or three times the size of the Cuban plant, but we have been unable
to find any reliable structural characters to serve for a specific
separation. The form of the perianth seems quite variable in all
conditions of the species, but is often broader in the var. longifolia
than in the type.

No. 180 Hep. Am., issued as Blepliarostoma nematodes, is ref-
erable to Telaranea nematodes longifolia ; it is mixed with Cepha-
lozia connivens and with a minute Lepidozia, probably a reduced
form of L. setacea.

Telaranea bicruris (Steph.)

Lepidozia bicruris Steph. Hedwigia, 24 : 166. //. j. 1885.

Brazil : Sao Francisco, Ule.

This plant, though a close relative of T. nematodes, seems en-
titled to specific distinction, differing in the almost invariably 2-
parted leaves, in the frequent continuation of the stems into leaf-
less flagella (rare in T. nematodes) and in the simpler 9 bracts.
It is described by Stephani as dioicous, though apparently he had
not seen $ plants. We have been unsuccessful in attempts to
find antheridia in the specimen kindly communicated by Herr
Stephani, and it certainly may be suspected that dioicism is to be

288 Howe : Notes on American Hepaticae

added to its distinguishing characters. All the perianths seen are
on very short postical branches.

III. Arachxiopsis

Arachniopsis diacantha (Mont.)

Jiingermannia dia^antha Mont. Ann, Sci. Nat. IV. 5 : 349.
1856.

Arachniopsis coactilis capillacea Spruce, On Cephalozia, 85.
1882; Trans, and Proc. Bot. Soc. [Edinb.] 15: 356. 1885.

Leaves mostly rigid, composed of 4-6 cells, these 4-7 times
as long as broad.

Peru : Weddell, Spruce.

Jungermannia diacantJia Mont, is represented in the Montagne
herbarium, now preserved in the Museum d'Histoire Naturelle of
Paris, only by two small sterile fragments fastened to pieces of
mica. This original material, which we have seen through the
courtesy of Mons. Hariot, agrees perfectly, so far as it goes, with
the specimen from Mt. Campana, Peruvian Andes, distributed by
Spruce as Arachniopsis capillacea in his " Hepaticae Spruceanae :
Amazonicae et Andinae." Montagne's description of the leaves
as "spiraliter 5/1 circa caulem dispositis " was evidently an error,
due, it may be, to a torsion of the stem.

Arachniopsis confervifolia (Gottsche)
Jiingermannia confervifolia Gottsche, Hepaticae Cubenses
Wrightianae.

Cephalozia cotfervifolia Aust. Bull. Torrey Club, 6 : 302.

1879.

Arachniopsis coactilis Spruce, On Cephalozia, 85. 1882. Trans.

and Proc. Bot. Soc. [Edinb.] 15: 355. //. 13. 1885. Exclud-
ing var. capillacea.

Leaves often flaccid, composed of 6-14 cells, these 2-3 times
as long as broad.

Cuba : Wright ; South America : on the tributaries of the
Amazon, Spruce. We have little hesitancy in considering Arach-
niopsis coactilis Spruce a synonym of the Cuban Jnngcrmannia con-
fervifolia Gottsche. The latter is more flaccid, showing a relation
to the South American specimens somewhat analogous to that

Howe : Notes on American Hepaticae 289

described above as existing between Jwigerntannia nematodes
Gottsche and Lepidozia chaetopJiylla Spruce. The Cuban plant is
autoicous and in structural characters is essentially identical with
the typical A, coactilis. Vestiges of underleaves are occasionally
met with in both and in both the leaf is sometimes reduced to a
single prong.

The specific separation of Aracliniopsis confervifolia from A.
■diacantha is possibly open to question, yet in the light of available
specimens the two appear distinct. Aracliniopsis coactilis filifolia
Spruce, judging from specimens distributed as Aracliniopsis filifolia
in Hepaticae Spruceanae, seems to us less entitled to specific rank.

Aracliniopsis confcruifolia resembles Telaranea nematodes in

outward appearance, but is easily distinguished by the characters

alluded to above under Telaranea. It is to be expected that

Aracliniopsis will be found to occur within the limits of the United

States.

IV. RicciA Campbelliana

Herr M. Heeg, of Vienna, has kindly called our attention
{in Hit) to the close resemblance between the Californian Riccia
-Campbelliana M. A. Howe (Mem. Torrey Club, 7 : 26. pi. gi.
f. i-ij. 1899) and the Mediterranean Riccia macrocarpa Levier
(Bull. Soc. Bot. Ital. 1894: 114. 1894). The similarity is indeed
very striking as is evident from specimens of R. macrocarpa which
we owe to Dr. Levier, but R. macrocarpa appears to be uniformly
dioicous as originally described by Levier and as described again
by Stephani (Bull. Herb. Boi'ss. 6: 343. 1898), while R. Camp-
belliana is uniformly monoicous ; and the areolae of the outer
face of the spores of Riccia macrocarpa are more perfect than in
R. Campbelliana. In view of these differences the claim of Riccia
Campbelliana to specific rank seems defensible for the present at
least. The species has recently been collected by Dr. Walter R.
Shaw at Claremont, Los Angeles County, Cal., thus extending its
known range about two hundred and fifty miles southward.

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