Annals
of the

Missouri Botanical

Garden

Volume XIV
1927

With Forty-one Plates and One Hundred and
Seventy-five Figures

Published quarterly at 8 West King Street, Lancaster, Pa., by the Board of Trustees
of the Missouri Botanical Garden, St. Louis , Мо.

Entered as — = — at the Post-Office n Lancaster, Pennsylvania,
r the act of March 3, 1879.

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions
from the Missouri Botanical Garden and the Graduate Labora-
tory of the Henry Shaw School of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden.

Information
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TABLE OF CONTENTS

Revision of the Genus Townsendia. . . Esther L. Larsen

Studies on South American Labiatae, III
Без, с zn Carl С. Epling

A Monograph of the Genus Pholiota. .L. О. Overholts
A Monograph of the Section Oreocarya

of Cryptantin:....<...2. ОК Edwin B. Payson
Cotton Wilt: A Pathological and Physiolog-

ical Investigation. ........ 1. à David C. Neal

Species of Cercospora on Smilax in the
United States. <r.. on ae e L. O. Overholts

PAGE
1—46

47-86
87-210

211-358

359-424

425-432
433-436

Annals
of the
Missouri Botanical Garden

Vol. 14 FEBRUARY, 1927 No. 1

A REVISION OF THE GENUS TOWNSENDIA!
ESTHER LOUISE LARSEN
Jessie R. Barr Research Fellow in the Henry Shaw School of Botany
of Washington University
HISTORY OF THE GENUS

The genus T'ownsendia was first described by William Jackson
Hooker in his ‘Flora Boreali-Americana’ іп 1834,2 It was named
in honor of David Townsend of Pennsylvania, an ardent student
of botany who contributed substantially to our knowledge of the
flora of his native state and especially to the genus Aster. Hook-
er, in establishing the genus T'ownsendia, recognized only one
species, T'ownsendia sericea, which he described in detail and
carefully illustrated.

omas Nuttall, in 1841, added four new species to the genus,
namely, Townsendia incana, T. spathulata, T. strigosa, and T.
grandiflora. The material from which Nuttall described these
species was collected by him “оп the Black Hills (or eastern chain
of the Rocky Mountains) near the banks of the Platte." During
the next four decades Т. Parryi Eaton,‘ Т. condensata Parry,

! An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University and sub-
mitted as a thesis in partial fulfilment of the requirements for the degree of Master
of Science in the Henry c School of Botany of Washington University.

? Hook. Fl. Bor. Am. 2: 834.

3 Nutt. Trans. Am. Phil. "n М. 8. 7: 304. 1841.

* Eaton, Am. Nat. 8: 212. 1874.

5 Parry, Am. Nat. 8: 213. 1874.

Issued June 8, 1927

ANN. Mo. Вот. Garp., Vor. 14, 1927 (1)

ГУ от. 14
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN

T. scapigera Eaton,* T. Rothrockit Gray; and T. Wilcoriana
Wood? were described by individual workers, thus making a total
of ten known species. In 1880 Dr. Asa Gray? published a sy-
nopsis of the group to which he added the following species as
new: T. florifer, T. Watsoni, T. arizonica, and T. glabella. This
synopsis formed the basis for Gray’s” treatment of the group in
the 'Synoptical Flora of North America, in which seventeen
species and four varieties of T'ownsendia were recognized. Since
1886 several species have been published and included in the
various manuals treating the flora of portions of the western
United States; but no revision of the group as а whole has ap-
peared since Dr. Gray's excellent treatment in the ‘Synoptical
Flora of North America.’

In 1894 Professor Thomas C. Porter revived Richardson's spe-
cific name exscapus and created the binomial T'ownsendia exscapa
(Richards.) Porter," a name which has been current in botan-
ical literature during the last thirty years.

Through the courtesy of Dr. A. W. Hill, Director of the Royal
Botanic Gardens, Kew, England, the writer has been privileged
to examine portions of the original material on which the genus
Townsendia was founded. А critical examination of this material
shows that the genus was based on two specifically distinct
elements, namely, specimens collected by Dr. John Richardson
at “Carlton House upon the Saskatchewan," a plant described
in 1823 in ‘‘Franklin’s Journey to the Polar Sea" as Aster
exscapus Richards." and specimens collected in the “Rocky
Mountains" by Thomas Drummond. These two plants differ
in the following important details: Richardson's specimen has
pubescent, linear-lanceolate flat leaves, which overtop the heads,
and the pappus of the ray-flower equals that of the disk-flower;
Drummond's specimen has more densely sericeous-pubescent sub-
terete leaves equalling but rarely exceeding the head, and the

* Eaton, Bot. King's Exp. 145. 1871.

1 Gray, Wheeler Rept. 6: 148. қ

* Wood, Bull. Torr. Bot. Club 6: 163. 1875.

? Gray, Proc. Am. Acad. 16: 82. 1880.

10 Gray, Syn. Fl. М. Am., ed. 2, 17: 166. 1886 and 1888.

и Porter, Mem. Torr. Bot. Club. 5: 321. 1894.

1 Richards. Frankl. Jour. Bot. App. 7, 748. 1823.

1927]
LARSEN— REVISION OF THE GENUS TOWNSENDIA 3

pappus of the ray-flower is much shorter than that of the disk-
flower. Тһе specimens collected by Drummond accord in every
detail with the description and illustration in Hooker's 'Flora
Boreali-Americana,' while those collected by Richardson present
several discrepancies when compared with Hooker's description
and illustration, particularly in the character of the leaf, pubescence
and pappus. Hence the Drummond plant is interpreted as the
type of Townsendia sericea Hooker, and this species therefore
must be taken as the type or standard species of the genus Town-
sendia. The Richardson plant, on the other hand, becomes the
type of Townsendia exscapa (Richards.) Porter.

GENERAL MORPHOLOGY

Roots.—Most members of the genus T'ownsendia develop rather
slender tap-roots. Тһе roots of some of the more caespitose
species, however, become very coarse and woody and more or less
branched. Тһе slender tap-root with a simple unbranched crown
is typical of the genus.

Stems.—The stem presents considerable variation in length and
in the extent to which it becomes branched. Townsendia Parryi
and T. formosa have simple, егесі, scapiform stems, while 7.
grandiflora, T. texensis, and T. strigosa have numerous ascending
branched stems. Тһе stem is usually herbaceous throughout its
entire length, but in some species, particularly those of xerophytic
regions, the basal portion becomes distinctly ligneous.

Leaves.— The leaf outline varies from linear-lanceolate to
obovate-spathulate. In T. Watsoni and T. Parryi both types
are present, the stem-leaves being linear-lanceolate and the basal
leaves obovate-spathulate. Тһе broader leaves are attenuated
at the base into а рейде, while the narrower ones are only ob-
scurely petiolate or sessile. Тһе range in the leaf size is from
0.5 to 6 em. in length and from 0.2 to 1.0 em. in width. Entire
leaf margins prevail throughout the genus. The surface is usually
pubescent, but sometimes it is glabrate, as in 7. glabella. In T.
spathulata, on the other hand, the leaves are villose-lanate. How-
ever, а strigose pubescence of closely appressed hairs as in 7.
eximia and T. strigosa is most prevalent in the genus.

[Vor. 14
4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Inflorescence.— The heads resemble those of the closely related
genus Aster. Тһеу are usually solitary and terminal and may be
borne on naked scapes or peduncles, or in the acaulescent forms
sessile among the rosulate leaves. In the branched species the
heads are occasionally disposed in clusters of two or three, but the
solitary condition is most characteristic.

Involucre.— The involuere is broadly campanulate and is com-
posed of two to six series of imbricated bracts. The outline of
the bract within the genus is quite variable, ranging from linear-
lanceolate to obovate, and from obtuse to acute or acuminate at
the apex. А lacerate-ciliate margin prevails throughout the
genus. Furthermore, in most species the involucral bracts also
have a membranaceous margin. Тһе character of the terminal
portion of the bract is important in the natural grouping of
species.

Pappus.—The plurisetose pappus consists of a single row of
rather coarse, slightly flattened bristles. Тһе pappus of the
ray-flower is somewhat shorter, or often reduced to a crown of
short squamellae. Тһе pappus in 7. formosa in both ray- and
disk-flowers is scarcely more than a vestige of the squamellate
crown. T'ownsendia eximia develops a coroniform pappus of
coalescent rigid squamellae sometimes bearing two or more
prolonged awns. Тһе condition in Т. eximia is analogous to that
of T. glabella where the short ray-pappus contains a few elongated
setae. The character of the pappus in some species is very con-
stant, while in others it is exceedingly variable.

Intermediate stages in the length of the ray-pappus from a
crown of short setae to a condition in which the setae equal those
of the disk-flower in length may be found in T. incana and T.
florifer. Тһе variability in the length of the pappus in certain
species, particularly in Т. sericea, has been emphasized by Gray,
Meehan,' and others. It is evident that the length of the ray-
pappus alone cannot be used in the differentiation of species.

Plate 5.

Corolla and Stamens.— The corolla of the disk-flower equals the
involucre in length while the ray is twice as long. "The color of
the ray varies within the genus from white through pink to

! Meehan, Nat. Flowers II. 1: 189. 1880.

1927]
LARSEN— REVISION OF THE GENUS TOWNSENDIA 5

purple but is never yellow. In both ray- and disk-flowers the
corolla is deciduous. The stamens are typical of the aster group.

Pistil.—The style appendages are lanceolate and have well-
developed stigmatic surfaces in both ray- and disk-flowers.
Nuttall referred to the ray-flowers of Т. strigosa and Т. grandiflora
as “infertile or neuter." А ray-flower from Nuttall’s type of
T. strigosa was found to have stigmatic surfaces exactly like those
of the disk-flowers, and the ovary contained a well-developed
ovule. Flowers were examined from all recorded species, and in
all cases the ray-flowers were found to be fertile. Moreover, in
those specimens which had reached maturity the ray-achenes were
well developed and appeared to be viable.

Achenes.—The achenes are ovate or oblong, much compressed,
and calloused margined, although those of the ray are sometimes
triangular. The hairs on the achene are bidentate or glochidiate-
tipped. These two types have the same morphological origin,
the bidentate forms being merely a forerunner of the glochidiate
type. The nature of these hairs is best seen under the low power
of the compound microscope. The type of pubescence is fairly
constant within the species and is important in the classification
of the group.

GEOGRAPHICAL DISTRIBUTION

The genus T'ownsendia is restricted in its distribution to the
western half of the North American continent. It extends from
western Manitoba and southern Alberta west to the Cascade
Mountains in Washington and Oregon, southward to the state of
Hidalgo in Mexico. Т. mexicana is endemic to Mexico. The
only other representative of the group occurring in that country
is Т. strigosa, a closely related species which extends from Wyo-
ming and Colorado through New Mexico and Arizona into ad-
jacent Mexico. The genus is best represented in Colorado where
ten out of the nineteen species recognized in this paper are found.
Material at hand would indicate that three of these are endemic
to that state. So far as known, Т. leptotes occurs only in the
Middle and South Park Region, Т. glabella in southwestern Colo-
rado, while Т. Rothrockii extends from the north-central part of
the state to the Uncompahgre Mountains. Two other species

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Townsendia. ff,

Townsendia exsca . s-e florifer
— — --- — Townsendia Rothrockii — 4 — + — Tounsendia grandiflora.
— — «e Townsendia елітіп. + + -+ -Townsendia incana.
seaccocecvesce Townsendia montana о о o T'ownsendia Parryi,
—x— x — x — Townsenilia sericea. 4 eee + moo 4 c Townsendia Watsoni.
———— же ір T'ownsendia strigosa. ннн Townsendia атіготіса
= — 41— 11—10 T'ownsetulia mertcana. овое ох — оо Townsendia texensis.
ownsendia scapigera. +++ + 4 + + + - T ownsendia formosa...
WRT ownsendia leptotes,

нана на ыы Т
.--ае--е---ө-- Townsendia spathulata.
Ф Townsendia glabella.

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 7

which seem to be restricted to rather local areas are Т. formosa of
southwestern New Mexico and adjacent Arizona and Т. texensis of
northwestern Texas. 'The difference between the mountain
species Т. sericea and the plains species Т. exscapa is very well
brought out by material collected in Colorado. Тһе latter has
by far the largest distribution of any member of the genus. It
extends from western Manitoba and eastern Alberta south
through the plains region into Texas, New Mexico, Arizona, and
southwestern Colorado. Т. sericea is found in the Rocky
Mountains from Alberta to Colorado and in the Black Hills of
South Dakota. Another species which has discontinuous dis-
tribution is Т. scapigera which extends from the Uintah Mountains
of Utah to northeastern California and is also found in the region
of Santa Fe, New Mexico.

ACKNOWLEDGMENTS

The writer is indebted to Dr. George T. Moore, Director of the
Missouri Botanical Garden, for the use of the library and her-
barium of that institution, and especially to Dr. J. M. Greenman,
Curator of the Herbarium, for advice and assistance. Acknow-
ledgements are due to Dr. A. W. Hill, Director of the Royal
Botanic Gardens, Kew, England, and to Mr. T. A. Sprague, of
the same institution, for the loan of a part of the original material
on which this genus was founded. She is also indebted to Mr.
D. C. Davies, Dr. P. A. Munz, Dr. Francis W. Pennell, Dr.
William R. Maxon, Dr. N. L. Britton, Dr. P. A. Rydberg, Dr.
B. L. Robinson, Dr. Edwin B. Payson, and Dr. M. O. Malte, for
the loan of material necessary to the study of this genus.

ABBREVIATIONS

The specimens cited in this paper are deposited in various
herbaria which are indicated by the following abbreviations:

C = National Herbarium of the Victoria Memorial Museum,
Ottawa, Canada; F = Herbarium of the Field Museum of
Natural History; G = Gray Herbarium of Harvard University;
Kew = Royal Botanic Gardens, Kew, England; M = Missouri
Botanical Garden Herbarium; NY = New York Botanical
Garden Herbarium; Pomona = Herbarium of Pomona College;

[Vor. 14
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Phil = Herbarium of the Academy of Natural Sciences of Phila-
delphia; В, = Rocky Mountain Herbarium; US = United States
National Herbarium.

TAXONOMY

Townsendia Hook. КІ. Bor. Am. 2: 16. 1834; DC. Prodr. 7:
273. 1838; Nutt. Trans. Am. Phil. Soc. №. S. 7: 304. 1841;
Torr. & Gray, Fl. №. Am. 2: 185. 1842; Benth. & Hook. Gen. РІ.
2: 268. 1873; Gray, Proc. Am. Acad. 16: 82. 1880; Syn. FI.
N. Am. 1°: 166. 1884, and ed. 2, 166. 1886 and 1888; Coulter,
Man. Rocky Mt. Region, 156. 1885; Engl. &Prantl, Nat. Pflanzen-
fam. 4%: 161. 1890; Nelson in Coulter & Nelson, Man. Cent.
Rocky Mts. 507. 1909; Wooton & Standley, Contr. U. S.Nat.
Herb. 19: 691. 1915; Rydb. Fl. Rocky Mts. 873. 1917, and
ed. 2, 873. 1922; Tidestrom, Contr. U. S. Nat. Herb. 25: 553.
1925.

Herbaceous, caulescent or acaulescent, glabrate or pubescent
perennials. Leaves alternate, sessile or petioled, linear-lanceolate
to obovate-spathulate, entire. Involucre of many imbricated,
appressed, lanceolate bracts, usually with lacerate-ciliate mem-
branaceous margins. Heads many-flowered. Ray-flowers nu-
merous, in a single series, pistillate, fertile, rays linear, 2-5-
dentate. Disk-flowers tubular, 5-lobed, perfect. Branches of
the style lanceolate, acutish, hairy toward the tip. Pappus uni-
seriate, that of the disk-flower composed of numerous, rather
rigid, barbellate-scabrous bristles as long as the corolla; that of
the ray-flowers similar or shorter, sometimes squamellate, with
a few longer setae intermixed. Achenes of the disk compressed,
obovate to oblong; those of the ray sometimes triangular, pubes-
cent with bidentate or glochidiate-tipped hairs, sometimes gla-
brate.

Type species: 7. sericea Hook. Fl. Bor. Am. 2: 16, pl. 119.
1834, which was based on the collection of Drummond made in
the “Rocky Mountains."

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 9

KEY TO THE SPECIES

А. Bracts of the was: attenuate-acuminate.

tems erect, в
с. Рарриз ini Р OE АН а 1. T. Parryi
CC. Pappus a crown of minute squammelate setae. ........... 2. T. formosa

BB. Stems ascending, branched.
D. Pappus of ray-flower coroniform-concreted; that

of disk-flower similar but with two stout awns........... 3. T. eximia
DD. Pappus in ray-flower a crown of short distinct squa-
mellae; that of disk-flower plurisetose............... 4. T. grandiflora

AA. Bracts of the involucre acute or obtuse.
E. Leaves glabrous or glabrate.
Е. Involucral bracts scarious-margined.
С. Pappus of ray-flower a crown of short squamellae....5. Т. lezensis

GG. Pappus of ray-flower рішгізебове...................... 6. T. glabella
FF. Involucral bracts not scarious-margined ............. 7. T. Rothrockit

EE. Leaves persistently pubescent.
Н. Achenes glabrous ог шаһғайе.......................... 8. Т. montana

НН. Achenes persistently pubescent.
J. Hairs on achenes bidentate.
K. Plants cinereous with short hirsute pubescence.
L. Involucral bracts linear-lanceolate.
Lea

s all obovate-spathulate................ 9. T. Watsoni

MM. Leaves of stem mostly linear-spathulate........ 10. $ өне
LL. Involucral bracts broadly lanceolate.......... IX

KK. Plants densely subsericeous, villous.............. I$. T. эзет чи

JJ. Hairs on achenes glochidiate-tipped.
N. Plants with branched ascending stems.
О. Involucral bracts in two series, equal.......... 18. T. mexicana
OO. Involucral bracts in m series, unequal...... 14. T. strigosa
NN. Plants depressed, caespito
P. Densely canescent, ақы
Q. Leaves 1.5-3 cm. long, narrowly spathulate

to оМалпсеоіміз аг а қ Өн; зеи. 15. T. incana
QQ. Leaves 1-1.5 cm duod obovate-spathulate....16. Т. arizonica
PP. Sparsely hirsute publie qned АСТ! 17. T. leptotes

NNN. Plants strictly nga 1: р
В. Leaves narrowly oblanceolate, somewhat glabrate,
distinctly fattened: ART EARS CER vi de И 18. T. exscapa
RR. Leaves linear-lanceolate, densely pubescent, subterete.19. Т. sericea

1. T. Parryi Eaton, Am. Nat. 8: 212. 1874; Gray, Proc. Am.
Acad. 16: 82. 1880; Syn. Fl. N. Am. 1°: 167. 1884, and ed. 2,
167. 1886 and 1888; Coulter, Man. Rocky Mt. Region, 156.
1885; Howell, Fl. Northwest Am. 306. 1897; Nelson in Coulter
& Nelson, Man. Cent. Rocky Mts. 508. 1909; Rydb. ЕІ.
Rocky Mts. 874. 1917, and ed. 2, 874. 1922

[Vor. 14
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Stem erect, naked and pedunculiform above, sparingly leafy
below, rarely branched, 5-35 cm. high; leaves rosulate, spathulate,
often apiculate, tapering into a petiole, 2-5 cm. long, .3-.5 ст.
broad; heads solitary, pedunculate or rarely sessile, 3—6 cm. in
diameter including the rays; involucre 3—6-seriate; bracts lance-
olate, acute, with narrow scarious margins, lacerately ciliate,
inner bracts acuminate; rays twice the length of the involucre,
blue; pappus the same in ray- and disk-flowers, persistent, pluri-
setose, a little longer than the achene; achenes pubescent with
bidentate hairs.

Distribution: Rocky Mountains from southwestern Canada to
Colorado, west to eastern Oregon.

Specimens examined:

Montana: Teton River at the foot of the Rocky Mts., on hard,
stony, gravelly plains, May 19, 1854, Doty 59 (M); bluffs, Mid-
vale, July 4, 1903, Umbach 254 (F); bluffs, Midvale, June 24,
1903, Umbach 142 (Е); canyon, Helena, July, 1892, Aiton (Е,
No. 90554); Helena, June, 1891, Kelsey (F, Nos. 397673 and
397674); Helena, June, 1892, Starz (M, No. 713519); mountains
about Helena, Anderson (M); near Butte, alt. 1846 m., July,
1893, Mrs. C. H. Moore (M); rocky canyon, dry ground, Bozeman
and vicinity, June 15, 1905, Blankinship 301 (F); Gallatin Co.,
May, 1888, T'weedy 228 (Е); mountain meadows, alt. 1537 m.,
Bozeman, June 1, Livingston, June 8, 1906, Blankinship 301а
(Е, М, В); on a clayey gravelly slope in the foothills ten miles
east of Monida, Madison Co., June 18, 1899, А. & Е. Nelson 5425
(M, R); Bridger Mts., June 11, 1897, Rydberg & Bessey 5132
(F, R); Belt Mts., July 17, 1886, Anderson (F, No. 360840);
Little Belt Pass, alt. 2154 m., Aug. 10, 1896, Flodman 828 (М);
1888, Kelsey (М, No. 783933); July, 1894, Mrs. Moore (М).

YELLOWSTONE NATIONAL Park: Electric Peak, alt. 2923 m.,
July 26, 1902, Smith 16 (F); Mt. Washburn, July 20, 1902,
Smith (Е, No. 121846); Mammoth Hot Springs, June 15, 1902,
Mearns (F, No. 121848); subalpine, alt. 2923 m., Aug., 1884,
Tweedy (Е, No. 211406); Swan Lake, alt. 2308 m., June, 1885,
Tweedy 695 (F); rocky hills near Mammoth Hot Springs, alt.
1846 m., July, 1893, Burglehaus (M); Mt. Washburn, July, 1912,
Evkenberry 59 (Е).

LARSEN—REVISION OF THE GENUS TOWNSENDIA 11

Ууомгма: Wind River Mts., alt. 2764 m., 1873, Parry 144
(F, M, co-type) ; Gros Ventres Fork, alt. 2000 m., June 10, 1860,
Hayden (M); slopes at timber-line, Wyoming Range, 15 miles
west of Merna, Sublette Co., July 18, 1922, E. & L. Payson 2762
(M, R); sage-brush slopes 20 miles west of Big Piney, Sublette
Co., July 10, 1922, E. & І. Payson 2632 (M, В); Gros Ventres
Fork, alt. 2400 m., June 5, 1860, Hayden (M); in the vicinity of
Green River Lakes, Sublette Co., alt. 300 m., Aug. 11, 1925, E. &
L. Payson 4642 (R).

COLORADO: dry ridge near Cottonwood Lake, east of Smoot,
Lineoln Co., alt. 3169 m., Aug. 2, 1923, E. & L. Payson 36938
(M).

ТІрлно: dry hillside south of Henry Lake, Fremont Co., alt.
1846 m., July 15, 1920, E. & L. Payson 2026 (M, R); exposed
rocky slopes, base to summit of mountains northeast of Henry
Lake, Fremont Co., alt. 2830 m., July 11, 1920, E. & L. Payson
1979 (M, R).

САМАРА: Moose Mts.," Rocky Mts., alt. 2061 m., June 30,
1897, Macoun (F, No. 227891); Mount Forget-me-not, July 16,
1897, Macoun (F, No. 227661).

OnEGON: subalpine ridges of the Wallowa Mts., alt. 2154 m.,
July 31, 1899, Cusick 2295 (F, M, R).

2. T. formosa Greene, Leafl. Bot. Obs. & Crit. 1: 213. 1906;
Wooton & Standley, Contr. U. S. Nat. Herb. 19: 692. 1915;
Nelson in Coulter & Nelson, Man. Cent. Rocky Mts., 508. 1909.

T. pinetorum Greene асс. to Nelson in Coulter & Nelson,
Man. Cent. Rocky Mts., 508. 1909, in synonymy.

An herbaceous perennial, spreading by short, stout stolons,
the sterile ones ending in a rosette of leaves, the others in а stout
upright monocephalous stem about 25 ст. in height; stem striate,
sparsely pubescent; leaves thin, glabrous except at the callose-
ciliate margin, basal leaves obovate-spathulate, 1.5—4 cm. long,
.5-1.5 em. broad, very obtuse, narrowed below into a sessile or
subpetiolate base, those of stem oblong-spathulate, sessile, gradu-
ally reduced towards the inflorescence; heads large, 4—6 cm. in
diameter including the rays; Азов 2-3-seriate; bracts with
broad membranaceous margins, minutely Hotê elite, those

[Vor. 14
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of the outer series broadly ovate, those of the inner series linear-
lanceolate, distinctly attenuate-acuminate; achenes glabrous.
Plate 2; pl. 5, fig. 19-24.

Distribution: known only from southwestern New Mexico and
adjacent Arizona.

Specimens examined:

New Mexico: Mogollon Mountains on or near the west fork
of the Gila River, Socorro County, alt. 2615 m., Aug. 8, 1903,
Metcalfe 413 (M, R); around the south end of the Black Range,
Sawyer's Peak, Grant Co., alt. 2770 m., Sept. 30, 1904, Metcalfe
1434 (M, co-type); Sacramento Mts., July 28, 1899, Wooton (В).

ARIZONA: White Mountains, Апр. 6-15, 1903, Griffiths 5340
(M); Bonita Creek, White Mts., July 23, 1912, Goodding 1235
(R); Thompsons Raneh, Black River, White Mts., July 13, 1910,
Goodding 561 (В).

3. Т. eximia Gray, Мет. Ат. Acad. (РІ. Fendl.) N.S. 4: 70.
1849; Walp. Ann. Bot. Syst. 2: 822. 1851-1852; Syn. Fl. N.
Am. 1°: 167. 1884, and ed. 2, 169. 1886 and 1888; Proc. Am.
Acad. 16: 83. 1880; Coulter, Man. Rocky Mt. Region, 156.
1885; Coulter & Nelson, Man. Cent. Rocky Mts. 508. 1909;
Wooton & Standley, Contr. U. S. Nat. Herb. 19: 692. 1915;
Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

T. Vreelandii Rydb. Bull. Torr. Bot. Club 28: 22. 1901;
Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

Herbaceous perennial, caudex sending up a number of simple
or branched stems 15-35 ст. high; leaves spathulate or the upper
ones lanceolate, 2-5 cm. long, nearly glabrate; heads terminal,
solitary; involucre 3-4-seriate, 1-1.5 em. broad, 2-3 em. in di-
ameter; bracts ovate-lanceolate and somewhat cuspidate-acumin-
ate with а narrow membranaceous lacerate-ciliate margin; ray-
flowers about 35-40, fertile, blue or purple, with а much-reduced
persistent pappus of rigid coroniform-concreted squamellae;
pappus of the disk-flowers containing two subulate corneous
stout awns which are slightly shorter than the achene; achenes
pubescent with glochidiate-tipped hairs, at maturity broadly
ovate with a cartilaginous margin. Plate 5, fig. 13-18.

Distribution: mountains of southern Colorado and northern
New Mexico.

1927]
LARSEN— REVISION OF THE GENUS TOWNSENDIA 13

Specimens examined:

COLORADO: side of Veta Mt., alt. 2600 m., July 19, 1900, Vree-
land 639 (NY, түре of T. Vreelandit).

New Mexico: Gallinas Valley above the Hot Springs, Las
Vegas, Sept. 12, 1881, G. Engelmann (M); Las Vegas, Sept.,
1881, G. Engelmann (M); dry hills and hillsides, Sandia Mts.,
Balsam Park, alt. 2500 m., July to Aug., 1914, Ellis 56 (M); La
Glorieta, 1879, Brandegee (F, No. 204786) ; Glorieta, 1881, Vasey
(F, No. 211503); sides of high mountains up Santa Fe Creek,
June 28, 1847, Fendler 353 (M, co-type) ; Santa Fe, 1891, Alcott
(M, Nos. 890716 and 890499); Canyoncito, Santa Fe Co., alt.
2210 m., June 18, 1897, A. A. ғ E. G. Heller 3726 (M); Al-
buquerque, Sandia Mts., Sept. 6, 1884, Jones 4157 (F, R);
Harvey's Upper Ranch in Pecos River National Forest, alt.
2985 m., Aug. 1, 1908, Standley 4621 (M); below Winsors Ranch,
in Pecos River National Forest, alt. 2550 m., July 19, 1908,
Standley 4412 (M); Rito de los Frijoles, Aug., 1910, Robbins 8189,
(R).

4. T. grandiflora Nutt. Trans. Am. Phil. Soc. N.S. 7: 306.
1841; Torr. & Gray, Fl. N. Am. 2: 186. 1842; Gray, Mem. Am.
Acad. (Pl. Fendl.) N.S. 4: 70. 1849; Proc. Am. Acad. 16: 83.
1880; Syn. Fl. N. Am. 17: 167. 1884, and ed. 2, 167. 1886 and
1888; Coulter, Man. Rocky Mt. Region, 156. 1885; Britton,
Man. Fl. Northern States and Canada, 944. 1901; Nelson in
Coulter & Nelson, Man. Cent. Rocky Mts., 509. 1909;
Wooton & Standley, Contr. U. 5. Nat. Herb. 19: 692. 1915;
Rydb. Fl. Rocky Mts. 873. 1917, and ed. 2, 873. 1922.

Caulescent, divaricately branched from the base, 5-20 ст.
high; leaves linear, sublanceolate, acute, nearly glabrous, 2-5
ст. long, 0.3-1 ст. broad, two or more of the uppermost usually
subtending the head; involucre usually 3-seriate; bracts ovate-
lanceolate and rigidly cuspidate-acuminate, with narrow mem-
branaceous lacerate-ciliate margins; heads 1.5-5 cm. in diameter
including the rays; pappus of ray-flower reduced to a crown of
short squamellae, that of the disk-flower plurisetose, longer than
the achene; achenes sparsely pubescent with glochidiate-tipped
hairs.

[Vor. 14
14 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: eastern South Dakota and Nebraska, west to
Wyoming and south to New Mexico; common throughout eastern
Colorado.

Specimens examined:

SourH DaxoTa: Running Water, Aug. 14, probably 1856,
H. Engelmann (M); Black Hills on upper Pole Creek, Aug. 1,
1856, H. Engelmann (M); Badlands, Cheyenne Valley, Wash-
ington Co., July 23, 1911, Visher 2138 (Е).

NEBRASKA: Warbonnet Canyon, alt. 1532 m., June, 1890, Т. A.
Williams (M); Pine Ridge, July 21, 1889, Webber (М); Eaglenest
Butte, 1853-4, Hayden (M).

WowiNa: stony slopes, Laramie Co., June 29, 1901, Nelson
8312 (M, R); open sandy slopes south of Sibylee, Albany Co.,
July 3, 1900, A. Nelson 7373 (M, R); Corlett, June 24, 1907,
Johnston 258 (M). |

CoLonApo: Douglas Co., 1892, Walker (Е, No. 376084);
Castle Rock, 1889, Walker (F, No. 376085); Florence, July 31,
1872, Brandegee 487 (M); Colorado Springs, July 19, 1872, Red-
field 478 (M); Soldier, June 13, 1899, Marshall 3138 (Е); Garden
of the Gods, alt. 2000 m., Aug. 14, 1922, Brumback & Davis 178
(F); Garden of the Gods, near Colorado City, July 18, 1872,
Porter (F, No. 318424); Eldorado Springs, alt. 1631 m., June 24,
1917, Clokey 2810 (F, R); in dry soil, Boulder Canyon, alt. 1692
m., Young (Е, No. 290193); Colorado Territory, 39—40° lat., alpine
and subalpine, 1864, Parry (M); Rocky Mts., 40-41? lat., Vasey
304 (M); infrequent, mesa slopes, Boulder, alt. 1690 m., June 23,
1921, Hanson C 159 (M); mountains, Larimer Co., alt. 2308 m.,
June 14, 1896, Crandall (M); Una, July 10, 1894, A. Nelson 385
(M); Platteville, Apr. 17, 1908, Johnston 492 (M); Horsetooth
Gulch, 10 miles southwest of Fort Collins, alt. 1385 m., June 30,
1893, Baker (M); Golden, Jefferson Co., July 4, 1915, Johnston
414 b (M); foothills near Golden, Castle Rock, July 1, 1885,
Patterson 49 (F, M); foothills near Golden, June 20, 1878, Jones
284 (F); Gold Hill, Aug. 12, 1875, Patterson (Е, No. 208980);
near Golden City, 1870, Greene (F, No. 15363); Fremont Co.,
near Canyon City, 1873, Greene (Е, No. 15364); 39-402 lat., alpine
and subalpine, 1864, Parry (M); Rocky Mts., 40? lat., 1862, Hall
(F); Rocky Mts., Hall & Harbour (F, No. 367351); Rocky Mt.

1927]
LARSEN—REVISION ОЕ THE GENUS TOWNSENDIA 15

flora, 39-41? lat., 1862, Hall & Harbour 289 (F, M); eastern
Colorado, Carleton (F, No. 353119) ; southern Colorado, Brandegee
(Е, No. 204740); Manitou, Aug. 11, 1884, Letterman 88 (M).

New Mexico: low prairie between Orate Creek and Rio Col-
orado, Aug. 21, 1847, Fendler 533 (M); Moro River Prairie,
Aug. 15, 1847, Fendler 15? (M).

5. T. texensis Larsen, n. зр.!

Herbaceous perennial; the caudex giving rise to ascending
branched stems 6-30 ст. high, leafy throughout; leaves oblance-
olate, 1-5 cm. long, 0.2-0.8 em. broad, apiculate, narrowed at the
base into а petiole, sparsely pubescent with closely appressed
hairs, occasionally glabrate; heads usually solitary, terminal,
sessile or short-pedunculate, 1.5-3 ст. in diameter including the
гау; involucre 4—5-seriate, 1-1.5 em. in diameter; bracts oblance-
olate, acute, pink-tipped, membranaceous-margined and lacerate-
ciliate; ray-flowers dark blue or purple with a reduced plurisetose
pappus scarcely longer than the breadth of the achene; pappus of
disk-flowers plurisetose, somewhat shorter than the corolla;
achene pubescent with glochidiate-tipped hairs. Plate 3.

Distribution: northwestern Texas.

Specimens examined:

Texas: Randall Co., “rocky bluffs of the Red River," Aug.
13, 1900, Eggert (М, Nos. 121021, түре, 121022, 121023) ; Canyon,
Aug. 13, 1900, Eggert (M, No. 720398); Randall Co., “rocky
banks of the Red River," Aug. 12, 1900, Eggert (M, Nos. 121028
and 121027); rocky bluffs of Paloduro, May 30, 1902, Reverchon
3320 (M); abundant on barren slopes, branch of Paloduro Can-
yon, Sept. 12, 1917, Young (M, Nos. 831212 and 831677); dry

! Herbaceis perennis; caule ramoso, ramis ascendentibus, 6-30 cm. altis, foliaceis;
foliis oblanceolatis, 1-5 cm. longis, 0.2-0.8 cm. latis, apiculatis, integris, basi in
petiolam sensim angustatis, visitique dense strigoso-pilosis, rarius glabratis; capitulis
qug solitariis, terminalibus, sessilibus vel brevi-pedunculatis, radio incluso
1.5-3 cm. in diametro; involucris campanulatis, 4—5-seriatis, 1-1.5 сіп. in diametro;
bracteis s involucri oblanceolatis, acutis ad apices roseis, marginibus membranaceis
lacerato ligulatis, ligulis atro-caeruleis vel purpurascenti-
bus, pappi setis multo reductis vix ` diametro achenii longioribus; floribus disci
numerosis, pappi setis multis paululo corollis brevioribus; acheniis glochideo-
pubescentibus.—Collected on “госку bluffs of the Red River," Texas, Aug. 13,
1900, H. Eggert (Mo. Bot. Gard. Herb., No. 121021, TYPE).

[Vor. 14
16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

open ground, caleareous soil, Channing, Hartley Co., June 19,
1918, Е. J. Palmer 14170 (M); calcareous open wound on plains,
Canyon, Randall Co., Oct. 13, 1918, Е. J. Palmer 14586 (М);
Canyon, Randall Co., June 12, 1917, Е. J. Palmer 12510 (М).

6. T. glabella Gray, Proc. Am. Acad. 16: 86. 1880; Syn. FI.
N. Am. 17: 169. 1884, and ed. 2, 169. 1886 and 1888; Coulter,
Man. Rocky Mt. Region, 158. 1885; Nelson in Coulter &
Nelson, Man. Cent. Rocky Mts. 510. 1909; Rydb. Fl. Rocky
Mts. 875. 1917, and ed. 2, 875. 1922.

Т. Bakeri Greene, Pittonia 4: 157. 1900.

Subacaulescent; caudex thick, woody, bearing tufted leaves;
leaves thick, pilose when young, soon becoming glabrous, linear-
spathulate, 2-4 ст. long, 0.2-0.5 ст. broad, gradually narrowed
into a slender petiole; heads solitary, on naked peduncles 2-5 em.
long; heads 1.5-2.5 ст. in diameter including the rays; involucre
2-3-seriate; bracts of the involucre oblong, with narrow mem-
branous margins finely lacerate-ciliate; ray-flowers blue or purple,
setae irregular, varying in length from 1 to 5 mm.; pappus of
disk-flowers regular, as long as the corolla; achenes sparsely pu-
bescent with glochidiate-tipped hairs.

Distribution: known only from southwestern Colorado.

Specimens examined:

COLORADO: Pagosa Springs, Aug. 13, 1917, Payson 1160 (М,
R); dry hills, Pagosa Springs, Archuleta Co., alt. 2160 m.,
June 29, 1921, Bethel, Willey & Clokey 4340 (F, M, R); hillside
near Dix, alt. 2615 m., Baker, Earle & Tracy 548 (Е, М, В); Los
Pinos (Bayfield), May 16, 1899, Baker 727 (Е, М, В, түре of T.
Bakeri); mature yellow pine forest, Piedra, June 31, 1924, Hazel
М. Schmoll 1348 (К); open places between pines, Piedra, June 21,
1914, Hazel M. Schmoll 1212 (R).

7. T. Rothrockii Gray, acc. to Rothrock in Wheeler Rept. 6:
148, t. 7. 1878; Gray, Proc. Am. Acad. 16: 85. 1880; Syn. Е.
М. Am. 17: 168. 1884, and ed. 2, 168. 1886 and 1888; Coulter,
Man. Rocky Mt. Region, 157. 1885; Nelson in Coulter & Nelson,
Man. Cent. Rocky Mts. 510. 1909; Rydb. Fl. Rocky Mts. 875.
1917, and ed. 2, 875. 1922.

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 17

Acaulescent; leaves broadly spathulate, approximately 2 cm.
long, glabrous, rosulate; heads sessile, approximately 2 cm. in
diameter including the rays; involucre 3-4-seriate; bracts oblong
or narrowly ovate, purplish, thickish-margined, distinctly ciliate;
ray-flowers with a much-reduced pappus; pappus of disk-flowers
equalling or exceeding the length of the corolla; achenes sparsely
pubescent with glochidiate-tipped hairs.

Distribution: alpine districts of Colorado.

Specimens examined:

COLORADO: loamy places of the foothills, Sheep and Engineer
(?) Mts., Uncompahgre River, alt. 3000-3500 m., Aug. 2, 1893,
Purpus 532 (F); South Park, alt. 4150 m., July, 1873, Rothrock
876 (Е, TYPE); South Park, Aug., 1873, Rothrock (Е, No. 304922
in part).

8. T. montana Jones, Zoe 4: 262. 1893; Rydb. Fl. Rocky
Mts. 874. 1917, and ed. 2, 874. 1922; Tidestrom, Contr. U. В.
Nat. Herb. 25: 554. 1925.

T. alpigena Piper, Bull. Torr. Bot. Club 27: 394. 1900.

T. dejecta Nelson, Bot. Gaz. 27: 267. 1904; Nelson in Coulter
& Nelson, Man. Cent. Rocky Mts. 510. 1909.

Caespitose from a multicipital caudex; leaves obovate-spathu-
late, 1-5 ст. long, 0.3-0.5 ст. broad, apiculate, narrowed at the
base into a petiole, pubescent in the early stages with appressed
strigulose hairs, more or less glabrate; heads 1.5-2.5 em. in di-
ameter including the rays, sessile or solitary on naked scapes,
1-5 cm. in length; involucre about 3-seriate; bracts of outer
series lanceolate, those of the inner series obovate, obtuse, pink-
tipped, membranaceous, lacerate-margined; pappus of disk-
flowers plurisetose, equalling the length of the corolla, pappus of
ray-flowers similar but somewhat shorter; achenes glabrous or
glabrate; achenes of ray-flowers occasionally hairy toward the
base with a few scattered bidentate hairs.

Distribution: western Wyoming to Oregon, southward into
Utah.

Specimens examined:

YELLOWSTONE NATIONAL Park: Mammoth Hot Springs, July
5, 1902, Mearns (F, No. 121847).

[Vor. 14
18 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

WYOMING: Piney Mt., 25 miles west of Big Piney, Sublette
Co., Summit, July 12, 1922, E. 4 L. Payson 2694 (М, В); cal-
careous slide rock, Teton Pass Mts., east of Victor, alt. 2831 m.,
July 22, 1920, E. & L. Payson 2078 (M, R); mountains near
Cottonwood Lake, east of Smoot, Lincoln Co., alt. 3200 m.,
Aug. 2, 1923, Payson & Armstrong 3706 (M, В); Sheep Mt.
(Ferry Peak), Snake River Range, near Alpine, Lincoln Co., July
11, 1923, Payson & Armstrong 3474 (M, В); in vicinity of Green
River Lakes, Sublette Co., alt. 3169 m., Aug. 5, 1925, E. & L.
Payson 4542 (M).

IpauHo: subalpine slopes of loose calcareous soil, base to summit
of mountains northeast of lake, Henry Lake, Fremont Co., alt.
2678 m., July 11,1920, E. & L. Payson 1986 (M, R) ; Mt. Chauvet,
July 29, 1897, Rydberg & Bessey 5131 (F, R).

Отан: canyon above Tropic, alt. 2154 m., Мау 29, 1894, Jones
(Pomona, No. 40754); mountains above Silver Lake, July 30,
1880, Jones (Pomona, No. 40756); Alta, above the Flagstaff
Mine, Aug. 7, 1879, Jones (Pomona, No. 40755, TYPE of Т. mon-
tana); loose stony soil, Uintah Mts., Dyer Mine, July 3, 1902,
Goodding 1238 (В, M, co-type of Т. dejecta).

OREGON: subalpine ridges of Wallowa Mts., alt. 2015 m., July
31, 1899, Cusick 2294 (F, M, co-types of T. alpigena).

9, T. Watsoni Gray, Proc. Am. Acad. 16: 84. 1880; Syn. FI.
М. Am. 1°: 168. 1884, and ed. 2, 168. 1886 and 1888; Nelson
in Coulter & Nelson, Man. Cent. Rocky Mts. 509. 1909; Tide-
strom, Contr. U. S. Nat. Herb. 25: 554. 1925.

T. strigosa Eaton, non. Nutt. Bot. King's Exp. 145. 1871;
Gray, Syn. Fl. №. Am. 12: 168. 1884, and ed. 2, 168. 1886 and
1888.

Caulescent, hirsute with a close appressed pubescence; the
caudex sending up a number of branched, sparingly leafy stems;
leaves mostly obovate-spathulate, 2-4 cm. long, 0.2-0.8 cm. broad,
narrowed into a petiole; heads on short bracteate or naked pe-
duncles, 0.3-1 cm. long, 1-1.5 em. in diameter including the rays;
involucre 2-seriate; bracts oblong-lanceolate, margins membran-
aceous, lacerate-ciliate; ray-flowers with а reduced pappus of
unequal capillary bristles shorter than the diameter of the achene;

1927] |
LARSEN—REVISION OF THE GENUS TOWNSENDIA 19

pappus of the disk-flower equalling or surpassing the corolla;
achenes pubescent with bidentate hairs.

Distribution: southeastern Oregon through Nevada to south-
western Utah.

Specimens examined:

Uran: Dugway, May 28, 1891, Jones (M); Glenwood, alt.
1692 m., May 24, 1875, Ward 92 (F, M).

NEVADA: 1891, A. J. Jones (М).

OnEGON: sandy soil near Vale, May, 1896, Leiberg 2067 (M);
common on hills, in the region of Malheur River, June 19, 1898,
Cusick 1985 (F, M).

10. T. florifer (Hook.) Gray, Proc. Am. Acad. 16:84. 1880; Syn
ЕІ. №. Am. 1::167. 1884, and ed. 2,167. 1886 апа 1888; Coulter,
Man. Rocky Mt. Region, 157. 1886; Howell, Fl. Northwest
Am. 306. 1897; Piper, Contr. U. S. Nat. Herb. 9: 563. 1906;.
Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922;
Tidestrom, Contr. U. S. Nat. Herb. 25: 554. 1925.

T. strigosa Gray in Wilkes’ Exp. 17: 344. 1874, not Nutt.,
fide Gray, Proc. Am. Acad. 16: 84. 1880.

T. florifer Gray var. communis Jones, Proc. Calif. Acad. II. 5:
697. 1895.

Erigeron ? florifer Hook. Fl. Вог. Am. 2:20. 1834.

Aplopappus florifer Hook. & Arn. Bot. Beechey Voy. 351.
1841, excl. var. 6.

Stenotus florifer Torr. & Gray, Fl. N. Am. 2: 238. 1842, excl.
var. В.

Caulescent, cinereous-hirsute; the caudex sending up a number
of simple or branched stems 5-18 cm. high, leafy throughout;
leaves linear or the lowest lanceolate-spathulate, acute, mostly
apiculate-acuminate; heads 2-3 cm. in diameter including the
rays, solitary, terminating the branches; involucre 2-3-seriate;
bracts linear-lanceolate and acute, of about equal length, the
membranaceous margins lacerate-ciliate; ray-flowers with pappus
similar to that of the disk-flowers but varying in length from about
the width of the achene to nearly as long as that of the disk-
flowers; pappus of disk-flower composed of coarse, white setae
which exceed the corolla in length; achenes densely pubescent
with bidentate hairs. Plate 5, figs. 25-28.

[Уог. 14
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: dry hills and plains, central Washington and
Oregon, southeastward through southern Idaho, Utah, to western
Wyoming.

Specimens examined:

WyominG: moist rich bottoms, Gros Ventres Fork, alt. ‚2000
m., June 10, 1860, Hayden (М).

fino: ditch banks along fields, Challis, Custer Co., alt. 1662
m., July 15, 1916, Macbride & Payson 3225 (М, В); clayey
hills, Kings Hill, Elmore Co., alt. 800 m., July 16, 1911, Nelson &
Macbride 1129 (M, R); dry stony hillsides and dry flats,
Arco, Blaine Co., alt. 1640 m., July 3, 1916, Macbride
& Payson 3095 (M, R); loose soil, Reynolds Creek, Owyhee
Co., alt. 1538 m., July 3, 1911, Macbride 1017 (M, R); gravelly
slopes, New Plymouth, Canyon Co., alt. 680 m., May 21, 1910,
Macbride 90 (M, R); near Nampa, July 1, 1892, Mulford (M).

Отан: Joseph City, Sevier Co., alt. 1692 m., Мау 13, 1899,
Jones 6379 (М); Marysvale, alt. 1846 m., Мау 31, 1894, Jones
5323 (М, Е); rim of Great Salt Lake Desert, May 6, 1889, Н.
Engelmann (M); sage-brush slopes, Milford, June 5, 1902, Good-
ding 1046 (R).

WasHINGTON: Ritzville, Adams Co., alt. 480 m., June 6, 1893,
Sandberg & Leiberg 169 (Е, М); Craigs Ferry, Kittitas Co., July
15, 1903, Cotton 1361 (M); Columbia River opposite Umatilla,
Apr. 20, 1882, Howell (Е, No. 206944); Columbia River opposite
Umatilla, Apr. 29, 1882, Howell (M); Wilson Creek, June, 1893,
Sandberg & Leiberg (M); Yakima region, Cascade Mts., June,
1882, Brandegee (М).

OREGON: near Lexington, Morrow Co., alt. 420 m., Мау 7,
1894, Leiberg 34 (F, M); near Umatilla, May 1, 1882, Howell (F,
No. 396898); on open plains, Cline Falls, Crook Co., May 22,
1905, Nelson 815 (M, R); dry banks of Deschutes River five miles
below Bend, July 30, 1920, Peck 9708 (М); stony hills west of
Silver Creek (and common westward), June 28, 1901, Cusick 2616
(F, M, R).

11. Т. scapigera Eaton, Bot. King’s Exp. 145. t. 17. 1871;
Gray, Proc. Am. Acad. 16: 84. 1880; Syn. Fl. М. Am. 17: 168.
1884, and ed. 2, 168. 1886 and 1888; Rydb. Fl. Rocky Mts. 874.

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 21

1917, and ed. 2, 874. 1922; Tidestrom, Contr. U. S. Nat. Herb.
25: 554. 1925; Jepson, Man. Fl. Plants California, 1044. 1925.

Aplopappus florifer var. % Hook. & Arn. Bot. Beechey Voy.
351. 1841 (fide Gray).

Stenotus florifer var. В Torr. & Gray, Fl. М. Am. 2: 238. 1842
(fide Gray).

T. scapigera var. caulescens Eaton, Bot. King's Exp. 145. 1841;
Gray, Proc. Am. Acad. 16: 84. 1880; Syn. Fl. N. Am. 17: 168.
1886, and ed. 2, 168. 1886 and 1888.

Т. scapigera var. ambigua Gray, Proc. Am. Acad. 16: 84. 1880;
Syn. Fl. №. Am. 17: 168. 1884, and ed. 2, 168. 1886 and 1888;
Tidestrom, Contr. U. S. Nat. Herb. 25: 554. 1925.

T. ambigua (A. Gray) Rydb. Fl. Rocky Mts. 874. 1917, and

ed. 2, 874. 1922.

Herbaceous perennial, canescent with fine appressed pubes-
cence; caudex bearing tufted leaves; leaves narrowly spathulate
to obovate, 1-3 em. long, 0.3-0.8 cm. wide, lamina sometimes
emarginate, narrowed into a petiole; flowering scapes 1-5 cm.
long, naked or 1-2-bracted, sometimes leafy; heads 2-2.5 cm. in
diameter including the rays; involucre 2—3-seriate; bracts oblong-
lanceolate, acute, margins lacerate-ciliate; pappus of ray-flower
similar to that of the disk but somewhat shorter; pappus of disk-
flower plurisetose, exceeding the length of the corolla; achenes
pubescent with bidentate hairs.

Distribution: Wyoming to northeastern California; also in New
Mexico.

Specimens examined:

Wyomine: Holm Lodge, about 40 miles west of Cody, Park
County, Aug. 26 and 27, 1922, von Schrenk (M).

New Mexico: Santa Fe, 1847, Fendler 351 (Phil.) ; without defi-
nite locality, Kern (in part) (Phil.).

Гтан: Rabbit Valley, alt. 2092 m., Aug. 6, 1875, Ward 523
(О. S., TYPE of T. scapigera var. ambigua); Deep Creek, June 6,
1891, Jones (Pomona, No. 40882).

NEVADA: Monitor Valley, alt. 1538 m., July, 1868, Watson 519
(US, түре of T. scapigera var. caulescens) ; eastern Nevada, 1883,
Meehan (Phil.).

CALIFORNIA: Buffalo Ravine near Surprise Valley, Apr. 1879,
Lemmon 29 (M).

[Vor. 14
22 ANNALS OF THE MISSOURI BOTANICAL GARDEN

12. T. spathulata Nutt. Trans. Am. Phil. Soc. N.S. 7: 305.
1841; Torr. & Gray, Fl. №. Am. 2: 186. 1842; Eaton, Am. Nat.
8: 213. 1874; Gray, Proc. Am. Acad. 16: 86. 1880; Syn. Fl. №.
Am. 1°: 169. 1884, and ed. 2, 169. 1886 and 1888; Coulter,
Man. Rocky Mt. Region, 158. 1885; Nelson in Coulter & Nelson,
Man. Cent. Rocky Mts. 510. 1909; Rydb. Fl. Rocky Mts. 875.
1917, and ed. 2, 875. 1922.

T. condensata Eaton, Am. Nat. 8: 213. 1874; Gray, Proc. Am.
Acad. 16: 83. 1880; Syn. Fl. №. Am. 17: 167. 1884, and ed. 2,
167. 1886 and 1888; Coulter, Man. Rocky Mt. Region, 157.
1885; Nelson in Coulter & Nelson, Man. Cent. Rocky Mts. 519.
1909; Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

T. Parryi var. alpina Gray, Proc. Am. Acad. 16: 83. 1880;
Syn. Fl. №. Am. 17: 167. 1884, and ed. 2, 167. 1886 and 1888;
Coulter, Manual Rocky Mt. Region, 156. 1885.

T. alpina (Gray) Rydb. Mem. N. Y. Bot. Gard. 1: 390. 1900;
Nelson in Coulter & Nelson, Man. Cent. Rocky Mts. 509. 1909;
Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

Caespitose perennial, 3-5 cm. high; leaves crowded, rosulate,
obovate to spathulate, 1-1.5 cm. long, 0.2-0.4 ста. broad, densely
subsericeous-villose to villose-lanate; heads sessile or pedunculate,
1-5 cm. in diameter including the rays; involucre usually serrate;
bracts pinkish, oblong-lanceolate, those of the inner series with a
weak attenuate apex, the narrow margins scarious, lacerate-
ciliate; pappus of ray- and disk-flowers similar, composed of
slender setae, as long as the corolla of the disk-flower; achenes
pubescent with bidentate hairs.

Distribution: alpine and subalpine regions of southwestern
Alberta and western Wyoming.

Specimens examined:

YELLOWSTONE NATIONAL Park: Electric Peak, alt. 2770 m.,
July 26, 1902, Е. С. Smith (Е).

Wyomina: high alpine ridge between the valleys of the Stinking
Water and the Yellowstone, 1873, Parry 142 (Е, М); northwestern
Wyoming, 1873, Parry 146 (Е, С, М, түре of T. Parryi var.
alpina); high alpine peak, Owl Creek Range, July, 1874, J. D.
Putnam (С, түре of Т. condensata); ‘‘Black Hills of Platte,"
Nuttall (РЫП); shale flats, Bush Ranch, June 10, 1910, Nelson
7054 (В).

1927]
LARSEN—REVISION ОЕ THE GENUS TOWNSENDIA 23

CANADA: High River, Rocky Mts., alt. 2308 m., July, 1884,
Dawson (G).

Note: Dr. C. C. Parry says in a note accompanying the type of
T. condensata Gray: ‘‘Single specimen from a high alpine peak,
Owl Creek Range, by J. D. Putnam. I take this to be a con-
densed alpine form of 145." The head of Putnam's specimen is
larger than in No. 145 and looks very much as if it were a fasci-
ation of several heads. Otherwise, these specimens are identical.
In the specimens cited the heads vary from 1 to 5 ст. in diameter.
The presence of intermediates indicates that the size of the head
cannot be used as a means of specific demarcation.

13. T. mexicana Gray, Mem. Am. Acad. N.S. (Pl. Fendl.) 4:
70. 1849; Walp. Ann. Syst. Bot. 2: 822. 1851-1852; Gray,
Proc. Am. Acad. 16: 86. 1880; Syn. Fl. М. Am. 17%: 169. 1884,
and ed. 2, 169. 1886 and 1888; Hemsley, Biol. Cent.-Am. Bot.
2: 118. 1881.

Caulescent; conspicuously cinereous with strigose pubescence;
stems decumbent, simple or branched, 5-17 cm. long, leafy
throughout; leaves linear, sometimes linear-spathulate, 1-2.5 cm.
long, 0.1-0.2 ст. broad; heads usually solitary on terminal pe-
duncles, 1-2 ст. in diameter including the rays; involucre dis-
tinctly 2-seriate; bracts of equal length and all very obtuse,
membranous-margined ; ray-flowers fertile with a much-reduced
pappus; pappus of the disk-flowers equalling the corolla in length;
achenes sparsely pubescent with glochidiate-tipped hairs.

Distribution: east central Mexico, southern Coahuila, Zacatecas
to Hidalgo.

Specimens examined:

Мех!со:

CoaHUILA: Saltillo, March 22, 1877, Gregg 327 (М, TYPE);
Saltillo, alt. 1650 m., June 5, 1909, Arséne 3387 (M); Saltillo,
Apr. 1-15, 1880, Ed. Palmer 499 (F).

ZACATECAS; near Conception Del Oro, Aug. 11-14, 1904, Ed.
Palmer 252 (M); vicinity of Cedros, Aug., 1908, Kirkwood 110
(F); low places, plains, Cedros, Aug., Lloyd 110 (M).

HipALGo: rocky flats and mountains, Ixmiquilpan, July, 1905,
Purpus 1845 (M, Е); calcareous bluffs near Tula, alt. 2080 m.,

[Vor. 14
24 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Aug. 6, 1896, Pringle 6573 (M); dry calcareous rocks near Tula,
alt. 2080 m., Sept. 16, 1902, Pringle 9967 (M, F); calcareous
plains near Pachuca, alt. 2350 m., Aug. 2, 1898, Pringle 7580
(F, R).

14. T.strigosa Nutt. Trans. Am. Phil. Soc. N.S. 7: 306. 1841;
Torr. & Gray, Fl. №. Am. 2: 186. 1842; Gray, Mem. Am. Acad.
N.S. (Pl. Fendl.) 4: 70. 1849, in part; Syn. Fl. N. Am. І?: 169.
1884, and ed. 2, 169. 1886 and 1888; Proc. Am. Acad. 16: 86.
1880; Coulter, Man. Rocky Mt. Region, 158. 1885; Nelson in
Coulter & Nelson, Man. Cent. Rocky Mts. 509. 1909; Rydb.
Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922; Tidestrom,
Contr. U. 8. Nat. Herb. 25: 554. 1925.

T. Fendleri Gray, Mem. Am. Acad. N.S. (Pl. Fendl.) 4: 70.
1849; Walp. Ann. Syst. Bot. 2:822. 1851-1852; Gray, Syn. Fl. М.
Am. 17: 169. 1884, and ed. 2, 169. 1886 and 1888; Proc. Am.
Acad. 16: 86. 1880; Coulter, Man. Rocky Mt. Region, 158.
1885; Nelson in Coulter & Nelson, Man. Cent. Rocky Mts. 509.
1909; Wooton & Standley, Contr. U. 5. Nat. Herb. 19: 692.
1915; Rydb. Fl. Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

Caulescent; cinereous with close strigulose pubescence, the
caudex sending up a number of simple or branched stems, 3-25
cm. high, nearly naked below, leafy towards the inflorescence;
leaves linear to linear-spathulate, 1-3 ст. long, 0.3-0.6 em. broad,
gradually narrowed into a slender petiole; heads subsessile and
solitary or terminating the lateral branches in 1—3-headed clus-
ters; heads about 1.5 cm. in diameter including the rays; involucre
2-—4-seriate; bracts oblong-lanceolate, membranaceous-margined
and lacerate-ciliate; ray-flowers about 15, rays pink or rose-
purple, with a much-reduced pappus; pappus of disk-flowers
plurisetose, as long as the corolla; achenes pubescent with glo-
chidiate-tipped hairs. Plate 4.

Distribution: dry sandy soil, southwestern Wyoming to New
Mexico and Arizona and adjacent Mexico.

Specimens examined:

WYOMING: dry flats 21 miles west of Green River, June 19,
1923, Payson & Armstrong 3205 (M, В); Church Buttes, Fort
Bridger, July, 1873, Porter (M, Phil.); without definite locality,
Parry (Phil) ; Green River, May 30, 1897, Nelson 3031 (M, R).

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 25

Сотовдро: “К. Mts. Platte," Nuttall (Phil., түре); Salida,
June 19, 1898, Baker, Earle & Tracy 1016 (M); Texas Cr., Fre-
mont Co., 1874, Brandegee 951 (M); Arkansas River near Puncha
Pass, Sept. 24, 1878, Jones 767 (М); Huerfano, Aug., 1867, Parry
93 (M, R); dry slopes, alt. 1662 m., Paradox, Montrose Co.,
June 17, 1912, Walker 93 (M); adobe plains of San Juan Valley,
July, 1875, Brandegee (М); southern Colorado, Brandegee (Phil.) ;
sands of Huerfano Ст., Sept., 1875, Brandegee (М); McElmo Cr.,
June 3, 1892, Eastwood (F, No. 82217); alkaline hillsides, Natur-
ita, 1662 m., May 19, 1924, Payson 321 (F, M, R); Arkansas
Valley, Sept., 1873, Wolf 517 (Е).

New Mexico: near Espanola, Santa Fe Co., alt. 1723 m.,
May 17, 1897, Heller & Heller 3547 (M); Mangas Springs, 18
miles northwest of Silver City, Grant Co., alt. 1323 m., Apr. 12,
1903, Metcalfe 15 (M, R); Mesilla Valley, Dona Ana Co., alt.
1184 m., Apr. 19, 1907, Wooton & Standley 3237 (M); sand hills
near Mesilla, May 4, 1906, Standley (М); gravelly hillsides, Santa
Fe, May-July, 1847, Fendler 350 (M, co-type of T. Fendleri);
El Paso, Apr., 1852, Parry (M); Santa Fe, 1891, Alcott (M, No.
890501); sandy banks of the Rio Grande and stony hills, El Paso,
March-June, 1851-2, Wright 1172 (Phil, M); without definite
locality, Kern (Phil); gravelly hills, Santa Fe, May, 1847,
Fendler 351 (Phil., M); Aztec, Мау 4, 1899, Baker 728 (Е, M, В);
loamy flats, Hillsboro (N. Percha), Sierra Co., alt. 1692 m., Oct.
28, 1904, Metcalfe 1510 (F, M); Mesilla Valley, Dona Ana Co.,
alt. 1154 m., Apr. 2, 1907, Wooton & Standley (F, M).

ARIZONA: dry spots in river bottoms, Rio Verde, Fort Whipple,
Sept. 6, 1865, Coues & Palmer 523 (M); sandy soil, Beaver Cr.,
Sept., 1903, Purpus 8300 (M); Smart 92 (Е); Voth 7 (Е).

15. T. incana Nutt. Trans. Am. Phil. Soc. N.S. 7: 305. 1841;
Torr. & Gray, Fl. М. Am. 2: 155. 1842; Walp. Rep. 2: 575.
1843; Gray, Syn. Fl. №. Am. 12: 169. 1884, and ed. 2, 169.
1886 and 1888; Proc. Am. Acad. 16: 86. 1880; Coulter, Man.
Rocky Mt. Region, 157. 1885; Nelson in Coulter & Nelson,
Man. Cent. Rocky Mts. 509. 1909; Wooton & Standley, Contr.
U. S. Nat. Herb. 19: 692. 1915; Rydb. Fl. Rocky Mts. 875.
1917, and ed. 2, 875. 1922; Tidestrom, Contr. U. S. Nat. Herb.
25: 554. 1925.

[Уоь. 1
26 ANNALS OF THE MISSOURI BOTANICAL GARDEN

T. Fremontii Torr. & Gray, Boston Jour. Nat. Hist. 5: 106.
1845.

T. incana var. ambigua Jones, Zoe 4: 264. 1893.

T. incana var. prolixa Jones, Contr. Western Bot. 13: 15.
1910.

Herbaceous perennial, strigulose-cinereous, caespitose; stem
usually 3-6 сш. high; leaves spathulate, sometimes apiculate,
petiolate, 2-4 ст. long and 0.2-0.5 сіп. broad, the uppermost
clustered at the base of the heads and seldom surpassing them in
length; heads 1-2.5 ст. in diameter including the rays, usually
sessile; involuere 2-3-seriate; bracts broadly lanceolate, their
scarious margins lacerate-ciliate; pappus of the disk-flower
plurisetose, equalling the length of the corolla; pappus of the
ray-flower similar to that of the disk-flower but only one-third
to one-half as long; achenes pubescent with glochidiate-tipped
hairs.

Distribution: Wyoming, south through western Colorado into
northwestern New жеме west into eastern Arizona and Utah.

Specimens examine

Үүоміма: “Black Hills of the Platte," Nuttall (Phil., TYPE);
on a stony flat, Granger, Uinta Co., June 14, 1899, A. & E. Nelson
5408 (M, R); Granger, June 10, 1898, A. Nelson 4622 (M, R);
deep hot sands, Aleova, Natrona Co., July 1, 1901, Goodding 166
(M, R).

COLORADO: dry rocky south slope, Norwood Hill, San Miguel
Co., alt. 2154 m., Aug. 11, 1912, Walker 448 (M, R); Grand
Junction, May, 1891, Eastwood (Е, Pomona); Grand Junction,
June, 1892, Eastwood (F, Pomona); Grand Junction, Apr. 15,
1891, Jones (Pomona, No. 39662) ; Grand Junction, alt. 1412 m.,
June 11, 1901, Baker 105 (M).

New Mexico: Aztec, May 6, 1899, Baker 729 (M, В).

Uman: stony slopes, Thompson Springs, alt. 1630 m., May-
Oct., 1899, Purpus 6765 (M); Thompson Springs, Мау 7, 1891,
Jones (Pomona, No. 39664, түре of Т. incana var. ambigua);
Richfield, alt. 1692 m., June 5, 1875, Ward 176 (F, M); Lower
Crossing, alt. 1384 m., July 2, 1898, Jones (M, No. 121121);
Westwater, May 6, 1891, Jones (Pomona, No. 39663); West-
water, alt. 1380 m., June 28, 1898, Jones (Pomona, No. 39629);

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 27

Chepeta Well, alt. 1540 m., May 23, 1908, Jones (Pomona, No.
39630, түре of T. incana var. prolixa).

ARIZONA: near Oraibi, 1900, Voth 19 (F); Hackberry, May 26,
1884, Jones 4516 (F, R).

16. T. arizonica Gray, Proc. Am. Acad. 16: 85. 1880; Syn.
Fl. М. Am. 1::169. 1884, and ed. 2,169. 1886and1888; Wooton
& Standley, Contr. U. S. Nat. Herb. 19: 692. 1915; Rydb. Fl.
Rocky Mts. 875. 1917, and ed. 2, 875. 1922; Tidestrom,
Contr. U. S. Nat. Herb. 25: 554. 1925.

T. arizonica X incana Jones, Zoe 2: 248. 1891.

Depressed subacaulescent and multicipital, branching from a
perennial root, forming a loose pulvinate tuft 3-4 cm. high, mi-
nutely sericeous-canescent; leaves short, obovate-spathulate, 2-3
em. long, 0.2-0.5 em. broad, seldom surpassing the foliose-fulcrate
heads; heads 1-1.5 cm. in diameter including the rays; involucre
2-3-seriate; bracts lanceolate, obtuse, with narrow, membrana-
ceous lacerate-ciliate margins; pappus of the disk-flower pluri-
setose, equalling the length of the corolla, that of the ray-flower
similar but shorter.

Distribution: southwestern Colorado and Arizona; doubtless
also in Utah.

Specimens examined:

Сотоваро: Naturita, June 1, 1917, Payson 989 (М, В); rocky
hillside, Naturita, alt. 1670 m., Apr. 21, 1914, Payson 242 (Е,
M, R); dry arroyo sides, Paradox, Montrose Co., alt. 1666 m.,
June 17, 1912, Walker 90 (M, R); Paradox, Montrose Co., alt.
1354 m., Walker 206 (R); Grand Junction, Mesa Co., May 31,
1921, Osterhout 6116 (R).

Arizona: Buckskin Mts., June 19, 1890, Jones (Pomona, No.
39642, түре of Т. arizonica X incana); Milford, alt. 1540 m.,
June 19, 1880, Jones 1794 (F, M); Ash Fork, May 13, 1883, Rusby
660 (M); “southern Utah, northern Arizona, &," Palmer 204
(M).

17. T. leptotes (Gray) Osterh. Muhlenbergia 4: 69. 1908;
Rydb. Fl. Rocky Mts. 875. 1917, and ed. 2, 875. 1922.
Т. sericea var. leptotes Gray, Proc. Am. Acad. 16: 85. 1880;

[Уоь. 14
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Coulter, Man. Rocky Mt. Region, 157. 1885; Gray, Syn. Fl.
№. Am. 1°: 169. 1884, and ed. 2, 1886 and 1888

T. lepotes Osterh. Muhlenbergia 4: 69. 1908 (doubtless a
typographical error, since the bibliographical citation in synon-
ymy refers to var. leptotes Gray).

Acaulescent; leaves pubescent with hirsute appressed hairs,
narrowly linear, attenuate at the base, 2-4 em. long, 0.1-0.2 em.
broad, surpassing the shortly pedunculate or sessile heads; heads
about 1.5 cm. in diameter including the rays; involucre 3-4-
seriate; bracts broadly linear, their membranaceous margin
scarcely wider than the lacerate ciliation; pappus of ray- and
disk-flowers similar, plurisetose, setae equalling the length of the
corolla of the disk-flower, rarely shorter; achenes only sparsely
pubescent with glochidiate-tipped hairs.

Distribution: known only from the Middle Park region of
Colorado.

Specimens examined:

Cotorapo: Middle Park, coll. of 1864, Parry (С, түре; М,
co-type, No. 121020); "Estes Park,” coll. of 1864, Parry (Е,
No. 209717); Kremmling, Grand Co., June 22, 1907, Osterhout
3487 (В); Kremmling, Grand Co., Мау 26, 1915, Osterhout 5221
(R).

18. T. exscapa (Richards.) Porter, Mem. Torr. Bot. Club 5:
321. 1894, in part, as to name-carrying synonym; Nelson in
Coulter & Nelson, Man. Cent. Rocky Mts. 509. 1909, іп part,
excluding synonym Т. sericea; Wooton & Standley, Contr. U. S.
Nat. Herb. 19: 692. 1915; Rydb. Fl. Rocky Mts. 875. 1917,
and ed. 2, 875. 1922, as to name only; Tidestrom, Contr. U. 8.
Nat. Herb. 25: 554. 1925, in part, excluding T. sericea and T.
mensana.

Aster ? exscapus Richards. Frankl. Jour. Bot. App. 7, p. 748.
1823.

T. sericea Hook. Fl. Bor. Am. 2: 16. 1834, in part, as to
synonym, Aster exscapus, and plant of Richardson; DC. Prodr.
7:278. 1838, in part; Walp. Rep. 2:575 and 957. 1843, in part,
as to Aster exscapus; Torr. & Gray, Fl. №. Am. 1: 185, 1842, in
part, as to Aster ехзсариз and plant of Richardson; Gray, Proc.

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 29

Am. Acad. 16: 85, 1880, in part, as to Aster exscapus; Syn. Fl. N.
Am. 17: 168. 1884, and ed. 2, 168. 1886 апа 1888, in part, as
to Aster exscapus.

T. Wilcoxiana Wood, Bull. Torr. Bot. Club 6: 163. 1875;
Gray, Proc. Am. Acad. 16: 84. 1880; Syn. Fl. М. Am., ed. 2, 1°:
168. 1886 and 1888; Coulter, Man. Rocky Mt. Region, 157.
1885; Rydb. Fl. Rocky Mts. 875. 1917, and ed. 2, 875. 1922.

T. sericea 8 papposa Gray, Mem. Am. Acad. N. 8. (Pl. Fendl.)
4:70. 1849; Proc. Am. Acad. 16: 84. 1880; Syn. Fl. №. Am. 12:
168. 1884, and ed. 2, 168. 1886 and 1888.

T. exscapa Wilcoxiana (Wood) A. Nels. in Coulter & Nelson,
Man. Cent. Rocky Mts. 510. 1909.

T. intermedia Rydb. in Britton's Manual Fl. N. States and
Canada, 944. 1901, and ed. 2, 944. 1905; Rydb. Fl. Rocky
Mts. 875. 1917, and ed. 2, 1922; Tidestrom, Contr. U. S. Nat.
Herb. 25: 554. 1925.

Depressed, acaulescent, rising from a woody root-stalk; leaves
linear-spathulate, 2—5 ст. long and 0.2-0.5 ст. broad, somewhat
apiculate, sparsely pubescent with fine appressed hairs, somewhat
glabrate; heads large, 2-3 cm. high and 3-5 ст. in diameter in-
cluding the rays; involucre 4—6-seriate; bracts linear-lanceolate,
obtuse, pink-tipped with membranaceous lacerate-ciliate margins;
pappus of ray- and disk-flowers similar, plurisetose of elongated
setae exceeding the length of the corolla of the disk-flower;
achenes pubescent with glochidiate-tipped hairs.

Plate 7, figs. 29-31.

Distribution: plains from southern Alberta and Saskatchewan,
south to Texas and west to Arizona; also in southern Colorado.

Specimens examined:

бостн DAKOTA: clay-hills, near Hot Springs, Мау 10, 1924,
McIntosh 21 (R);hills near Hot Springs, June 2, 1924, McIntosh
166 (R).

Kansas: Trego Co., Rich 718 (В, М,180-түРЕ of T. intermedia);
Clark Co., Curtis (M, No. 121174); arid sterile slopes near Cul-
lison, March 31, 1888, Norris (M, No. 121175); gravelly hills col-
lected within a radius of five miles of Osborne City, April 16,
1894, Shear 2 (M).

OKLAHOMA: vicinity of Camp Supply, Woodward, Арг. 5, 1925,
Wilcox (F, M); hillside, Alva, Apr. 17, 1913, Stevens 216 (M).

[Vor. 14
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Texas: infrequent, rocky slopes, alpine, March 22, 1919,
Hanson (M); rare on high mountains, Limpia, March, 1914, Allen
36 (М); calcareous bluffs, Falls Creek, Hood Co., Арг, 1884,
Reverchon 1588 (Е, М).

СогокАро: near foothills, Fort Collins, Apr. 19, 1898, Crandall
3132 (Е, К); plains, Colorado Springs, Мау 4, 1878, Jones 25 (Е);
Cheyenne Canyon, May 4, 1891, Smith (M, No. 121177); Canyon
City, Apr. 1875, Brandegee (M); Rocky Mountain Fl., 39-412
lat., 1862, Hall & Harbour 290 (Е, No. 456674; М, No. 121176);
Los Pinos, May, 1899, Baker 730 (Е, М, В); vicinity of New
Winsor, May 11, 1899, Osterhout (R, F, No. 118369) ; Evans, 1909,
Johnston 253 A (M); dry hillside, Naturita, Payson 326 (M, R).

New Mexico: Raton Mts., Colfax Co., March 23, 1848,
Gordon 42 (M); sloping hillsides on grassy plains, Santa Fe,
Apr.-May, 1847, Fendler 349 (М, co-type of Т. sericea 8 рар-
роза); near Silver City, March 29, 1889, Greene (Е).

ARIZONA: Flagstaff, May-Oct., 1902, Purpus 4 (М); vicinity
of Flagstaff, alt. 2154 m., June 2, 1898, MacDougal 31 (F, R);
Prescott, 1876, Palmer (F, No. 208510); Bright Angel, May 18-
27, 1903, Griffiths 4361 (M); Fort Whipple, May, 1865, Coues &
Palmer 365 (M); Fort Whipple, Apr. 20, 1865, Coues & Palmer
316 (M).

CANADA: at Carlton House, Richardson (Kew, түрк); sand
hills, Aweme, Manitoba, May 28, 1900, Criddle (M); quite rare
at Briggs Creek, Elbow River, Alberta, June 26, 1897, Macoun
(C); dry slope, Medicine Hat. Alberta, May 9, 1894, Spread-
borough (C); sandy hills, Aweme, Alberta, May 20, 1905, Criddle
900 (M).

19. T. sericea Hook. Fl. Bor. Am. 2: 16, pl. 119. 1834, in
part, excluding synonym; DC. Prodr. 7: 273. 1838, in part;
Nutt. Trans. Am. Phil. Soc. №. S. 7: 304. 1841; Walp. Rep. 2:
575 and 957. 1843, in part, excluding Aster exscapus; Torr. &
Gray, Fl. №. Am. 2: 185. 1842, excluding Aster erscapus and
the plant of Richardson; Gray, Proc. Am. Acad. 16: 85. 1880,
in part; Syn. Fl. М. Am. 12: 168. 1884, and ed. 2, 168. 1886 and
1888, іп part, as to plant from “Rocky Mountains in lat. 54°.”

T. mensana Jones, Contr. Western Bot. 13, p. 15. 1910.

1927]
LARSEN—REVISION OF THE GENUS TOWNSENDIA 31

T. exscapa Porter in Mem. Torr. Bot. Club 5: 321. 1894,
in part, as to synonym Т. sericea; Nelson іп Coulter & Nelson,
Man Cent. Rocky Mts. 509. 1909, in part, as to synonym T.
sericea; Rydb. Fl. Rocky Mts. 875. 1917, and ed. 2, 875. 1922,
in part, as to synonyms; Tidestrom, Contr. U. S. Nat. Herb. 25:
554. 1925, in part, as to Т. sericea and T. mensana.

Depressed, acaulescent, rising from a woody perennial caudex;
leaves linear-lanceolate to subterete, 1-4 cm. long, 0.2-0.3 cm.
wide, clustered at the base of the sessile heads, canescent with
dense closely appressed sericeous pubescence, the older leaves
conspicuously exceeding the head; heads 1-1.5 cm. in diameter
including the rays; involucre 4—5-seriate; bracts pinkish, linear-
lanceolate, acute with narrow membranaceous lacerate-ciliate
margins; pappus of the disk-flower plurisetose, equalling the
length of the corolla; pappus of the ray-flower variable from re-
duced squamellae little longer than the breadth of achene to a
condition similar to that in the disk-flower; achenes pubescent
with glochidiate-tipped hairs. Plate 6; pl. 7, fig. 32-34.

Distribution: Rocky Mountains, Alberta to Colorado and
Utah; also in the Black Hills of South Dakota.

Specimens examined:

Soutu Daxora: dry hillside near Pringle, alt. 1508 m., Apr. 19,
1909, Murdoch 3510 (Е).

Montana: Helena, 1892, Newton (F); Gallatin Co., June, 1888,
Tweedy 20 (F); Custer, Apr. 12, 1890, Blankinship 147 (M);
chiefly on the plains, Helena, May 18, 1887, Anderson (M, No.
121170).

WYOMING: Laramie, May 8, 1897, A. Nelson 2862 (Е); Laramie
Hills, Apr. 28, 1896, A. Nelson 1883 (Е); Laramie plains, Apr. 12,
1894, A. Nelson 7 (F, M); Laramie, Albany Co., May, 1899,
A. Nelson 7055 (М, В); March 8, 1860, Hayden (М, No. 121166);
March 25, 1860, Hayden (M, No. 121161); Deer Creek, west of
Fort Laramie, Apr. 15, 1860, Hayden (M, No. 121160); head of
Muddy Creek, May 4, 1860, Hayden (M, No. 121167); Shoshone
Mts., May, 1907, Hapeman (M, No. 867626).

COLORADO: near Boulder, Apr. 1901, Ramaley 654 (В); Rocky
Mts., 39-41° lat., 1862, Hall & Harbour 290 (Е, Nos. 314634 and
17721); Larimer Co., March 30, 1896, Baker 1250 (Е, М); West

[Vor. 14

32 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Cliff, Custer Co., Apr. 1888, Cockerell (F, No. 352961); steep
slope of “Тһе Mesa," alt. 2738 m., Apr. 23, 1911, Murdoch 4503
(F, M); Georgetown, June, 1873, Wolf 416 (F); Middle Park,
1861, Parry 35 (F); Denver, May, 1894, Bethel (F, No. 91820);
dry hills, Mt. Vernon, canyon, Jefferson Co., alt. 1730 m., Apr.
13, 1920, Clokey 4338 (F, R); foothills west of Fort Collins, alt.
1692 m., March 3, 1896, Baker (M, No. 121168) ; Gregory Canyon,
Boulder, Boulder Co., May 21, 1912, Уез 368 (М); infrequent,
upper mesas near Boulder, alt. 1477 m., March 19, 1921,
Hanson c160 (M); South Park, Aug., 1873, Rothrock (F, No.
304922 in part).

Отан: Theodore, Benches of the Uintas, alt. 2308 m., May 14,
1908, Jones (Pomona, Хо. 40603, түре of T. mensana).

CANADA: Rocky Mountains, Drummond (Kew, турЕ; also C,
No. 7710); clay banks, Medicine Hat, Apr. 22, 1894, Spread-
borough & Macoun (C, M); hillsides, Sweet Grass Hills, July 15,
1895, Macoun (C); Fort McLeod, Alberta, coll. of 1888, Cowdry
(C).

SPECIES ExcLUDED

Townsendia Wrighti Gray, Bot. Mex. Bound. Surv. p. 78.
1859 — Aster Wrightii Gray, Pl. Wright. Part II, p. 75. 1853.

List or ExsiccATAE

The distribution numbers are printed in italics. Тһе number in parenthesis is
the species number used in this revision.

Aiton, G. B. (1).

Anderson, F. W. (1); (19).

Arséne, Bro. G. 3387

Baker, С. F. (4); 727 (6); `728 (14); 105,
m (15); 730 (18); 1250 (19).

Baker, C. F., Earle, F. S., and Tracy,
S. M. 548 (6); 1016 (14).

Bethel, E. (19).

Bethel, E., Willey, F. S., and Clokey, I.
W. 4840 (6).

Blankinship, J. У. 301, 301a (1); 147
(19).

Brandegee, T. S. (3); 487 (4); (10); 951
(14); (18).

Brumback, Miss F. M., and Davis, Miss
С. А. 178 (4).

Burglehaus, Е. Н. (1).

Carleton, M. Н. (4).

Clokey, 1. W. (4); 4838 (19).

Cockerell, T. D. A. (19

Cotton, J. 8. 1861 (10).

Coues, Е., and Palmer, E. 523 (14); 315,
365 (18).

Cowdry (19).

Crandall, C. S. (4); 3132 (18).

Criddle, M. 900 (18).

Curtis, C. (18).

Cusick, W. C. 2295 (1); 2294 (8); 1935
(9); 2616 (10).

Dawson, С. M. (12).

1927]

LARSEN— REVISION OF THE GENUS TOWNSENDIA 33

Doty, T. 59 (1).
Drummond, Thomas (19).
Eastwood, Miss A. (14); (15)
Eggert, H. (5).

Eikenberry, W. L. 59 (1).

Engelmann, H. (4); (10).

Fendler, A. 353 (3); 157, 533 (4); 851
(11); 350, 851 bs 349 (18).

Flodman, J. H. 8. ).

Goodding, L. N. p (8); 166 (15); 1235,
561 (2); 1046 (10).

Gordon, А. L. 22 (18).

Greene, E. L. (4); (18).

Gregg, J. 327 (13).

Griffiths, D. 4361 (18); 5340 (2).

Hall, E. (4).

Hall, E., and Harbour, J. P. 289 (4);
290 (18); 290 (19

Hanson, H. C. c159 (4); (18); c160 (19).

Bspéman, Н . (19).

Hayden, F. ү. (1); (4); (19); (10).

Heller, А. A., & Е. G. 3726 (3); 3547 (14).

Howell, T. J (10).

Johnston, E. L. 263, 414b, 492 (4); 253a
(18).

Jones, A. (9).
Jones, M. А 4157 (3); 284 (4); (8); 0);
5328, 6879 (10); (11); 767 (14); 45
(15); do (16); 25 (18); (19).
Kelsey, F. D. (1).
Kern, R. H. (11); (14).
Kirkwood, J. E. 110 (13).
Leiberg, J. B. 206? (9); 34 (10).
9 (11).

Macbride, J. F., and Payson, E. B. 3095,
3225 (10

MacDougal, D. T. 31 (18).

Macoun, J. (1); (18); (19).

McIntosh, А. C. 21, 166 (18).

Marshall, W. F. 3138 (4).

Mell and Knopf (18).
Metcalfe, O. B. 413, 1434 (2); 15, 1510
(14).

Moore, Mrs. C. H. (1).

Mulford, Miss I. (10).

Murdoch, J. 3510, 4503

Nelson, A. 385, 7373, ud an 815 (10);
4622 (15); 7, 1883, 2862, 7055 (19);
7054 (12); 3031 (14).

Nelson, А., and E. 5425 (1); 5408 (15).

Nelson, А., and Macbride, J. Е. 1129
10)

(10).

Nelson, E. 815 (10).

Newton, G. W. (19).

Norris, H. W.

Nuttall, T. (12); (14); (15).

More G. E. 6116 (16); 5221, 3487
(17); (18).

Palmer, E. J. 14170, 14586 (5).

Palmer, Ed. 252, 499, (13); 204 (16);

8

(18).

Parry, C. C. 144 (1); T no 145 (12);
93, 94, (14); (17); 35 (19).

Patterson, H. N. 49 (4 4

Payson, E. B. 1160 (6); 321 (14); 989,
242 (16); 326 (18).

Payson, E. B., and L. B. 1979, 2026,
2632, 2762, 4642, 3693 (1); 4542, 1986,
2078, 2694, 4542 (8).

Payson, E. B., and Armstrong, G. M
3693 (1); sir, 3706 (8); 3205 (14).

Peck, М. E. 970 ).

Porter, Т. С. (4); E

Pringle, C. G. 6573, 7580, 9967 (13).

Purpus, C. A. 532 (7); 1845 (13); 8300
(14); 6765 Are i (18).

Putnam, J. D. (12).

Ramaley, F. 654 ngs

Redfield, J. H. 478 (4).

Reverchon, J. on (5); 1533 (18).

Rich, J. A. 718 (18).

Richardson, John (18).

Rothrock, J. T. yd (19).

Robbins, W. W. 8189 (3).

Rusby, H. H. € а

Rydberg, P. А., and Bessey, Е. А. 5132
(1); 181 (8).

Sandberg, J. Н., and Leiberg, J. B. 169

10).

Schmoll, uM M. 1348, 1212 (6).
Shear, C. L. 2 (18).

Smart, D. 92 (14).

mith. B. H. (6).

[V or. 14, 1927)

34 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Smith, E. C. 16 (1); (12); (18).
Spreadborough, W. (18
Spreadborough, W., and Мола % . (19).
Standley, P. C. 4412, 4621 (3); (14).
Starz, E. (1).

Stevens, G. W. 216 (18).

Tweedy, F. 228, 695 (1); 20 (19).
Umbach, L. M. 142, 254 (1).

Vasey, G. R. (3); 304 (4).

Vestal, A. A. 368 (19).

Visher, S. S. 2138 (4).

Von Schrenk, H. (11).

Voth, H. R. ? (14); 19 (15).
Vreeland, F. K. 639 (3).

Walker, E. P. 93 (14); 448 (15); 90, 206
16).

Walker, Mrs. S. B. (4).
Ward, L. F. 92 a 628 (11); 176 (15).
м 11).

(4).
Wolf, J. 517? an; 416 (19).
Wooton, E. O.
Wooton, E. O. = fundies, P. C. 3237

(14).
Wright, C. 1172 (14).
Young, M. S. (5).
Young, R. T. (4).

INDEX TO SPECIES

New species and combinations are printed in bold face type; synonyms in italics,
and previously published names in ordinary type.

г соо ЕЕ cic. 39
БОР ИИ; M 19
са, о и 21
AE Wrighti: В 32
ШЫРТ Құла УТ ЛУТ 28
ОН ТУГЕ 28
gS асым 19
ЖИІ Дс 19
о И АРА 19
о, O 19
ONE BRO. 21
TOWRA 8
о ccd sue x CREE ERA
лы И и 22
РОСТУ 21
a АО. 27
arizonica X incana. 27
КЕ сл ни 16
КИЕ, OTe On РС OP 22
ere С 17
а КЕС 12
1. c TR 28
ОРА р 31
exscapa Wileoziana............. 29
IONE ЖИККЕ ОЛЛО M 24
^^ ЖОЧИ E КА 19
€ var. соттитів........... 19
ПОРО ee eek eee 11
ime TEE залға ST ees 26
с. I Sic x Do ыт ITI 16

он Ls УБ edis калеа 13
Tr MEDI CCP 25
incana var. ambigua............ 26
incana var. proliza............. 26
CONG КТГ” 29
ИЛАШ: осуз ere Are T YT 28
ee, ee errr 27
WAG Суы асы ДЫ 80
иа газа аруа ерге 23
КО УЛУГУ КТУ 17

EER EVEN РІ» УЛ Т 9
Рату п. Рю 22
PIR РЕТТИ Tf
BIEN. ane ee yer 16
ос: 2а 212255252222. 20
scapigera var. ambigua ......... 21
scapigera var. Merci re 21
и EET ERSTE Y TUTTO TY TTL 30
dons MER TRIGO TITEL 28
sericea var. 1ершез.............. 27
sericea B рарроға,;.:........... 29
кокс е vies Fou apa 22
| ПАРА 24
MNA Байт: uaa ox eI. 18
b land БЕ, ее. 19
BEN. cono о В 15
VNEEUN о 12
О тан 18
а ПОРА 29

[Уоь. 14, 1927]
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 2
Townsendia formosa Greene
New Mexico
From a co-type, Metcalfe No. 1434, in the Missouri Botanical Garden Herbarium.

ANN. Mo. Вот. Ganp., Vor. 14, 1927

Рі,лте 2

лома де.
ФАК LM
i "ua

г
4

"ә
M 12:35 Д
bs

/ &. 1 i
( L4 fec of ;
]-^ J
4% erb af
Noal FY PLANTS OF SEW MEXICO
DP боа феа ч ЖУЙЕЫ, еле
: ‹
а.
Ж x o^ nam
>

LARSEN—REVISION OF THE GENUS TOWNSENDIA

[Vor. 14, 1927]
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 3
Townsendia texensis Larsen
Northwestern Texas

From the type specimen, Eggert, in the Missouri Botanical Garden Herbarium
No. 121021.

PLATE 3

ANN. Mo. Вот. Garp., Vor. 14, 1927

ше

елки

EGGERT, HERBARIUM AMERICANUM,

idi

i

LARSEN—REVISION OF THE GENUS TOWNSENDIA

[Vor. 14, 1927]
40 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 4
Townsendia strigosa Nuttall

. Achene of the ray-flower with pappus attached, Х 5.
Corolla of the ray-flower, Х 5.

Fig. 1. Style-branches of the они, greatly enlarged.
Fig. 2. Style-branches of the disk-flower, Х 5.
Fig. 3. Corolla of the disk-flower ары. dio stamens, X 5.
Fig. 4. A seta of the disk-flower, greatly enlarged.
Fig. 5. Disk-flower, X 5.
Fig. 6. Glochidiate-tipped hair, greatly enlarged.
Fig. 7. Style-branches of the ray-flower, X
Fig. 8. Style-branches of the ray-flower, greatly enlarged.
Fig. 9. A seta of the ray-flower, greatly enlarged.

0

L

12.

"i
"evt
ка =

Fig. otograph of the type specimen in the Herbarium of the Academy of
Natural imi Philadelphia.

PLATE 4

Ann. Mo. Вот. Garp., Vor. 14. 1927

РОА

n up =.
„=“, айма. Ч
(Ж ысы Б:

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с :
V Ёла Pr А-
47
A [on bt
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LARSEN—REVISION OF THE GENUS TOWNSENDIA

42

[Уох. 14, 1927]
ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 5

T. eximia Gray.

ig. 13. Ray-flower, Х 214.

. 14. Pappus of the ray-flower, greatly enlarged.

. 15. Disk-flower, showing a single elongated seta in the pappus, X 214.
. 16. Pappus of the disk-flower, greatly enlarged.

. 17. Achene of the disk-flower with the corolla and pappus attached.

. 18. Glochidiate-tipped hair.

. 19. Ray-flower, X 214.

Fig. 20. тэ рни of the ray-flower, greatly enlarged.
. 21. Disk-flower, Х 214.

. 22. Style-branches of the disk-flower, X 5.

. 23. Style-branches of the disk-flower, greatly enlarged.
ig. 24. A seta of the pappus, greatly e

T. florifer Gra

Fig.
Fig.
Figs.

y.
25. Bidentate hairs, greatly enlarged.
26. Disk-flower, X 5.

27 and 28. Ray-flowers from the same head showing the variability of the
pappus, Х 5.

ANN. Mo. Вот. Ganp., Vor. 14, 1927 PLATE 5

^ At
А 9H м Ы Ma »
м ~i 4 C
N N /
s \ /
NC A. ху 14

i^
SN
XX A
45 М\ М,
ЕА لي‎
Le X
== N
~ ұл,

17

|

LARSEN—REVISION OF THE GENUS TOWNSENDIA

[Vor. 14, 1927)
44 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 6
Townsendia sericea Hook.

From a co-type, Drummond, in the National Herbarium of the Victoria Memorial
Museum, Ottawa, Canada, No. 7710.

ANN. Mo. Вот. Ganp., Vor. 14, 1927

PLATE 6

Ex Herbario Musei Britannici.

pwn eper Sca S4 tr tren Rook Б;
“” fe ж
C feo ез Cey coce Фе Сэм kass
„Гаа ғас Jana ew ~
» / a /
eol 2726. me pu

LARSEN—REVISION OF THE GENUS TOWNSENDIA

[Vor. 14, 1927]
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 7
Townsendia exscapa (Richards.) Porter.

Fig. 29. Photograph of type specimen in Kew Herbarium.
Fig. 30. Ray-flower, X 5.
Fig. 31. Disk-flower, Х 5.

endia sericea Hook.
Fig. 32. Photograph of type specimen in Kew Herbarium.
Fig. 33. Ray-flower, X 5.
Fig. 34. Disk-flower, 5.

PLATE 7

ANN. Mo. Вот. Garp., Vor. 14, 1927

\%

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zw
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1

LARSEN—REVISION ОЕ THE GENUS TOWNSENDIA

STUDIES ON SOUTH AMERICAN LABIATAE. ПП
SYNOPSIS OF THE GENUS SATUREIA

CARL CLAWSON EPLING
Instructor in Botany, University of California, at Los Angeles
Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany
of Washington University
SATUREIA (Tourn.) L.

Gardoquia Ruiz et Pavon, Prodr. 86, pl. 17. 1794.

Rizoa Cav. in Anal. Cienc. Nat. 3: 133. 1801.

Xenopoma Willd. in Ges. Naturforsch. Fr. Berlin, Mag. 5: 399.
1811.

Thymus Kunth in Humboldt et Bonpland, Nov. Gen. et Sp.
Pl. 2: 315. 1817 (non L.); Swartz, Prodr. Veg. Ind. Occ. 89.
1788.

Micromeria Benth. in Bot. Reg. 15: sub pl. 1282. 1829.

Herbae perennes vel suffrutices in regione nostra, habitu di-
verso internodiis saepius foliis aequilongis vel brevioribus; foliis
maximam partem parvis, non rugosis, venis saepe prominulis,
margine integra vel subserrata; floribus in cymulis nunc patenti-
bus nunc densis in foliorum superiorum axillis, rarius solitariis;
calycibus tubulosis, fere cylindratis maturis non auctis, saepius
13-venis, venis prominulis, dentibus aequalibus vel patenter bi-
labiatis, erectis, intus saepius hirsutis; corollae tubo saepius
exserto, frequenter subincluso, superne gradatim ampliato, intus
frequenter villoso; labro erecto, subintegro vel bifido, breviore,
labiolo quam labro paulo Ep c trifido, lobo medio majore et
subpatente; staminibus 4, supra tubi medium saepius sitis,
didymis, posticis paulo brevioribus, filamentis glabris, connectivo
crasso late divaricatis, rarius subparal-
lelis; stylo paulo exserto, inaequaliter bifido; nuculis oblongis,
saepius apice obtusis, rarius apiculatis.

Satureia in the region under consideration is a genus of а
certain diversity of habit and flower character, the species of

1 Issued June 8, 1927
ANN. Mo. Вот. GarD., Vor. 13, 1927 (47)

[Vor. 14
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN

which, however, form a continuous and interlocking series, so
that subgeneric lines are difficult to distinguish. Аз concluded by
Briquet,! the proposed genera Gardoquia, Xenopoma, and Micro-
meria are confluent, and the study of species herein described
and unknown to him have strengthened this conclusion.

The primary differentiating characters of the genus Gardoquia
were the long, recurved corolla-tube, the erect emarginate upper
lip of the corolla, and the didymous cordate anthers. The genus
was maintained by Bentham with the following note “Genus а
Micromeria non nisi longitudine corollarum, limbique lobis sub-
erectis, distinctum,” * while Micromeria (including X enopoma) in
turn was separated from Satureia on the basis of the narrower
calyx and the character of its venation. With the present series of
species these characters are insufficient. Rizoa is wholly synony-
mous with Gardoquia, being based on Gardoquia multiflora R. & P.

CoNsPECTUS SPECIERUM

А. Flores in axillis solitarii, pedicellis calyces subaequantibus vel longioribus
elatis, bracteolis «ренді З P E tea ОЬ ЭҢ 1. S. Brownei
AA. Flores 3-6 et ultra in axillis vel solitarii, pedicellis quam calycibus
brevioribus, bracteolis praesentibus
B. Herbae же ааны humiles, altitudine 5-20 em., floribus in axillis

solita
C. addi p pedicellorum medium sita; folia obovata, margine

integra.
D. Calyces 13-venis, dentibus duobus anticis quam posticis sub-
on gioribus, erectis раза ее 3 3. S. pusilla

Е СВО 38. 8. Darwinii

CC. — ad — basim sita; folia ovata vel —
vata, margine leniter crenata...................... 31. S. nubigena
BB. Suffrutioes vel ition altitudine 30-200 cm., floribus 1—6 et ed in

C. Calyces 1.5-3.5 mm. longi, corollae tubo incluso vel paulo exserto
rarius quam calyce duplo longiore.
D. Calyces 1.5-2 mm. longi, corollae tubo incluso.

E. Folia utrinque glabra, anguste lanceolata, acuta...... 37. S. axillaris
EE. Folia puberula vel pubescentia, oblonga, obtusa. . .39. S. oligantha

DD. Саусез 3-3.5 mm. longi, corollae tubo breviter exserto.
E. Folia ovata vel rotundato-ovata, leniter crenata....... 34. S. vana

1 Briquet, J. in Engler u. Prantl, Nat. Pflanzenfam., ed. 1, IV. Abt. 3a, 296. 1897.
? Bentham, G. in DC. Prodr. 12: 235. 1848.

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 49

Е. Folia lanceolata, acuta; flores saepius tres in axillis.
es Cree > Ко л. 36. S. simulans
FF. Folia oblanceolata, oblonga vel ovata, obtusa; flores saepius

solitari.
G. Calyces subturbinati; corollae 7 mm. longae; folia ovalia

vel elliptica, 3—5 mm. longa................ 38. 8. brevicalyx
GG. Calyces tubulosi; corollae 9 mm. longae; folia aret
olata vel oblonga, .5-1.5 cm. longa.......... phe еген

СС. Calyces 4.5-10 mm. (3.5-4 mm. in S. rigidula et ics uii
mm. longi, corollae tubo saepius longe exserto quam ub di
duplo triplove EI

D. Folia 2-5 cm. longa, sat tenuia, ovata vel elliptica, crenato-
serrata; flores 3-3.5 em. longi
E. Flores pedicellis gracilibus caly baequantil levati; caly-
сз о ААА nance ee be 3 2. S. multiflora
EE. je €— quam calycibus brevioribus; — extus
РЕТТЕРІ 2 . 8. elliptica
DD. Folia таго 25 ст. longa, saepius .5-1.5 cm. vel ye es
diversiore saepius integra; flores saepius 2-2.5 cm. longi.

E. Folia obovata vel oblanceolata, omnino obtusa; یا‎ dentes
acuminati (acuti in S. guamaniensis) etiam subaristati.
F. Folia rhomboideo-obovata, vel subrotun
G. Folia rhomboideo-obovata, subtus var Й flores fere
lon 6. S.

3 em. lop... SECURUM oe as Matthews
GG. Folia rotunda, subtus parce villosa; flores vix 6 7e Fps
44 0а тал» «ve Г РО, O S. breviflora
FF. Folia oblanceolata.
С. Flores in axillis solitarii................... 5nd Miet noo
GG. Flores 3-7 in axillik. s.a АО aa . S. Gilliesii

EE. тош. eiliptico-lanseclatt, ovata, vel linearia, raro hne
ubaristati (S. Lindeniana,

rigidula, glabrata).
Е. Folia 1-2.5 ст. longa, acuta (frequenter obtusa in S. tomentosa)
pagina superiore subglabra, inferiore incana (sericea in
S. acutifolia).

С. Calyces vix 5 mm. longi, tubo extus villosissimo. .12. 5. discolor
GG. Calyces 6-8 mm. longi, tubo extus appresso-villoso vel
puberulo.
H. Folia ovata, crenata, subtus ee .9. S. tomentosa
НН. Folia elliptico-lanceolata, subinte,
2 na superiore sericea, ена tomentosa; E

Ца... а. У ел . S. acutifolia
ad Pagina superiore glabra, inferiore tome margo
ӘТЕР” . S. Pavoniana

FF. Folia 2-15: mm. longa utrinque glabra MEHR in S. An-
drei) saepius subnitida, venis prominulis, ascendentibus,
rectis, mom margo leniter vel nihil revoluta.

а

8.
J. Calycis dentes subaristati, 2 mm. longi....... 17. S. glabrata

[Vor. 14
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN

JJ. 2. dentes lanceolati, acuti, 1-1.5 mm. longi.
. Folia utrinque acuta, internodiis multo longiora.

ЖАРҒЫ МДҰ И СА, 18. S. taxifolia
KK. Folia utrinque obtusa, internodiis —Ó
E КЕКСЕ ТТ УТТА К SENG Рур 8 pallida
HH. Folia praecipue subtus puberula.............. S. Andrei
GG. Folia 2-5 mm. longa.

H. Folia in basi angustata, haud cordata.
J. Margo integra; calycis dentes erecti...........
JJ. Margo serratula; calycis dentes patentes. . tá Ж ech
HH. Folia in basi cordata.
J. Corolla 18-20 mm. longa; calycis dentes tres postices

о Хо” МАЛИНО ird RC 23. S. connata
JJ. — 20-25 mm. longa; calycis dentes на. postices
ere ad. medium bal... TA 2. S. Jamesoni

FFF. Folia ^ mm. longa, pubescentia, saepe Rm rien
superiore glabra in S. Lindeniana), venis obscuris; margo
saepius valde revoluta.

G. Calycis dentes duo antici subulati, fere aristati.
H. Dentes antici 3 mm. к posticos "e
. S. Lindeniana
HH. Dentes. anti 1. bn mm. „кай, обо та
а Тени 26. S. rigidula

H. Folia linearia, utrinque sericeo-villosa; cms { ж

рае рота РС ЧО 7. S. sericea
HH. Folia ovata, Pr guperiore glabrata vel Me ied
calyces 5 m ngi.
J. Foliorum maii valde revolutae; pagina inferiore
Г СОС TE TOITERTEIO P T 28. S. revoluta
JJ. Foliorum margines revolutae; pagina inferiore pu-
berula, venis prominulis................. 24. S. argentea

Sect. HEsPEROTHYMUS Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. За, 300. 1897.

Мете sect. Hesperothymus Benth. Lab. Gen. et Бр. 371.
1834.

Herbae prostratae, caulibus repentibus, foliis ovato-rotundis,
subglabris; floribus in foliorum axillis solitatim dispositis, pedi-
cellis calyces maximam partem superantibus, rarius paulo brevi-
oribus, bracteolis duobus nunc praesentibus nunc absentibus;
calycibus tubulosis, dentibus erectis subaequalibus, obscure bi-
labiatis, intus hirsutis; corollae tubo paulo exserto.

This section seems to the present author to be most nearly

1927]
EPLING—SOUTH AMERICAN LABIATAE. ІП 51

allied to sect. Gardoquia (Ellipticae) through Satureia Douglasii
(Benth.) Briquet.

1. Satureia Brownei (Swartz) Briq. in Engler u. Prantl, Nat.
Pflanzenfam., ed. 1, IV. Abt. За, 300. 1897.

Herba жен саше prostrato, repente, ramis filiformibus,
ascendentibus, glabris vel tenuiter pilosis, quadratis, angulis
acutis, submarginatis; foliis saepius tenuiter membranaceis minus
quam internodiorum longitudine, saepius .5-1 cm. longis, rarius
1.5 cm., ovato-rotundatis, apice obtusioribus, in basi saepius sub-
truncatis et abrupte in petiolo angustatis, margine sinuato-
степафа vel subintegra, omnino glabris vel subtus sparse sed
patenter pilosis, petiolo gracillimo laminam aequante vel sub-
nullo; floribus saepius solitariis, rarius duobus in axillis; calycibus
glabris, rarius pilosis, 3-4 mm. longis, 13-venis, dentibus .8-1
mm. longis, subaequalibus, ovato-triangulis, acutis, ciliolatis,
tubo intus ad dentium basim piloso-annulato, pedicello filiforme
ealyce aequilongo vel saepius longiore; corollis violaceis, fauce
saepe variegatis, 4.5-5 mm. longis, tubo superne ampliato, intus
glabro, labro 1 mm. longo, emarginato, labiolo longo 1.5 mm., lobo
medio majore, patente, frequenter emarginato; staminibus didy-
mis, supra tubi medium sitis, posticorum filamentis saepius 1 mm.
longis, anticorum 2 mm., antheris .7 mm. latis, thecis paulo di-
vergentibus; stylo e éctollu subexserto; nuculis .8-1 mm. longis,
oblongis, atris.

Subsp. eubrownei nom. nov.

Thymus Brownei Swartz, Prodr. Veg. Ind. Occ. 89. 1788, et
Fl. Ind. Осс. 2:1011. 1800; Benth. Lab. Gen. et Sp., 372. 1834.

Ramulis foliisque fere glabris; petiolis laminas aequantibus vel

arte dimidia brevioribus; foliis nec sessilibus; florum pedicellis
saepius calycibus aequilongis vel paulo brevioribus.

Specimens examined:

VENEZUELA: Сайрап, 1250 m., June 7, 1885, Jahn 201 (known
as *Poleo") (US)!; Paramo de la Sal, 3000 m., Mérida, Sept. 1,

1 The following abbreviations are used herein: ASP, Academy of Natural Sciences
of Philadelphia; FM, Field Museum of Natural History; GH, Gray Herbarium;

MBG, Missouri Botaniéal Garden; NY, New York Botanical Garden; UC, University
of California; US, United States National Herbarium.

[Vor. 14
52 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1921, Jahn 569, 610 (US); between Antimano and Aguas Negros,
900-1500 m., Apr. 6-7, 1913, Pittier 6017 (US); prope coloniam
Tovar, 1854—5, Fendler 869 (MBG; GH ; NY); Paramo del Tam-
bor, 2400 m., Mérida, Nov. 14, 1921, Jahn 736,738 (US) ; between
Colonia Tovar and Lagonazo, in meadows and forests, 1700-
2300 m., trailing, forming colonies on road, Feb. 21, 1921, Pittier
9262 (US); Paramo del Jabón, 3000-3200 m., Oct. 2, 1910, Jahn
41 (US); Paramo de Рійапео, 2600 m., March, 17, 1915, Jahn
410 (US).

Сотомвтл: Sabana de Bogotá, Bro. Ariste-Joseph А 908 (US);
Cuesta de Tocotá, road from Buenaventura Cali, western Cordil-
lera, 1500-1900 m., Dec. 1905, Pittier 724 (US); grassy wayside,
north of Caramanta, Antioquia, 2000-2200 m., repent herb with
odor of Hedeoma, Sept. 19, 1922, Pennell 10777 (US; ASP);
forest, 2000-2500 m., Las Minitas, south of Caldás, Antioquia,
Sept. 21-22, 1922, Pennell 10941 (US; ASP); forest, 2000-2500 m.,
Huila, Aug. 1-8, 1917, Rusby & Pennell 625 (US; MBG; GH;
NY); meadow, 2000-2100 m., Balsillas, Huila, Aug. 3-6, 1917,
Rusby & Pennell 727 (US; MBG; GH; NY); forest, 2600-2900
m., Pinares above Salento, Caldas, Aug. 2-10, 1922, Pennell 9243
(ASP; US); cliff near Rio San Andreas, 2500-2800 m., Caloguala,
Coconuco, El Cauca, June 14, 18, 1922, Pennell 7157 (US; ASP);
Quaranda, July 8, 1876, Andre 1016, mountains southeast of
Bogota, 2800 m., June 6, 1875, Andre 1016 (NY); Popayan, 1600-
2000 m., Lehmann B.T. 1141 (blooms in Feb.) (NY); grassy open,
1500-1600 m., Rio Quindio, Salento, Caldas, July 27-30, 1922,
Killip & Hazen 9040 (US; ASP); around Huila, Rio Paez Valley,
El Cauca, 1600-1900 m., “а diminutive plant on wet sand and
in moss; fl. purplish pink," Jan. 1906, Pittier 1242 (US) ; wet slope,
open forest, Susumoco, southeast of Quetamo, 1200-1400 m.,
Sept. 5, 1917, Pennell 1735 (cor. violet) (NY); Popayan, Hartweg
1335 (NY); moist soil near stream, 2000-2200 m., San Isidro,
Puracé, El Cauca, June 10-11, 1922, Pennell & Killip 6430 (US;
ASP) ; field, 2000-2200 m., Santa Elena, above Santuario, Caldas,
Sept. 7-13, 1922, Pennell 10592 (“corolla pale amparo-purple
with markings of violet-purple") (US; ASP); thicket below San
Jose, 2300-2500 m., San Antonio, El Cauca, July 1, 1922, Pennell
7654 (US; ASP).

1927]
EPLING——SOUTH AMERICAN LABIATAE. III 53

Ecuapor: in Andibus Ecuadorensibus, 1857-9, Spruce 5091
(GH); vic. of Huigra, Sept. 12, 1918, Rose 22627 (known as “ Po-
leo"; growing in water; strong odor of pennyroyal) (US).

Sect. Garpoquia Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. За, 300. 1897.

Gardoquia Ruiz et Pavon, Prodr. 86. pl. 17. 1794.

Frutices fruticulive ramulis saepius ascendentibus, brevioribus;
foliis saepe fasciculatis, forma diversioribus, maximam partem sub-
integris, glabris vel pubescentibus; floribus 1-6 in foliorum axillis,
pedicellis singulis bracteolis duobus ornatis (S. multiflora exclusa)
quam calycibus brevioribus elatis; calycibus tubulosis, cylin-
dratis, dentibus maximam partem bilabiatis, tribus posticis an-
ticos superantibus, acutis, etiam subaristatis, intus nudis vel
hirsutis, saepius erectis; corollae tubo maximam partem calycem
duplo triplove superante.

Gardoquia merges into Xenopoma imperceptibly in habit, as
illustrated by S. breviflora and S. tomentosa or S. boliviana and
S. Gilliesii, and in floral characters, as in S. rigidula, breviflora,
argentea, boliviana and vana. Тһе closest alliance between the
two sections appears to lie between S. boliviana and members of
the subsection Obovatae, or between S. breviflora, tomentosa and
vana.

A. Ellipticae
Foliis 2-5 cm. longis, sat tenuibus, ovalibus vel ellipticis, glabris
vel subglabris, crenatis; floribus 3-15 in cymulis in foliorum
axillis, corollis 3-3.5 ст. longis.

2. Satureia multiflora Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia multiflora Ruiz et Pav. Syst. Veg. 149. 1798; Benth.
Lab. Gen. et Sp., 398. 1834, et in DC. Prodr. 12: 235. 1848.

Rizoa ovatifolia Cav. Anal. Cienc. Nat. 3: 133. 1801.

Frutex aromaticus erectus, ramis altitudine circa 1 m., ramulis-
que subglabris, purpureis, quadratis, angulis obtusis, inter-
nodiis quam foliis sat longioribus, ramulis lateralibus brevibus,
gracilibus; foliis membranaceis, 3-5 ст. longis, 1-2 cm. latis,
lanceolatis vel ovatis, apice obtusis, in basi rotundatis, margine

[Vor. 14
54 ANNALS OF THE MISSOURI BOTANICAL GARDEN

convexa, leniter crenata, crenarum culminibus inter se 4-5
mm. distantibus, petiolis gracilibus, 5-12 mm. longis, glabris;
floribus 3-7-15 in cymis laxis, raro solitariis, pedunculis gracil-
limis .5-1.5 cm. longis elevatis, bracteolis parvis 1-1.5 mm. longis
subulatis ornatis, pedunculis secondariis .3-1 cm. longis; calycibus
membranaceis, 8 mm. longis, in basi paulo angustatis, extus
glabris, fauce intus nudis; dentibus circa 1 mm. longis, lanceolatis,
acutissimis, subaequalibus, duobus anticis tamen paulo longi-
oribus approximatisque; corollis ut videtur purpureis, 20-30 mm.
longis, maximam partem circa 25 mm., tubo extra calycem multo
dilato, extus puberulis vel subglabris, intus pilosis, labro longo
cirea 4 mm., retuso, labiolo aequilongo lobis rotundatis medio
lateralibus paulo longiore, in basi angusto; staminibus didymis,
paulo supra tubi medium sitis, anticis corolla longioribus, posticis
labro subaequilongis, thecis late divergentibus, .7-.8 mm. longis;
stylo paulo exserto; nuculis oblongis, fuscis, 1.2 mm. longis.

Specimens examined:

Снил: unknown collector 245 (NY); 1828-34, Gay 148 (NY);
Gay (СН); Chanco, Reed (GH); Valdivia, Ufergebüsch des Calle-
Calle, Jan. 17, 1898, Buchtien (US) ; Chiquayante (? Chiguailante),
Feb. 19, 1892, Kuntze (US); Budi, Jan. 1923, Bro. Claude-Joseph
2016 (US); Temuco, Jan. 1920, Bro. Claude-Joseph 1043 (US);
bushy slopes above Aranco, 50-100 m., March 6, 1925, Pennell
12953 (GH).

ARGENTINA: Panguipulli, July 1924, Bro. Claude-Joseph 2627
(US).

3. Satureia elliptica Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, ТУ. Abt. За, 300. 1897.

Gardoquia elliptica Ruiz et Pavon, Syst. Veg. 149. 1798;
Benth. Lab. Gen. et Sp. 399. 1834; et in DC. Prodr. 12: 235.
1848.

Stachys speciosa Hook. Bot. Mise. 2: 235. 1831.

Frutex erectus odoratissimus, ramosus, ramis teretibus cortice
discedente, ramulis glabratis, nitidis, quadratis, angulis acutis;
internodiis plerumque foliis aequantibus vel longioribus; foliis
20-30 mm. longis, 10-15 mm. latis, ellipticis, apice obtusis, in
basi acutiusculis et ad petiolum 3 mm. longum angustatis,

1927]
EPLING— SOUTH AMERICAN LABIATAE. ПІ 55

membranaceis, planis, rugosis, margine supra medium breviter
serrata, utrinque glabris vel sparse ciliatis, subtus pallidiore,
venis prominentioribus; floribus 4-12 in axillis, pedunculis brevis-
simis elatis, bracteolis linearibus, quam pedicellis brevioribus
ornatis; calycibus 8-11 mm. longis, extus villosis, fauce intus
nudis, dentibus duobus anticis fere 3 mm. longis, lanceolato-
acuminatis, approximatis, tribus posticis connatis, cirea 1 mm.
longis; pedicellis 3-5 mm. longis, villosis; corollis coccineis, in-
terdum flavis, 35-40 mm. longis, extus hirtellis, tubo extra
calycem multo dilato, intus ad basim hirsuto, labro 4.5-5.5 mm.
longo, leniter emarginato, ovato, labiolo subaequilongo, lobo
medio lateralibus paulo longiore; staminibus supra tubi medium
insertis, didymis, anticis labro longioribus, posticis aequilongis,
thecis late divergentibus, 1 mm. longis; stylo e corolla 10 mm.
exserto, ramis 1 mm. longis, planis, acutis; nuculis 2 mm. longis,
oblongo-obovatis.

Specimens examined:

Рево: Obrajillo, Wilkes Exp. (US); rock ledges at cascades of
Rio Chillon, above Obrajillo, Lima, 3100-3300 m., June 13-23,
1925, Pennell 14401 (СН); abundant on open rocky slopes,
Huaros, Lima, 3200-3600 m., June 23, 1925, Pennell 14734 (GH).

4. Satureia Loesneriana Mansfeld in Bot. Gart. Berlin-Dahlem,
Notizbl. 9: 287. 1925.

"Frutex 1 m. altus. Rami juniores quandrangulares, brevis-
sime puberuli. Foliorum lamina 9-22 mm. longa, 3-8 mm. lata,
elliptica vel rarius oblonga vel obovata, apice acuta, brevissime
mucronulata, basi in petiolum brevissimum angustata, margine
+ remote et breviter dentata vel integra, subrevoluta vel plana,
penninervis, utrinque glanduloso-punctata, nervis utrinque prom-
inulis (vel supra vix conspicuis), initio utrinque breviter puberula,
postea praecipue supra glabrescens vel glabra. Verticillastri
usque 16-flori. Calyx circ. 5.5 mm. longus, bilabiatus, labio
tridentato dentibus 0.6 mm. longis, labiolo dentibus 1.5 mm.
longis, dentibus extus et intus brevissime puberulus. Corolla
circ. 15 mm. longa, labio sat profunde emarginato 2.5 mm. longo,
labiolo tripartito 3 mm. longo, extus pubescens, tubo intus antice
piloso. Stamina didynamia + exserta, filamenta antica 5, pos-
tica 3.5 mm. longa, antheris divergentibus. Stylus exsertus.

[Vor. 14
56 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

“Peru: Prov. Huamachuco, Depart. Libertad, über Huama-
chuco, Grassteppe mit eingestreuten immer oder regengrünen
Straüchern, 3400-3500 m. ü. M., 1 m. hoher Strauch, Blüten
violett (Fl. 3. VII. 1914.—WEBERBAUER n. 7008!).

“Die Art gehórt in die Nahe von S. elliptica (Ruiz et Pav.)
Briq. und S. Matthewsii Briq., die erstere Art weicht schon durch
eine viel längere Korolla ab, die letztere besitzt nach der Be-
schreibung eine längere, innen kahle Krone und kürzere Kelch-
zühne."

Mihi ignota; ad S. ellipticam affinis videtur. Inquire praeterea
іп S. rugosani Виа. in Engler u. Prantl. Nat. Pflanzenfam., ed. 1,
IV. Abt. За, 300. 1897.

B. Obovatae

Foliis saepius 1-1,5 cm. longis, obovatis vel oblanceolatis,
glabris vel pubescentibus, margine integra vel crenata; floribus
1-6 in verticillastris sat densis in foliorum axillis, corollis circa
2-2.5 cm. longis (6 mm. in S. breviflora).

5. Satureia Gilliesii Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. За, 300. 1897.

Gardoquia Gilliesii Grah. in Edinb. Phil. Jour. 1831: 377. Sept.
1831; Benth. in DC. Prodr. 12: 235. 1848.

G. chilensis Benth. in Hook. and Arn. Bot. Beechey's Voyage,
58. 1841.

Satureia chilensis Briq. in Ann. Conserv. et Jard. Bot. Genéve
2:191. 1898.

Fruticulis suaveolens altitudine ad 1-1.5 m., caulibus in basi
lignosis, procumbentibus, ramis teretibus, cortice discedente,
ramulis pubescentibus, quadratis, angulis obtusis, internodiis
ramulorum sterilium quam foliis brevioribus, his itaque densis,
eis fertilium sat distantibus; foliis 7-10 mm. longis, rarius 10-20
mm., anguste oblanceolatis apice rotundatis, in basi ad petiolum
brevem attenuatis, margine integra, revoluta, utrinque subglabris,
rarius patenter puberulis; floribus 3-7 saepius circa 5 in axillis,
pedunculis .5-2 mm. longis, bracteolis foliis conformalibus
minoribus; calycibus 7-12 mm. longis, in basi paulo angustatis,
extus saepius puberulis, fauce intus nudis, dentibus 2-3 mm.
longis, lanceolato-acuminatis, subaristatis, duobus anticis quam

1927]
EPLING—SOUTH AMERICAN LABIATAE, III 57

posticis tribus saepe longioribus approximatisque; corollis circa
12-18 mm. longis, coccineis, extus pubescentibus, tubo ampliato,
intus pilosis, labro longo 2-3 mm., emarginato, sinu .5 mm. pro-
fundo, labiolo paulo longiore, lobo medio lateralibus superante,
in basi angustato; staminibus didymis, supra tubi medium sitis,
posticis vix e tubo exsertis, anticis corolla aequilongis, thecis
late divergentibus, .5 mm. longis; stylo paulo exserto; nuculis
non visis.

Specimens examined:

Рево: Dombey 291 (СН).

CHILI: Valparaiso, June 1885, Rusby 1061 (GH); Valparaiso,
July 1851, Gillies (GH); Valparaiso, Apr.-July, 1856, Harvey
(GH); ex regione inferiori Andium Chilensium juxta tepidaria
Cauquenes, 3000-5000 m., May 14-17, 1882, Ball (GH; NY);
Maule, Reed (GH); Valparaiso, Arnott (NY); no locality, 1884-85,
Statin (NY); no data, Wilkes Exp. (ASP; US ; NY); Santiago,
montaíia, Dec. 5, 1920, Bro. Claude-Joseph 1321 (US); Valpar-
aiso, Eights (US); Valparaiso, in Gebüschen, Sept. 5, 1895, Buch-
tien (US); Isle of St. Marys, Eights (US); mountains east of San-
tiago, 769 m., Dec. 27, 1900, Hastings 310 (US ; NY; UO); San-
tiago, Jan. 1919, Bro. Claude-Joseph 840 (US); Valparaiso, Feb.
1922, Bro. Claude-Joseph 1619 (US); Rio Blanco, Jan. 1924,
Bro. Claude-Joseph 2479 (US); Valparaiso, Mertens (MBG);
Santiago, Jan. 1920, Bro. Claude-Joseph 956 (US) ;near Valparaiso,
June 1885, Rusby 1061 (ASP; US); no data, Styles (ASP); in
fruticetis apricis collium St. Jago, Quillota, Feb., 1829, бері.-
Oct., 1829, Bertero 291, 1016 (MBG; GH; NY); Prov. Curicé,
hacienda Monte Grande, 1000 m., Гес., 1924, Werderman 547
(UC); Cerro Echaurrina (San Fernando), Prov. Colchagua, Oet.,
1925, Montero 13 (GH).

6. Satureia Matthewsii Briq. in Ann. Conserv. et Jard. Bot.
Genéve. 2: 189. 1898.

Frutex ut videtur elatus, ramis ascendentibus, teretibus, gla-
bris, mox ligneis, ramulis gracilibus, glabris, nodis inter se 3-5
mm. vel in ramulis sterilibus 1-1.5 em. distantibus ; foliis 1-1.5
cm. longis, elliptico- vel rhomboideo-obovatis, frequenter subro-
tundatis, apice saepius obtusis, in basi rotundato-extenuatis,

ГУ ол. 14
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN

utrinque glabris, dense glanduloso-punctatis, margine integra,
revoluta; verticillastris 1-3 floribus, in foliorum ad ramulorum
apices axillis approximatis, bracteis lineari-spatulatis, quam
pedicellis paulo longioribus; calycibus fere 1 cm. longis, anguste
tubulosis, extus puberulis, fauce intus nudis, patenter bilabiatis,
labia superiore erecta, 3 mm. longa, dentibus subulatis ad medium
connatis, inferiore 2.5 mm. longa, dentibus patentibus, subulatis,
ad basim connatis, pedicellis 2 mm. longis elatis; corollis circa
2 ст. longis, tubo 18 mm. longo, intus subnudo, extus hirtello,
superne ampliato, labro 3 mm. longo, emarginato, labiolo paulo
longiore, lobo medio majore; staminibus supra tubi medium sitis,
didymis, breviter exsertis, thecis valde divergentibus; stylo 5-6
mm. longo, exserto; nuculis non visis.

Specimens examined:

Рено: Near San Felipe, Cajamarea, 1950 m., May, 1915,
Weberbauer 7109 (FM).

7. Satureia guamaniensis Mansfeld in Bot. Gart. Berlin-Dahlem
Notizbl. 9: 286. 1925.

S. obovata Briq. in Engler u. Prantl, Die Nat. Pflanzenfam., ed.
1, IV. За, 300. 1897, non Lag. Gen. et Sp. Nov. 18. 1816.

Gardoquia obovata Ruiz et Pavon, Syst. Veg. 150. 1798;
Benth. Lab. Gen. et Sp., 403. 1834, et in D.C. Prodr. 12: 236.
1848.

Frutex altitudine circa 1 m., ramis teretibus, ramulis quadratis,
pubescentibus glabratisve, nodis approximatis; foliis 5-7 mm.
longis, obovatis oblanceolatisve, in basi ad petiolum brevissimum
angustatis, utrinque glabris, subtus pallidioribus; margine in-
tegra, revoluta; floribus in axillis solitariis, bracteolis foliis con-
formalibus, quam pedicellis longioribus; calycibus 5-6 mm. longis,
extus glabratis, fauce intus hirsutis, dentibus subaequalibus, lan-
ceolatis, acutiusculis, labia postica tamen longiore, ore itaque ob-
liquo, pedicellis 2 mm. longis elatis; corollis coccineis, 22-24 mm.
longis, tubo extra calycem valde dilato, fauce diametro 8 mm.,
extus pubescentes, intus leniter piloso, labro 5 mm. longo, emar-
ginato, sinu 1 mm. profundo, labiolo aequilongo, lobis aequalibus;
staminibus paulo supra tubi medium sitis, didymis, anticisecorolla
5-6 mm. exsertis, posticis e tubo exsertis; thecis late divaricatis,
.6 mm. longis; stylo e corolla 8 mm. exserto; nuculis non visis.

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 59

S. guamaniensis, as suggested by its author,! is scarcely separ-
able from Gardoquia obovata R. & P.

Specimens examined:

PERU: between Cuancabamba and Oyavaca, 3200 m., May,
1912, Weberbauer 6323 (FM, type collection of S. guamaniensis
Mansf).

8. Satureia breviflora Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia breviflora Benth. Lab. Gen. et Sp. 401. 1834, et in
DC. Prodr. 12: 237. 1848.

Frutex ramulis quadratis, pubescentibus, angulis obtusis, inter-
nodiis saepius folia superantibus; foliis .5-1 cm. longis, rotundatis
vel rotundato-ovatis, obtusis, subsessilibus, pagina superiore
viride, puberula, venis impressis, inferiore cano-tomentosa venis
prominenter costatis, margine subintegra vel frequenter dentato-
crenata, crenis fere 1 mm. altis; floribus 3-6 in axillis, brevissime
pedunculatis, verticillastris saepius ad ramulorum brevium latera-
lium extremitates congestis, foliis floralibus calyces subae-
quantibus, bracteolis linearibus 1-2 mm. longis; calycibus 3.5-4
mm. longis, extus hirtellis, fauce intus nudis, dentibus 1-1.5 mm.
longis, anguste lanceolatis, acutissimis, duobus anticis quam
posticis paulo longioribus, pedicellis gracilibus cirea 1 mm. longis;
corollis 6—7 mm. longis, tubo intus ad staminum anticorum bases
piloso, labro 1 mm. longo, emarginato, labioli lobis subaequalibus,
fere 2 mm. longis, medio paulo majore; staminibus in fauce sitis,
didymis, vix e tubo exsertis, thecis paulo divergentibus; stylo e
tubo paulo exserto; nuculis non visis.

Specimens examined:

Есглров: Quitensian Andes, Jameson (US).

C. Discolores

Foliis saepius 1-2 cm. longis, lanceolatis vel elliptici-lanceolatis,
pagina superiore glabra, sericea vel villosula, inferiore cano-
tomentosa vel tomentella, margine saepius integra (crenata in

1 “Die am nächsten stehende Art S. obovata (Ruiz et Pav.) Briq. unterscheidet sich
nach der (nicht mehr ausreichenden) Beschreibung nur durch längere Blatter,

grau behaarte zweige an der Basis behaarte kelch und innen Kahle Krone.”—Mans«
feld, i.c.

[Vor. 14
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN

S. tomentosa) ; floribus 1-6 in verticillastris sat densis in foliorum
axillis; corollis 1.5-3 cm. longis.

9. Satureia tomentosa Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, ГУ. Abt. За, 300. 1897.

Gardoquia tomentosa Kunth in Humboldt et Bonpland, Nov.
Gen. et Sp. 2: 314. 1817; Benth. in DC. Prodr. 12: 237. 1848.

Frutex ramosus altitudine ad 1 m., ramis duris, ascendentibus,
subteretibus, cortice saepe discedente, internodiis saepius 2-3
em. longis, ramulis gracilibus, ascendentibus, quadratis, angulis
obtusis, superne cano-puberulis vel tomentosis, internodiis
maximam partem quam foliis brevioribus; foliis 1-1.5 em. longis,
saepius ovatis, frequenter rotundatis vel ellipticis, apice acutis vel
obtusis, in basi cuneato-angustatis, margine revoluta, saepius
obscure crenata, pagina superior rugosa, nunc glabra nunc hir-
tella, inferiore albo-tomentosa, rarius villosa, venis subtus prom-
inentioribus, petiolis 1-2 mm. longis elatis; verticillastris
saepius 3-6-floribus, rarius breviter pedunculatis, in axillis
foliorum superiorum dispositis, his calyces paulo superantibus;
calycibus 7 mm. longis, extus molliter appresso-villosis, sub-bi-
labiatis, labia superiore longiore, dentibus supra medium con-
natis, vix .5 mm. longis, triangulo-lanceolatis, inferiore 2 mm.
longa, dentibus liberis, lanceolato-subulatis, approximatis; pedi-
cellis 2 mm. longis elatis; corollis rubris, 2-3 ста. longis, extus
pubescentibus, tubo superne gradatim ampliato, intus tenuiter
piloso, labro subrecto 5 mm. longo, emarginato, labiolo subaequi-
longo, lobo medio lateralibus nune aequilongo nunc fere duplo
longiore; staminibus breviter exsertis, didymis, supra tubo me-
dium sitis, thecis divergentibus; stylo exserto, circa 5 mm. longo;
nuculis atris, oblongis, angustis, 1.5 mm. longis.

A species of some variability in habit and pubescence and to
which S. Kunthii (grandiflora), elegans, and pulchella may ap-
parently be referred. Considering the variability of the species
as shown by the fairly large series of specimens examined, there
is nothing in Kunth’s descriptions to permit of their differenti-
ation.

Specimens examined:

? COLOMBIA: Tobacumdo, June 9, 1876, Andre 3598 (NY); no
data, Andre K 460 (NY).

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 61

EcvApon: Banos, Prov. Tunguragua, 2000 m., Tate 607 (US);
ad sepes, in planitie Rumibamba, Quito, Hartweg 1340 (NY);
ad pontem Guapalo prope Quito Hartweg 1339 (NY); Punin,
Quebrada Chalan, 2779 m., Oct. 28-Nov. 4, 1923, Anthony &
Tate 423 (US); ad vicum Guapalo prope Quito, 2076 m., Jameson
659 (ASP); no data, Jameson (US); Quitensian Andes, 1855,
Couthouy (СН); Riobamba, Aug. 11, 1920, Holway 870 (US; GH);
bei Riobamba, Ambato, Gualabamba, 1800-2700 m., Nov. 25,
1880, Lehmann 149 (US).

Peru: thickets and stream banks, 15 mi. southeast of Huanuco,
3230 m., Мау 31-Лше 3, 1922, Macbride & Featherstone 2084
(forma villosa) (MBG; FM).

10. Satureia thymoides Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. За, 300. 1897.

Gardoquia thymoides Kunth in Humboldt et Bonpland, Nov.
Gen. et Sp. РІ. 2: 314. 1817; et Benth. іп DC. Prodr. 12: 238.
1848.

“Frutex ramosissimus; ramis tetragonis, pubescentibus. Folia
opposita, brevissime potolni ovata, subcordata, acuta, sub-
serrata, margine revoluta, venosa, supra glabriuscula, subtus
cano-pubescentia, tres lineas longa, duas lineas lata. Petioli
pubescentes. Flores verticillati, pedunculati, semipollicares; ver-
ticillis multifloris, distantibus; internodiis quinque aut novem
lineas longis. Calyx tubulosus, decemsulcatus, quinque-den-
tatus, pubescens; dentibus inaequalibus, acuminato-subulatis.
Corolla calyce triplo longior, flava (?) ex Bonpl., pubescens; tubo
interne pubescente; fauce longissima, inferne barbata; limbo bi-
labiato, purpureo-maculato; labio superiore emarginato; inferiore
trifido; laciniis obtusis. Stamina quatuor, didynama, distantia,
subinclusa. Filamenta glabra. Antherae arcuatae. Stylus
exsertus, glaber. Stigma bifidum. Fructus ignotus. Crescit in
Andibus Quitensibus?”

Ut videtur ad Discolores referenda et ab affinibus imprimis
foliis subsessilibus, ovatis, subcordatis, parvis (3 lin. longis)
separanda est.

11. Satureia foliolosa Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. За, 300. 1897.
Gardoquia foliolosa Benth. in DC. Prodr. 12: 238. 1848.

[У от. 14
62 ANNALS OF THE MISSOURI BOTANICAL GARDEN

"folis parvis subsessilibus ovatis obtusis integerrimis supra
pallidis puberulis glabratisve subtus canescenti-tomentellis, ver-
ticillastris 2-6-floris, calyce subsessili hirsutissimo dentibus acutis-
simis patentibus tubo suo vix brevioribus. Ad Chapada de
Santa Martha (Purdie!). | Frutex decumbens, ramosissimus,
dense foliosus. Rami nunc pilis longis hirsutissimis, nunc fere
glabri. Folia 2-3 lin. longa, floralia conformia. Calyces 2 lin.
longi, latiuscule campanulati, incurvi. Corolla villosa, semipol-
licaris? ei G. discoloris similis videtur, sed in specimine nondum
aperta (v.s.)."

Planta mihi ignota similis S. discolori est, tamen ut videtur
foliis ovatis, subsessilibus, minoribus (2-3 lin. longis) et ramis
nune fere glabris, nune densissime pilis longis ornatis imprimis
differt.

12. Satureia discolor Briq. in Engler и. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

АНЯ discolor Kunth in Humboldt et Bonpland, Nov. Gen.
et Sp. Pl. 2: 312. 1817; Benth. in DC. Prodr. 12: 238. 1848.

Frutex ramosissimus, ramis teretibus, duris, cortice discedente,
ramulis quadratis, puberulis, nodis approximatis; foliis densis,
membranaceis, 5-10 mm. longis, maximam partem anguste el-
lipticis, rarius lanceolatis vel oblanceolatis apice acutiusculis, in
basi ad petiolum brevem angustatis, margine integra, revoluta,
pagina superiore subglabra пес nitida, inferiore patenter pal-
lidiore, albo-tomentella, venis prominentioribus; floribus in axillis
solitariis, bracteolis foliosis sed minoribus; calycibus 4.5-5 mm.
longis, extus piloso-villosis, fauce intus subglabris, dentibus tubum
aequantibus, lanceolato-acuminatis, approximatis, tribus posticis
connatis, lanceolatis, acutis, 1.5 mm. longis, omnibus pubescenti-
bus, vix villosis, pedicellis gracilibus, 2 mm. longis, corollis pur-
pureis (Bonpland), 14-15 mm. longis, extus villosulis, tubo extra
calycem dilato, intus ad faucem piloso; labro fere 2 mm. longo,
emarginato, sinu .5 mm. profundo, labiolo aequilongo, lobis sub-
aequalibus; staminibus didymis supra tubi medium sitis, anticis
longioribus, omnibus inclusis, thecis divergentibus, .6 mm. longis;
stylo e corolla vix сеа nuculis non visis.

Specimens examine

VENEZUELA: prope nia Tovar, 1854-5, Fendler 2058

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 63

(MBG; GH); Silla de Caracas, 2600 m., Apr. 27, 1884, Jahn 288
(US); Silla de Caracas, 2000-2640 m., Dec. 26-29, 1918, Pittier
8360 (US); Silla de Caracas, 2461 m., May 21, 1874, Kuntze 1640
(1-3 ft. tall; fl. violet) (US; NY).

13. Satureia acutifolia Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. За, 300. 1897.

Gardoquia acutifolia Benth. in DC. Prodr. 12: 236. 1848.

Frutex ut videtur, ramulis quadratis, sericeo-villosis, ad ex-
tremitates albis, mox fuscis, ramorum internodiis foliorum longi-
tudine subaequilongis, sed foliis ramulorum fertilium (vel ?
juvenilium) fasciculatis; foliis 1.5-3 em. longis, elliptico-lanceo-
latis, utrinque acutis vel in basi angustatis, margine patenter
revoluta, integra, pagina superiore sericeo-villosa, inferiore albo-
tomentosa, petiolis 1-2 mm. longis elatis; floribus tribus in axillis,
bracteolis 2-3 mm. longis subtentis, calycibus 8 mm. longis, extus
sericeis, 13-venis, labiis 2 mm. longis, dentibus duobus anticis
anguste lanceolatis, acuminatis, liberis, tribus posticis supra
medium connatis; pedicellis 1-2 mm. longis elatis; corollis ut
videtur rubris, 2.5 (?-3.0) ст. longis, extus sparse villosis, tubo
superne gradatim ampliato, labro erecto, fere 5 mm. longo, labiolo
subaequilongo; staminibus ut videtur inclusis, stylo paulo ex-
serto; nuculis non еге

садыр examined

Рево: ? Matthews (NY).

14. Satureia mantaroensis Mansfeld in Bot. Gart. Berlin-
Dahlem, Notizbl. 9: 287. 1925

"Frutex usque 2 m. altus. Rami juniores quandrangulares,
dense puberuli. Foliorum lamina 7-14 mm. longa, 5-7 mm. lata,
elliptica vel subobovata vel oblonga, supra brevissime puberula
subtus pallidior, tomentella, plana, nervis utrinque prominulis,
apice acuta vel obtusa, basi angustata, margine integerrima vel
vix conspicue et remote dentata; petiolus cire. 2 mm. longus,
tomentellus. Verticillastri plerumque biflori, foliis floralibus
conformibus. Pedicelli circ. 2 mm. longi, cano-puberuli. Calyx
8 mm. longus extus cano-puberulus, dentibus intus breviter
pilosis, bilabiatus (3/2), dentes labii cire. usque ad medium con-

[Vor. 14
64 ANNALS OF THE MISSOURI BOTANICAL GARDEN

nati, dentes labioli paullo breviores, liberi. Corolla 20-23 mm.
longa (labio emarginato circ. 5 mm. longo, labiolo trilobato 4.5
mm. longo), extus dense pubescens, intus tubo antice piloso et
paulo supra basin annulo pilorum vestita. Stamina didynamia,
anticis longioribus cire. 9 mm. longis, posticis 4-5 mm. longis,
antheris divergentibus, anticis et stylo exsertis. Stylus 32 mm.
longus.

“Peru: Depart. und Prov. Huancavelica, südliche Talwand des
Mantaro über Iscuchaca, Grassteppe mit eingestreuten Stráuch-
ern, 3600 m.u.M., bis 2 m. hoher Strauch mit roten Blüten (Fl.
15. УТ. 1910.—WEBERBAUER n. 5677 !).

“Die Art steht nach der Linge der Krone zwischen S. Pavoni-
ana Briq. (Gardoquia incana Ruiz et Pav.) und S. discolor (Kunth)
Briq. in der Mitte; S. discolor hat ferner eingeschlossene Stamina
und inne kahle Kronróhre, S. Pavoniana oberseits bleiche Blatter
(nach der nicht mehr ausreichenden Beschreibung). 8. thymoides
(Kunth) Briq. weieht durch den fast herzfórmigen Blattgrund
und zurückgerollten deutlich gezihnten Blattrand ab, steht aber
sonst besonders im Blütenbau der S. mantaroénsis wohl am
nächsten.”

Ex descriptione ab S. Pavoniana vix distincta videtur.

15. Satureia Pavoniana Briq. in Ann. Conserv. et Jard. Bot.
Genève 2: 189. 1

Gardoquia incana Ruiz et Pavon, Syst. Veg. 150. 1798; Benth.
Lab. Gen. et Sp., 401. 1834, et in DC. Prodr. 12: 237. 1848.

Frutex altitudine 30 ст.-2 m., caulibus erectis vel ascendenti-
bus, lignosis, subteretibus, cortice discedente, internodiis saepius
3-4 cm. longis, ramulis saepius 5-10 cm. longis, gracilibus, canes-
centibus, ascendentibus, internodiis quam foliis saepius breviori-
bus; foliis 1-1.5 cm. longis, ellipticis, rarius ovatis, acutis, in basi
angustatis, margine maximam partem integra planaque, rarius
denticulata vel subrevoluta, pagina superiore pallidiore, non
rugosa sed venis tamen saepe prominentioribus, inferiore dense
cano-tomentella, petiolis 1-2 mm. longis, verticillastris 1-3-
floribus, decussatim instructis in foliorum superiorum axillis dis-
positis, his calyces paulo superantibus, bracteolis linearibus or-
natis; calycibus incanis, paulo arcuatis, 6-8 mm. longis, subbi-

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 65

labiatis, fauce intus subnudis, labiis circa 1.5 mm. longis, dentibus
lanceolato-subulatis, posticis tribus ad medium connatis, anticis
fere liberis brevioribus, pedicellis 2-3 mm. longis elatis; corollis
rubris, fere 3 сш. longis, arcuatis, extus pubescentibus, tubo
superne gradatim ampliato intus infra staminum praecipue an-
ticorum bases piloso, labro subrecto, 5 mm. longo, emarginato,
labiolo subaequilongo, lobo medio longiore; staminibus didymis,
omnibus breviter exsertis supra tubi medium sitis; stylo cirea 5
mm. exserto; nuculis cirea 1.5 mm. longis, obidni angustis,
atris.

Specimens examined:

Peru: Yanahuanca, densely shrubby northeastern slope,
3076 m., June 16-22, 1922, Macbride & Featherstone 1196 (FM;
MBG); sunny blackberry patch, Mito, 2769 m., July 8-22, 1922,
Macbride & Featherstone 1411 (FM; MBG); dry hills, Oroya,
Lima, 3076-3384 m., 1919, Kalenborn 82 (US; MBG); no data,
Dombey (GH).

D. Striatae

Foliis 2-15 mm. longis, forma diversis, utrinque glabris, saepius
subnitidis, venis prominulis, ascendentibus, rectis, parallelis,
margine subintegra, floribus solitariis vel tribus in foliorum axillis;
corollis 1.5-3 cm. longis.

16. Satureia pallida sp. nov.

Frutex ramosus, altitudine ut videtur cirea 1 m., ramis as-
cendentibus, teretibus, lignosis, cortice discedente, Panini di-
varicatis, glabris, quadratis, angulis acutis, internodiis foliis sub-
aequilongis vel brevioribus; folis 1-1,5 cm. longis, maximam
partem ovalibus vel роо омон obtusis, іп basi saepius
rotundatis, utrinque glabris et pallidioribus, margine leniter
revoluta, integra, petiolis 1-2 mm. longis elatis; floribus 1-8 in
foliorum supremorum axillis dispositis; calycibus in speciminibus
visis 8-9 mm. longis, puberulis, subbilabiatis, fauce intus nudis,
labiis fere 1.5 mm. longis, dentibus posticis tribus ad medium
connatis, anticis liberis, omnibus subulatis, acutiusculis, pedicellis
3—4 mm. longis, bracteolis linearibus ad medium ornatis elatis;
corollis 2.5 cm. vel ultra longis, arcuatis, extus pubescentibus,
tubo superne gradatim dilato, intus infra staminum praecipue

[Vor. 14
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN

anticorum bases piloso, labro subrecto, 4 mm. longo, emarginato,
labiolo subaequilongo, lobo medio longiore; staminibus didymis,
omnibus breviter exsertis, supra tubi medium sitis; stylo exserto;
nuculis circa 1.5 mm. longis, ellipticis, atris.

A species allied to 5. Pavoniana and at first thought to be а
glabrous variety of it. However, study of other members of the
type collection, together with an unnumbered collection by Bang
at the New York Botanical Garden which is apparently conspeci-
fic, suggests a range of leaf variation too great to permit of re-
ference to S. Pavoniana. Тһе flowers of the two species are very
similar, differing chiefly in the size of the calyces and in the more
acute, even acuminate, calyx teeth of 8. pallida. The species
is intermediate with the groups Discolores and Striatae.

Specimens examine

Bouivia: Bang, no date (NY); Turedon, Bolivian plateau,
1891, Bang 1127 (US; NY, түре; GH).

17. Satureia glabrata Briq. in Engler и. Prantl, Die Nat.
Pflanzenfam., ed. 1, ТУ. Abt. За, 300. 1897.

Gardoquia glabrata Kunth in Humboldt et Bonpland, Nov. Gen.
et Sp. Pl. 2: 313. 1817; Benth. in DC. Prodr. 12: 236. 1848.

Frutex ramosissimus, ramis teretibus, cortice discedente,
ramulis densis quadratis patenter pubescentibus; foliis confertis,
6-9 mm. longis, ellipticis vel elliptico-lanceolatis utrinque acuti-
usculis, subsessilibus, utrinque glabris, pagina superiore nitida,
inferiore pallidiore, punctata, venis prominentibus, margine
subintegra, vix revoluta; floribus tribus in axillis, rarius solitariis
pedunculis subnullis, bracteolis primis omnino foliosis, secundariis
conformalibus, minoribus ornatis; calycibus 7 mm. longis, extus
glabris, fauce intus leniter hirsutis, dentibus subaequalibus, fere
3 mm. longis, longe acuminatis, ore obliquo, pedicellis 2-2.5 mm.
longis; corollis 27 mm. longis, extus pubescentibus, intus glabris,
labro 7 mm. longo, emarginato, sinu 1 mm. profundo, labiolo
paulo breviore, lobo medio lateralibus subduplo longiore; sta-
minibus didymis, supra tubi medium sitis, posticis labro paulo
2... anticis corollam superantibus; thecis late divergenti-
bus, .7-.8 mm. longis; stylo e corolla 7-8 mm. exsertis; nuculis
non visis.

1927]
EPLING— SOUTH AMERICAN LABIATAE. ІП 67

Specimens examined:
Есодров: vic. of Tablón de Oña, Sept. 27, 1918, Rose 23083
(US).

18. Satureia taxifolia Briq. in Engler и. Prantl, Nat. Pflanzen-
fam., ed. 1, ТУ. Abt. 3a, 300. 1897.

Gardoquia taxifolia Kunth in Humboldt et Bonpland, Nov.
Gen. et Sp. Pl. 2: 312. 1817; Benth. in DC. Prodr. 12: 236.
1848.

Thymus taxifolius Willd. ex Benth. in Linnaea 11: 342. 1837.

Frutex ramosissimus altitudine 1-2 m., ramis teretibus, cortice
discedente, ramulis densis, quadratis, puberulis; foliis densis, ad
ramorum extremitates saepius imbricatis, maximam partem 10-
12 mm. longis, saepius lanceolato-oblongis, frequente oblanceo-
latis etiam lineari-oblongis, apice obtusis, rarius acutiusculis, in
basi ad petiolum brevissimum angustatis, utrinque glabris, pagina
superiore nitida, inferiore pallidiore, venis prominentibus, margine
revoluta, integra vel praecipue ad apicem obscure serrata; floribus
in axillis solitariis, rarius didymis, bracteolis linearibus quam
pedicellis duplo longioribus ornatis; calycibus 6-7 mm. longis,
cano-puberulis, fauce intus nudis, dentibus subaequalibus, circa
1 mm. longis, lanceolatis, acutiusculis, labia postica longiore, ore
itaque obliquo, pedicellis 1 mm. longis; corollis coccineis, 25-27
mm. longis, extus pubescentibus, tubo extra calycem multo
dilato fauce intus sparse piloso, labro 3-3.5 mm. longo, emargin-
ato, sinu .5 mm. profundo, labiolo aequilongo, lobis subaequal-
ibus; staminibus supra tubi medium sitis, didymis, anticis corolla
longioribus, posticis e tubo exsertis; thecis late divaricatis, .6
mm. longis; stylo e corolla 4-5 mm. exserto; nuculis non visis.

Specimens examined:

Ecvapon: Loja, between San Lucas and Oña, 2200-3100 m.,
Sept. 7, 1923, Hitchcock 21527 (US); Loja, between La 'Toma and
Loja, 1800-2600 m., Sept. 4, 1923, Hitchcock 21437 (US); in
montibus, Loja, Hartweg 808 (cited by Bentham as 888) (NY);
vicinity of Loja, Sept. 29-Oct. 3, 1918, Rose 23279 (US).

19. Satureia Andrei sp. nov.
Frutex habitu foliisque fere 5. glabratae, foliis ramulisque
tamen utrinque tomentellis, eis 10-12 mm. longis, oblanceolatis,

[Vor. 14
68 ANNALS OF THE MISSOURI BOTANICAL GARDEN

obtusis, in basi angustatis, venis prominulis, margine subserrata,
subrevoluta, petiolis subnullis; floribus in axillis solitariis, brac-
teolis deciduis ornatis; calycibus 5.5 mm. longis, extus puberulis,
13-venis, dentibus 1 mm. longis, lanceolatis, acutis, subaequalibus
sed tamen leniter bilabiatis, intus hirsutis; corollis extus villosis
ut videtur circa 15 mm. longis, antheris generis vix exsertis, stylo
exserto 4—5 mm.; nuculis non visis.

Specimens examined:

Corounia: Cisne, Oct. 30, 1876, Andre 4331 (NY, түре, haud
satis est).

20. Satureia striata Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia striata Ruiz et Pavon, Syst. Veg. 148. 1798; Benth.
Lab. Gen. et Sp. 404. 1834; et in DC. Prodr. 12: 238. 1848.

Frutex strictus, altitudine .5-2 m., eaule in basi crasso, duro,
terete, striato, ramis ramulisque gracilibus, confertis, fastigiatis,
subquadratis, pubescentibus; foliis densis, subconduplicatis, 2-5
mm. longis, plerumque ovatis, obtusis, in basi rotundato-an-
gustatis, vix cordatis, glabris vel vena media subtus ciliata, pagine
superiore nitida, venis impressis, inferiore punctata, venis pulchre
elevato-striatis lateralibus cirea 4-6, pedicellis subnullis, pubes-
centibus; floribus in axillis solitariis, bracteolis foliis conformali-
bus; calycibus 6-7 mm. longis, profunde suleatis, 13-venis, gla-
bris, fauce intus nudis, dentibus 1 mm. longis, subaequalibus,
lanceolatis, acutis, patentibus, subrecurvis, pedicellis gracilibus
2 mm. longis; corollis rubris, 20-25 mm. longis, extus molliter
villosis, tubo extra calycem multo ampliato, intus infra labiolum
pilosis, labro erecto, 5-6 mm. longo, emarginato, sinu 1 mm.
profundo, labiolo 4 mm. longo, lobis subaequalibus, rotundatis,
medio patentibus; staminibus didymis, posticis e tubo paulo ex-
sertis, anticislabrum subaequantibus, omnibus supra tubi medium
sitis; thecis divergentibus, .5—.6 mm. longis; stylo e corolla 3 mm.
exserto; nuculis non visis.

Specimens examined:

Рено: Mito, July 23-Aug. 14, 1922, Macbride & Featherstone
1734 (FM; MBG); 12 mi. ont of Рапао, 3,076 m., July 4-10,
1922, Macbride & Featherstone 2213 (“а common bilide shrub”)
(FM; MBG).

у a4 PEE E

1927]
EPLING—SOUTH AMERICAN LABIATAE. III 69

21. Satureia plicatula sp. nov.

Suffrutex procumbens, caule duro, ramisque teretibus, ramulis
subquadratis, puberulis, nodis approximatis, saepe in ramulis
lateralibus brevibus confertissimis; foliis 3-5 mm. longis, ovatis,
apice acutis, subacuminato-mucronulatis, in basi rotundatis, sub-
sessilibus, utrinque subglaucis et minutissime puberulis, venis
paginae superioris leniter impressis, inferioris pulchre elevatis,
lateralibus parallelis plicatulis, media prominentiore, margine
serratula пес revoluta; floribus in axillis solitariis, bracteolis
pedicellis aequilongis, lineari-carinatis, mox deciduis ornatis,
calycibus 6 mm. longis, extus pubescentibus, fauce intus hirsutis,
dentibus anticis 1.3 mm. longis, subulatis, patentibus, tribus
posticis connatis, subulatis, subpatentibus, pedicellis 2-2.5 mm.
longis; corollis coccineis 25 mm. longis, tubo extra calycem multo
dilato, extus pubescente, intus praecipue ad staminum bases
piloso, labro 4.5 mm. longo, emarginato, sinu .5 mm. profundo,
labiolo paulo longiore, lobo medio lateralibus duplo longiore,
rotundato, in hoc specimine emarginato, in basi angustato;
staminibus paulo supra tubi medium sitis, didymis, omnibus e
tubo exsertis, anticis corolla subaequilongis; thecis divergentibus,
.7 mm. longis, stylo e corolla 5 mm. exsertis; nuculis non visis.

Specimens examined:

PERU: pendant from river cliff ledges, 2153 m., Шаба, Aug. 21,
1922, Macbride & Featherstone 2238A (FM, түре).

22. Satureia Jamesoni Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia Jamesoni Benth. Lab. Gen. et. Sp. 404. 1834, et in
DC. Prodr. 12: 239. 8

Frutex ut videtur strictus ramosissimus altitudine .5-1 m.,
caule in basi crasso, duro, terete, striato, ramis ramulisque graci-
libus, confertis, fastigiatis, subquadratis, pubescentibus; foliis
parvis, confertis, 2 mm. longis, plerumque ovatis, obtusis, in basi
rotundato-angustatis, utrinque tenuissime puberulis, subcon-
duplicatis, margine revoluta integra, venis subtus prominentiori-
bus, petiolis minimis, floribus solitariis vel tribus in axillis, brac-
teolis foliis conformalibus minutissimis; calycibus 5-7 mm. longis,
profunde sulcatis, 13-venis, subglabela. fauce intus nudis, den-

[Vor. 14
70 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tibus 1 mm. longis, subaequalibus, lanceolatis, acutis, patentibus,
subrecurvis, pedicellis gracilibus 2 mm. longis; corollis rubris,
20-25 mm. longis, extus pubescentibus, tubo extra calycem multo
ampliato, intus infra labiolum piloso, labro erecto, 3 mm. longo,
emarginato, sinu 1 mm. profundo, labiolo breviore, lobis sub-
aequalibus, rotundatis, medio patente; staminibus didymis,
posticis vix e tubo exsertis, anticis labiolo paulo longioribus, om-
nibus supra tubi medium sitis, thecis divergentibus, .5-.6 mm.
longis, stylo e corolla рема exserto; nuculis non visis.

Specimens examine

COLOMBIA: Ayapel, Audi K 1564 (NY) (doubtful station).

EcuApon: Provinces Azuay and Cañar, Paramo between
Cuenca and Huigra, 2700-3000 m., Sept. 12-13, 1923, Hitchcock
21693 (US); no data, Lehmann 4676 (US); vicinity of Cañar,
Sept. 15, 1918, Rose 22720 (US).

23. Satureia connata nom. nov.

S. microphylla Briq. in Engler u. Prantl, Nat. Pflanzenfam., ed.
1, IV. Abt. За, 300. 1897; non Guss. Fl. Sic. Prodr. 2: 120. 1828.

Gardoquia microphylla Kunth in Humboldt et Bonpland, Nov.
Gen. et Sp. Pl. 2: 311. 1817; Benth. Lab. Gen. et Sp. 404.
1834, et in DC. Prodr. 12: 238. 1848.

? Satureia ericoides Willd. ex Benth. in Linnaea 11:328. 1837.

Frutex ut videtur altitudine cirea 1 m., ramis ascendentibus,
teretibus, cortice discedente, ramulis quadratis pubescentibus,
nodis approximatis; foliis 2-3 mm. longis, subconduplicatis,
ovatis, apice obtusis, in basi cordatis, subsessilibus, margine
integra, subrevoluta, pagina superiore subglabra, nitida, in-
feriore puberula, venis prominentioribus; floribus in axillis soli-
tariis, bracteolis omnino foliosis, pedicellos breviter superantibus;
calycibus 5 mm. longis, extus puberulis fauce intus glabris,
dentibus anticis ovatis, acutis, 1 mm. longis, erectis, tribusposticis
connatis, subnullis, labia itaque subintegra, pedicellis 1-1.5 mm.
longis; corollis соссіпеів, 15-20 mm. longis, tubo extra calycem
multo dilato, extus pubescente, intus infra labiolum piloso, lobo
2.5 mm. longo, emarginato, sinu 1 mm. profundo, labiolo paulo
breviore, lobis subaequalibus; staminibus in hoc specimine sub-
aequilongis, paulo supra tubi medium sitis, 1.5 mm. longis, in

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 71

tubo omnino inclusis, thecis parallelis, .5 mm. longis; stylo e
corolla 1-2 mm. exserto; nuculis non visis.

Specimens examine

Есодров: ranch odes Ibarra nud Tulcán, Carchi, 3000 m.,
Aug. 10-11, 1923, Hitchcock 20793 (near type locality) (US; NY).

E. Revolutae

Foliis 2-6 mm. longis, ovatis vel linearibus, pagina superiore
glabra, sericea vel pubescentia, inferiore tomentosa vel puberula,
margine patenter saepius valde revoluta, integra; floribus soli-
tariis in axillis (5-9 in verticillastris sat densis in 8. rigidula);
corollis 9-25 mm. longis.

This group represents the extremes of adaptation to an arid
habitat to be found in the genus. Тһе conformation of the leaves
of S. Lindeniana is especially noteworthy.

24. Satureia argentea Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. За, 300. 1897.

Gardoquia argentea Kunth in Humboldt et Bonpland, Nov.
Gen. et Sp. Pl. 2: 313. 1817; Benth. in DC. Prodr. 12: 237.
1848.

Suffrutex procumbens, ramosus, ramis teretibus, cortice dis-
cedente, ramulis pubescentibus, quadratis, nodis approximatis;
foliis 3-4 mm. longis, lanceolatis, acutiusculis, in basi rotundato-
angustatis, sessilibus, margine patenter revoluta, integra, vena
media subtus prominentiore, utrinque argenteo-canis; floribus in
axillis solitariis, bracteolis omnino foliosis; calycibus 4.5-5 mm.
longis, extus argenteo-canis, fauce intus beds dentibus anticis
1 mm. longis, ovatis, acutis, tribus posticis connatis, vix .5 mm.
longis, acutis, pedicellis 2 mm. longis elatis, corollis coccineis,
14 mm. longis, tubo extra calycem valde dilato, extus villoso,
intus infra labiolum piloso, labro erecto, 2.5 mm. longo, emar-
ginato, sinu .5 mm. profundo, labiolo quam labro paulo breviore,
lobis subaequalibus; staminibus didymis, posticis 2-3 mm.
longis, vix e tubo exsertis, anticis 2-4 mm. longis, plerumque e
corolla breviter exsertis, thecis paulo divergentibus, .6—.7 mm.
longis; stylo e corolla 3-4 mm. exserto; nuculis non visis.

Specimens examined:

Реко: pendant from river cliff ledges, 2153 m., Llata, Aug. 21,
1922, Macbride & Featherstone 2238 (MBG,).

[Vor. 14
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN

25. Satureia Weberbaueri Mansfeld in Bot. Gart. Berlin-Dahlem,
Notizbl. 9: 285. 1925.

“Frutex 0.5 m. altus. Rami juniores quadrangulares, breviter
puberuli. Foliorum lamina 6-11 mm. longa, 2-4 mm. lata,
lamina majorum oblongo-elliptica v. lanceolato-elliptica, lamina
minorum saepe linearis v. lanceolato-linearis, apice obtusiuscula,
basi in petiolum angustata, margine revoluta (praecipue in foliis
minoribus), supra et subtus, cano-brunneo-puberula, nervis supra
immersis, subtus prominulis; petiolus 1 mm. longus vel subnullus,
puberulus. Verticillastri apice ramulorum brevium spicas for-
mantes, 2-6-Ногі; folia floralia conformia, calyces aequantia vel
superantia; bracteae linares. Calyx 8 mm. longus, cano-brun-
neo-puberulus, costatus, bilabiatus (3/2), dentibus labii paullo
connatis, dentibus labioli subliberis, intus puberulis. Corolla
circ. 15 mm. longa, labio emarginato et labiolo trilobato circ. 3
mm. longis, extus + pubescens, intus antice disperse pilosa.
Stamina didynamia, filamenta antica 6 mm. longa, exserta,
postica 4 mm. longa, subexserta; antherae divergentes. Stylus
20 mm. longus.

“Peru: Depart. Libertad, Prov. Santiago de Chuco, bei der
Hacienda Angasmarea, lockeres, von Grassteppe durchsetztes
Gestrauch, 3000-3100 m., 16 m. hoher Strauch mit purpurnen
Blüten (Fl. 8. VII. 1914.—WEBERBAUER n. 7016!).

“Die nachststehenden Arten S. argentea (Kunth) Briq. und
S. sericea (Presl) Briq. unterscheiden sich schon durch die dich-
tere, weisze Behaarung."

Mihi omnino ignota.

26. Satureia rigidula nom. nov.

S. fasciculata Briq. in Engler u. Prantl, Nat. Pflanzenfam., ed.
1, IV. Abt. За, 300. 1897, nec Rafin. Préc. Découv. 39. 1814;
пес Tausch in Syll. Ratisb. 2: 248. 1828.

Gardoquia fasciculata Benth. Pl. Hartweg. 243. 1839; et in
DC. Prodr. 12: 239. 1848.

Frutex altitudine 40—80 cm., ramis erectis vel ascendentibus,
virgatis, ramulis numerosis, brevibus, ascendentibus, omnibus
subteretibus, pubescentibus, saepe incanis, ramulorum inter-
nodiis foliis subaequilongis; foliis in axillis fasciculatis, sessilibus.

А, tA УЛ Ж ААН

1927]
EPLING— SOUTH AMERICAN LABIATAE. ПТ 73

3-5 mm. longis, saepius oblongo-linearibus et in basi .5-1.5 mm.
latis, frequenter tamen anguste triangulis et in basi latioribus,
acutis, fere ad venam mediam revoluta, pagina superiore puberula,
inferiore cano-tomentosa; verticillastris saepius 5-9-floribus, de-
eussatim instructis, in spicam densam 1-4 ст. longam ad ramu-
lorum apices confertis, bracteis subulatis, calycibus brevioribus;
ealycibus 3.5-4 mm. longis, extus puberulis, subbilabiatis, fauce
intus nudis, dentibus lanceolato-subulatis, superioribus circa 1.5
mm. longis, inferioribus fere 2 mm. longis, pedicellis subnullis;
corollis violaceis, cirea 10 mm. longis, extus pubescentibus, tubo
superne ampliato, decurvo, intus infra stamines pubescentibus,
labro bifido, sinu cirea .5 mm. profundo, labiolo trifido, lobo
medio paulo majore, staminibus didymis, ad corollae medium
sitis, anticis subexsertis, posticis inclusis, thecis divergentibus;
stylo paulo exserto; nuculis oblongis, angustis, circa 1 mm. longis,
fuscis.

Specimens examined:

?СогомвтА: Rio Chota, June 6, 1876, Andre 3583 (NY).

EcvApom: plains of Pamasqui and San Antonio, 2615 m.,
Jameson (NY); in planitie Rumibamba necnon juxta pontem
Guapalo prope Quito, Hartweg 1338 (NY); по data, Lehmann 6847
(US); Alausf, Chimborazo, 2500 m., July 19, 27, 1923, Hitchcock
20719 (US); crescit in apricis prope Quito, 2461 m., Jan. 21, 1856,
Jameson 181 (GH); 1857-9, Spruce 6062 (GH); Alausí, Chim-
borazo, 2500 m., July 19, 27, 1923, Hitchcock 20705 (US).

27. Satureia sericea Briq. in Engler и. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia sericea Presl in Benth. Lab. Gen. et Sp. 402. 1834;
Benth. in DC. Prodr. 12: 238. 1848.

Suffrutex procumbens vel erectus, altitudine circa 1 m., ramis
ascendentibus, teretibus, cortice discedente, ramulis quadratis,
sericeo-villosis, gracilibus; foliis 5-7 mm. longis, oblongo-lineari-
bus, 1-2 mm. latis, obtusis, in basi ad petiolum brevissimum
angustatis, margine integra, valde revoluta, utrinque sericeo-
villosis, argenteis, subtus albis, saepe in axillis fasciculatis; floribus
tribus in axillis, rarius solitariis, brevissime pedunculatis, brac-
teolis primis omnino foliosis, secondariis conformalibus sed

[Vor. 14
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

minoribus; calycibus 6-7 mm. longis, extus sericeo-villosis,
dentibus intus supra faucem pubescentibus duobus anticis 1.5
mm. longis, posticis 1 mm., omnibus lanceolatis, acutis, pedicellis
1-2 mm. longis elatis; corollis coccineis extus villosis, 22-24 mm.
longis, tubo extra calycem valde dilato, intus infra labiolum
piloso, labro erecto, 3 mm. longo, emarginato, sinu 1 mm. pro-
fundo, labiolo paulo breviore, lobis subaequalibus, medio tamen
paulo longiore; staminibus didymis, supra tubi medium sitis,
anticis 7 mm. longis, e corolla exsertis, posticis 4 mm. longis,
tubo subaequalibus, antheris .7-.8 mm. longis, thecis divergenti-
bus; stylo 6-8 mm. e corolla exserto; nuculis non visis.

Specimens examined:

Рен: sunny slopes, **Chunmis," Chasqui, 3230 m., Sept. 27,
1922, Macbride & Featherstone 1765 (MBG; FM); grassy rocky
canyon slope, 2153 m., Llata, Aug. 21, 1922, Macbride & Feather-
stone 2243 (MBG; FM); hacienda, 9 mi. up river from Yanahuan-
ca, 3282 m., northwest grassy slope, June 21, 1922, Macbride &
Featherstone 1272 (MBG; ЕМ); no data, Matthews (NY).

28. Satureia revoluta Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Gardoquia revoluta Ruiz et Pavon, Syst. Veg. 149. 1798;
Benth. Lab. Gen. et Sp., 405. 1834; et in DC. Prodr. 12: 239.
1848.

S. insignis Mansfeld Bot. Gart. Berlin-Dahlem, Notizbl. 9: 288.
1925.

Suffrutex procumbens, caulibus teretibus, cortice discedente,
ramis ramulisque pubescentibus, subteretibus quadratisve, gracili-
bus, confertis et subfastigiatis; foliis densis, 2-3 mm. longis,
ovatis vel patenter triangulis, apice obtusis, in basi truncatis,
margine rarius convexa, valde et pulchre revoluta, pagina superiora
viride, puberula, inferiore dense albo-tomentosa, petiolis 1 mm.
longis elatis; floribus saepius solitariis in axillis, bracteolis omnino
foliosis; мин 4,5-5 mm. longis, extus cano-puberulis, fauce
intus nudis, patenter bilabiatis, labiis 1.5 mm. longis, dentibus
parte dimidia breviore, lanceolatis, acutis, pedicellis maturis 3—4
mm. longis; corollis coccineis 20-22 mm. longis, tubo extra caly-
cem multo dilato, extus molliter villoso intus infra labiolum pi-

АС Es}

1927]
EPLING—SOUTH AMERICAN LABIATAE. III 75

losis, labro егесфо 3 mm. longo, emarginato, sinu 1 mm. profundo,
labiolo aequilongo vel paulo longiore, lobis subaequalibus; sta-
minibus didymis supra tubi medium sitis, omnibus e tubo ex-
sertis, anticis labro subaequilongis thecis subparallelis, .5 mm.
longis; stylo e tubo 5-6 mm. exserto; nuculis поп visis.

The present author has found no characters in the type col-
lection of S. insignis as represented in the Field Museum to
justify specific Scit from S. revoluta as understood by him.

Specimens examined

Peru: Wilkes Exp. (08; СН); between Cuaneabamba and
Oyavaca, 3200 m., May 1912, Weberbauer 6333, (FM, type collec-
tion of S. SAPE Mansf.); Oroya near Lima, 1919, M. Kalen-
born 162 (US; MBG); ? Culluy, July, Matthews 666 (GH).

29. Satureia Lindeniana Briq. in Ann. Conserv. et Jard. Bot.
Genéve 2: 191. 1898.

Suffrutex ut videtur, altitudine cirea 15 cm., ramis e caudice
lignoso erectis, virgatis, subteretibus, pubescentibus, nodis inter
se 2-5 mm. distantibus, ramulis brevibus, erecto-ascendentibus;
foliis cirea 4 mm. longis, erectis, sessilibus, oblongo-linearibus,
apice obtusis, in basi rotundato-angustatis, marginibus integris,
valde replicatis et infra venam mediam conniventibus, paginam
inferiorem cano-tomentosam itaque toto adumbrantibus, facie
superiore pubescente, inferiore glabra; floribus oppositis, soli-
tariis in axillis, calycibus subtubulosis, 6 mm. longis, extus pu-
berulis, paulo arcuatis, patente bilabiatis, labia superiore 2 mm.
longa, dentibus ad medium connatis, breviter triangulari-lance-
olatis, inferiore 3 mm. longa, dentibus lanceolato-subulatis inter
se liberis et а labia superiore sinu distinctiore separatis, pedicellis
1 mm. longis elatis; corollis 9 mm. longis, extus pubescentibus,
tubo intus nudo, superne ampliato, leviter arcuato, labro sub-
erecto, 1 mm. longo, ovato, emarginato, labiolo 2 mm. longo,
patente, lobo medio majore obovato; staminibus sub labro as-
cendentibus; stylo pauio exserto; nudus non visis.

Specimens examine

COLOMBIA: Nevada Sta. Marta, Purdie (GH).

30. Satureia rugosa Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, ТУ. Abt. За, 300. 1897.

[Vor. 14
76 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

Gardoquia rugosa Benth. Lab. Gen. et. Sp., 399. 1834; et in
DC. Prodr. 12: 236. 1848.

“fruticosa, villosa, foliis petiolatis ovato-rhomboideis serrato-
dentatis basi rotundatis coriaceis rugosis utrinque villosis subtus
vix canescentibus, verticillastris laxis, multifloris, calycis villosi
dentibus subulatis ciliatis, fauce intus subnuda, corollis calyce vix
duplo longioribus. Іп Peruvia (Ruiz et Pavon!). Frutex ra-
mosissimus, ramis duris tetragonis junioribus rufo-villosis. Folia
1-11% pollicaria, reticulato-venosa, floralia minora et cymas
superantia. Cymae breviter pedunculatae. Bracteae oblongae,
villosae, calyce breviores. Pedicelli breves. Calyces 3 lin. longi,
tenues, virides, villosissimi. Corolla villosa, labio superiore
erecto, brevissime emarginato, inferiore subpatente, lobis oblongis
inter se subaequalibus obtusis integerrimis. Stamina didynama,
antheris sub labio superiore per paria approximatis. Species
aequo jure ad Melissam, Micromeriam, vel Gardoquiam referenda
(v.s. olim in herb. Lamb.)." (Benth. in DC. Prodr. 12: 236.
1848).

Sect. Xenopoma (Willd.) Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. За, 300. 1897.

Xenopoma Willd. in Ges. Naturforsch. Fr. Berlin Mag. 5: 399.
1811.

Suffrutices fruticulive, rarius herbae perennes humiles, ramulis
divaricatis, foliis obovatis, subintegris, pubescentibus; floribus
1-6 in foliorum axillis, praecipue in speciebus humilibus solitariis,
pedicellis quam calycibus brevioribus, singulis bracteolis duobus
ornatis elatis; calycibus tubulosis, frequenter subturbinatis, den-
tibus subaequalibus obscure bilabiatis acutis, intus nudis vel hir-
sutis, erectis vel patentibus; corollae tubo omnino incluso vel
breviter exserto.

31. Satureia nubigena Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. За, 300. 1897.

Thymus nubigenus Kunth in Humboldt et Bonpland, Nov. Gen.
et Sp. Pl. 2: 316. 1817.

Micromeria nubigena Benth. Lab. Gen. et Sp. 381. 1834; et
in DC. Prodr. 12: 222. 1848.

Thymus humifusus Willd. ex Benth. in Linnaea 11: 342. 1837.

1927]
EPLING— SOUTH AMERICAN LABIATAE. III 77

Herba procumbens, fragrans, caulibus reptantibus, ramosis-
simis, ramis ramulisque filiformis, atris, puberulis hispidisve,
subteretibus, foliis 3-4 mm. longis, approximatis, plerumque
ovatis, apice obtusis, in basi ad petiolum angustatis, frequenter
rotundato-ovatis et in basi sub-truncatis, utrinque puberulis
saepius villosulis vel hispidulis rarius subglabris, margine integra
revoluta; floribus in axillis solitariis, bracteolis parvis, linearibus,
parte tertia pedicellorum longis ornatis; calycibus 3-4 mm. longis,
superne patenter dilatis, hirsutulis, sub-15-venis, fauce subnudis,
dentibus .6-.9 mm. longis, lanceolato-acuminatis, aequilongis,
anticis tamen patenter majoribus quam posticis tribus, pedicellis
1.5-2 mm. longis elatis; corollis 5-6 mm. longis, tubo superne sat
ampliato, nectarostegio e pilis intus fauce areolam formante,
labro emarginato, sinu parte dimidia labri longitudinis profundo,
labioli lobo medio majore, rotundato, 1-1.5 mm. longo; stamini-
bus parvis, circa 1 mm. longis, didymis, supra tubi medium sitis,
thecis paulo divergentibus; stylo corolla aequilongo; nuculis .8-
.9 mm. longis, oblongis, atris.

Specimens examined:

VENEZUELA: ‘‘Poleo de paramo," Paramo Sto. Domingo, 3600
m., Mérida, Sept. 14, 1922, Jahn 1151 (US); Laguna Verde,
Paramo Mucuchies, Mérida, 3384 m., 1922, de Bellard 19 (US);
summit of Páramo, Quirorá, 3200 m., Mérida, Feb. 24, 1922,
Jahn 882 (US); Sierra de Nevada de Mérida, Laguna del Gallo,
4070 m., Dec. 1910, Jahn 74 (US).

COLOMBIA: in monte ignivomo Azufral, May 18, 1876, Andre
3249 (NY); Hacienda de Antisana, Hartweg 1337 (NY); Ruiz,
3000 m., 1918, Dawe 750 (NY); Canaan, Mt. Puracé, El Cauca,
open near Rio Anambiu, 2900-3200 m., June 11-16, 1922, Killip
6732 (ASP; US); dry open, 3500-4000 m., Paramo de Ruiz, To-
lima, Dec. 16-17, 1917, Pennell 3015 (US; МВС; СН; NY);
Westabhange des Paramó de Ruiz, El Cauca, 3000-3500 m., Sept.
11, 1883, Lehmann 3111 (US).

Wovanon: Andes near Quito, 4000 m. » Couthouy (NY); Andes
of Quito, 4000 m., Jameson 217, ^ ora in graminosis alpinis
(vernacule n de los Andes"), 4000 m., Jan. 21, 1856 (US;
GH); Paramo de Tuza, 3400 m., Jan. 31, 1881, Т іскен 3092 а
(US); between Urbina and Mt. Chimborazo, 3600-4500 m., Prov.

[Vor. 14
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Chimborazo, Oct. 4, 1923, Hitchcock 21987 (US; NY); La Rin-
conada between Ibarra and Tulcan, 3000 m., Carchi, Aug. 10-11,
1923, Hitchcock 20787 (US; NY); paramo between Ofia and
Cuenea, 2700-3300 m., Azuay, Sept. 9-10, 1923, Hitchcock 21685
(U$).

Peru: mountain of ? Pellshum, “very plentiful,’ Jameson
(ASP); on mossy rock, Tambo de Vaca, 4000 m., June 10-24,
1923, Macbride 4399 (FM ; MBG).

Var. glabrescens Benth in DC. Prodr. 12: 222. 1848.

Ramis foliisque puberulis vel subglabris nec villosulis hispidu-
lisve.

Specimens examined:

CoLoMBIA: dry grassy paramo, 3700-4200 m., Paramo del
Quindio, Caldas, Aug. 15-20, 1922, Pennell & Hazen 10001
(US; ASP); grassy paramo, ‘‘Llano de Paletara," 2950-3100 m.,
June 15-17, 1922, Pennell 6930 (ASP; US); no data, Lehmann
4719 (US); Popayan, in paramo de Guanacas, Hartweg 1336 (NY).

Ecuapor: Quitensian Andes, 1855, Couthouy (GH).

Реко: 6 mi. south of Mito, 3076 m., Aug. 1-5, 1922, Macbride
& Featherstone 1821 (NY; MBG).

32. Satureia Darwinii Виа. in Engleru. Prantl, Nat. Pflanzenfam.,
ed. 1, IV. Abt. 3a, 300. 1897.

Micromeria Darwinii Benth. іп DC. Prodr. 12: 222. 1848.

Fruticulus humilis, prostratus, tegetes faciens, caulibus in basi
lignosis, ramis numerosis, subteretibus, pubescentibus, internodiis
maximam partem minus quam foliorum longitudine; foliis 4-5
mm. longis, obovatis vel subspatulatis, obtusis, in basi angustatis,
paginis ambobus pubescentibus; floribus in axillis solitariis, pedi-
cellis cirea 1 mm. longis, bracteolis duobus supra medium positis;
calycibus vix 4 mm. longis, 15-venis, extus pubescentibus, den-
tibus 1 mm. longis, ovato-lanceolatis, obtusis, subaequalibus, in-
eurvis, intus hirsutis, corollis 5 mm. longis, tubo incluso, labro
bifido, labioli lobo medio majore; staminibus inclusis, vix 1 mm.
longis, thecis sub-parallelis; nuculis von visis.

Specimens examined:

ARGENTINA: S. Patagonia in patches everywhere, Nov. 15,
1896, Peterson (NY); Patagonia, 50/30, 1882, Moreno 186 (NY);
Killikaike, Patagonia, Brown 65 (NY).

Жы, ЧС =

1927)
EPLING— SOUTH AMERICAN LABIATAE. III 79

33. Satureia pusilla Macl. in Rept. Princeton Univ. Exp. to
Patagonia 8: 698. 1905.

Micromeria раза (sic) Phil. Anal. Univ. Chile 90: 556. 1895.

Herba perennis humilis, altitudine 5-6 ст., caulibus in basi
duris, repentibus, ramis numerosis, gracilibus, puberulis, sub-
teretibus; foliis 3-4 mm. longis, obovatis, in basi angustatis,
utrinque puberulis, subsessilibus; floribus in axillis solitariis,
pedicellis circa 2 mm. longis, bracteolis duobus ad medium
positis; calycibus 3.5 mm. longis, extus hispidissimis, setis paten-
tibus, 13-venis, bilabiatis, dentibus tribus posticis cirea .4 mm.
longis, anticis circa .6 mm., omnibus ovato-triangulis, acutis,
saepe purpurascentibus, intus hirsutis, margine ciliolata; corollis
vix 5 mm. longis, labro bifido, labiolo longiore, lobo medio majore;
staminibus inclusis, vix 1 mm. longis, thecis parallelis; nuculis
non visis.

Specimens examined:

ARGENTINA: Gregory Bay, Magellan Sts., Nov. 23, 1886, Saf-
ford (NY, type collection).

34. Satureia vana sp. nov.

Frutex foliosus altitudine 2 m., ramis teretibus, ramulis di-
varicatis, quadratis, puberulis, internodiis folia subaequantibus,
foliis 8-12 mm. longis, late ovatis vel subrotundatis, obtusis, in
basi saepius cuneatis, rugosis, margine supra medium subserratis,
paginis ambobus puberulis, petiolis circa 1 mm. longis; floribus
3-6 in foliorum axillis, verticillastris subsessilibus, bracteolis 1
mm. longis ornatis; calycibus 3.5 mm. longis, in basi leniter an-
gustatis, 13-venis, extus praecipue ad venas hispidulis, dentibus
circa 1 mm. longis, ovatis, acutis,
tamen longioribus, intus tenuiter hirsutis, pedicellis subnullis;
corollis 7-8 mm. longis, extus villosulis, tubo superne тайа
dilato, labro 1.5 mm. longo, bifido, labiolo paulo longiore, lobo
medio majore; staminibus didymis, breviter exsertis, thecis
divaricatis; nuculis non visis.

A nondescript species singularly devoid of any marked char-
acter, but combining characteristies of sections Gardoquia and
Xenopoma.

Specimens examined:

[Vor. 14
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

PERU: rainy-green formation, 3100 m., Carumas, Moquegua,
Feb. 21-March 6, 1925, Weberbauer 7259 (FM, түре).

35. Satureia boliviana Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. За, 300. 1897.

Micromeria boliviana Benth. Lab. Gen. et Sp., 731. 1835;
et in DC. Prodr. 12: 222. 1848.

Xenopoma bolivianum Griseb. in Goett. Abh. 25: 272. 1879.

Satureia Kuntzeana Briq. in Engler а. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Frutex ramosus altitudine 30-90 cm., caulibus teretibus glabris,
cortice discedente, ramis ramulisque puberulis, gracilibus, quad-
ratis, angulis acutis, submarginatis; foliis .5-1.5 cm. longis, mag-
nitudine formaque diversis, eis in ramulis sterilibus diffusis,
ovato-ellipticis, obtusis vel subacutis, in basi cuneato-angustatis,
margine frequenter obscure serrata et revoluta, plerumque 1-1.5
em. longis, eis in ramulis florentibus densis saepius oblongis,
obtusis, in basi angustatis, margine integra et revoluta, plerum-
que .5-1 em. longis, omnibus utrinque puberulis floribus soli-
tariis in axillis, subsessilibus et in paniculis brevibus lateralibus
dispositis (frequenter tamen in ramis substerilibus tribus in axillis
pedunculo breve elatis );calycibus З mm. longis, tubulosis, den-
tibus lanceolatis-acutis, subaequalibus, corollis 8-9 mm. longis,
pubescentibus, tubo superne patenter ampliato, intus ad medium
breviter piloso, labro 1 mm. longo, emarginato, labiolo 2 mm.
longo, lobis subaequalibus, rotundatis, medio paulo majore;
staminibus didymis 2-2.5 mm. longis, supra tubi medium in-
sertis, thecis paulo divergentibus; stylo corolla aequilongo;
nuculis non visis.

Satureia Kuntzeana was based by Briquet on a collection by
Kuntze at Tunari, 4000 m., Арг. 1892. It was referred by him
to the Section Gardoquia to a position near S. microphylla (S.
connata nom. nov.) and S. Jamesoni with the following note “‘ Les
corolles sont blanches au lieu d'étre écarlates, c'est encore une
exception qui fortifie notre conception des Gardoquia comme
simple section du genre Satureia.”’ A sheet at the U. S. Nat.
Herb. bearing the above-cited collection-data and labelled
in Kuntze's hand “Satureja Kuntzeana Briq.," is assuredly 8.

1927]
EPLING—SOUTH AMERICAN LABIATAE. III 81

boliviana. It corresponds in every way to Briquet’s description
of the type.

Specimens examined:

Bourva: Cotani, 2450 m., Sept. 1911, Buchtien 5878 (US); La
Paz, 2653 m., Aug. 15, 1901, Williams 1674 (US; NY); Larecaja,
Sorata, 2650-3300 m., Feb. 1857-July, 1858, Mandon 517 (СН;
NY); an sonnigen Abhingen, 3600 m., La Paz, Sept. 3, 1906,
Buchtien 488 (US); Tiahuanaco, 3900 m., Nov. 1913, Buchtien 438
(GH; NY); La Paz, sonnige Abhàánge, 3700 m., Nov. 3, 1906,
Buchtien (GH); vic. of La Paz, Aug. 15, 1914, Rose 18894 (US;
МУ); по data, Bang (NY); viciniis Achacache, monticula Arichaca
in petrosis, 4000 m. alpine, Jan. 1859, Mandon 518 (apparently a
dwarf from 15 to 20 em. tall, otherwise about the same) (NY);
Tunari, 4000 m., Apr. 5, 1892, Kuntze (type collection of S.
Kuntzeana Briq.) (US); Sirupaya bei Yanacachi, 2100 m., Nov.
28, 1906, Buchtien 315 (US) ; Unduavi, 3076 m., Oct. 1885, Rusby
1500 (ASP; US; СН; NY).

Var. tarijense comb. nov.

Xenopoma bolivianum Griseb. var. tarijense Wedd. іп Griseb.
Symb. Fl. Argent. 272. 1879.

Foliis patenter serratis, crenarum culminibus acutis, inter se
1-2 mm. distantibus, pagina inferiore pallidiore; corollae tubo
calyce duplo longiore.

Specimens examined:

ARGENTINA: Cuesta de Tesanca et del Inca, May 25, 1873,
Lorentz & Hieronymus 1087 (NY).

36. Satureia simulans sp. nov.

Frutex ramosus foliosusque altitudine circa 2 m., ramis tereti-
bus vel subquadratis, cortice discedente, ramulis ascendentibus,
pubescentibus, quadratis, angulis obtusis; foliis 1-2 cm. longis,
lanceolatis, acutis, in basi rotundato-cuneatis, utrinque breviter
pubescentibus, margine revoluta, subserrata, petiolis 1-2 mm.
longis elatis, floribus 3-6 in axillis, pedunculis brevibus elatis,
rarius in axillis superioribus solitariis, bracteolis cirea 1 mm.
longis ornatis; calycibus 3.5 mm. longis, extus hirsutis, fauce intus
nudis, dentibus fere 1 mm. longis, lanceolatis, acutioribus, duobus
anticis quam posticis tribus paulo longioribus et saepe patentibus,

[Vor. 14
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN

pedicellis subnullis; corollis 6-8 mm. longis, tubo intus piloso,
superne gradatim dilato, labiis 1.5-2 mm. longis, labro emargin-
ato, sinu .5 mm. profundo, labioli lobis subaequalibus, rotundatis,
medio ad basim angustato; staminibus didymis, supra tubi
medium sitis, vix e tubo exsertis, thecis parallelis, .4 mm. longis;
stylo e tubo paulo exserto; nuculis von visis.

Planta aspectu 5. bolielana speciminibus substerilibus valde
similis, floribus tamen differt.

Specimens examined:

Botrvia: Unduavi; Nord Yungas, 3300 m., Nov. 1910, Buchtien
2954 (US); Unduavi, shrub 2 m., 3300 m., Nov. 1910, Buchtien
2955 (US, TYPE); Sorata, Apr. 19, 1920, Holway 550 (US).

37. Satureia axillaris (Rusby), comb. nov.

Bystropogon ахШате Rusby, Mem. Torr. Bot. Club 6: 108.
1896.

B. uniflorus Busby in Виа. Bull. l'Herb. Boiss. 4: 802. 1896.

Suffrutex diffusus, ramosus, altitudine ad 1 m., caulibus ramis
ramulisque gracillimis, glabris, quadratis vel caulibus teretibus,
angulis submarginatis; foliis 10-25 mm. longis, 1.5-3.5 mm. latis,
lineari-lanceolatis, rarius oblongis, apice acutiusculis, in basi ad
petiolum brevem vel subnullum angustatis, utrinque omnino, gla-
bris, margine subrevoluta, obscurissime et sparse serrata; floribus
in axillis solitariis, bracteolis parvis, subulatis, quam pedicellis
brevioribus subtentis; calycibus 1.5-2 mm. longis, tubulosis,
dentibus lanceolatis, acutis, patentibus, subaequalibus, fauce
intus glabris; corollis calyces vix superantibus, 2.5 mm. longis,
tubo superne ampliato, intus glabro, labro emarginato, labioli lobo
medio patente, rotundato, 1 mm. longo, staminibus minutissimis
ad tubi medium sitis; stylo corolla aequilongo; nuculis non visis.

Specimens examine "i:

Bourva: Bolivian plateau, 1891, Bang 1125 (type of Bystro-
pogon axillare Rusby) (US; ASP; MBG; GH); Sierra de Santa
Cruz, 1600 m., May, 1892, Kuntze (NY).

38. Satureia brevicalyx sp. nov.
Suffrutex erectus altitudine 50-60 cm., caule ut videtur virgato,
puberulo, quadrato, angulis obtusis, internodiis 2-4 cm. longis,

1927]
EPLING—SOUTH AMERICAN LABIATAE. III 83

ramulis numerosis brevibus, ascendentibus, gracilibus, puberulis;
internodiis 3-10 mm. longis; foliis 3-5 mm. longis, ellipticis vel
ovalibus apice saepius obtusis, in basi rotundato-angustatis,
margine revoluta, subintegra, utrinque molliter hirtellis, petiolis
cirea 1 mm. longis elatis; floribus solitariis in axillis, numerosis,
bracteis subulatis, quam pedicello brevioribus subtentis; calyci-
bus 3 mm. longis, late tubulosis, subbilabiatis, fauce intus nudis,
dentibus ovatis, obtusis, subaequilongis; corollis 5-6 mm. longis,
extus pubescentibus, tubo leniter arcuato, superne ampliato,
intus infra stamines hirtello, labro subrecto, ovato, emarginato,
labiolo quam labro paulo longiore, lobo medio obovato, majore;
staminibus 1 mm. longis, inclusis, thecis parallelis, supra tubi
medium sitis; stylo paulo exserto; nuculis ovatis, fuscis, .5 mm.
longis.

Specimens examined:

Perv: Panticalla Pass, 3600 m., July 16, 1915, Cook & Gilbert
1877 (US); “Bolivian and Peruvian Andes" (GH); Cuzco,
March, 1925, Herrara 825 (US, TYPE).

39. Satureia oligantha Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. 3a, 380. 1897.

M ا‎ EY Benth. Lab. Gen. et Sp. 381. 1834; et in
DC. Prodr. 12: 222. 1848.

Xenopoma eugenioides Griseb. in Goett. Abh. 19: 237. 1874.

Micromeria eugenioides Hieronym. in Acad. Nac. Cordoba,
Bol. 4: 413. 1881.

?Bystropogon minutus Briq. in Bull. Herb. Boiss. 4: 803.

96

Satureia Gilliesii Briq. in Engler u. Prantl, Nat. Pflanzenfam.,
ed. 1, IV. Abt. За, 300. 1897 (sub Sect. Xenopoma).

S. eugenioides Loes. ex Robt. Fries in Nov. Acta Soc. Upsal.
IV. 1: 107. 1905.

Suffrutex ramosus altitudine circa .5 m., in basi lignosus, caule
terete, cortice discedente, ramis ramulisque puberulis vel tenuiter
pubescentibus, gracilibus, subtortis, quadratis, angulis acutis;
foliis 3-8 mm. longis, approximatis, oblongis, obtusis, utrinque
puberulis vel pubescentibus, glandulosis, margine subrevoluta;
pedicellis .5-1 mm. longis, subsolitariis, tribus tamen frequenter

$ | [Vor. 14
84 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in axillis inferioribus pedunculo breve elatis, bracteolis subulatis,
pedicellis aequilongis ornatis; calycibus parvis, 1.5-2 mm. longis,
campanulatis, dentibus subaequalibus, acuminatis, patentibus et
paulo recurvis, fauce intus subvillosis; corollis calyces vix super-
antibus, 2.5 mm. longis, tubo superne ampliato, intus glabro,
labro emarginato, labioli lobo medio patente, rotundato, 1 mm.
longo, staminibus minutissimis ad tubi medium sitis; stylo corolla
aequilongo; nuculis oblongis, .7 mm. longis.

Specimens examined:

Рено: above Chivay, Coilloma, Arequipa, March, 1914, Weber-
bauer 6891 (FM).

ВошутА: Oruro, Тарасагі, 4000 m., March 17, 1892, Kuntze
(type collection of Bystropogon minutus) Briq. (NY).

ARGENTINA: Sierra Farmatura, 1873, Lorentz & Hieronymus
(NY); Cuesta de la Muschaca, Catamarca, Feb. 1876, Schicken-
danz 954 (NY); Sierra de Tucuman, Jan. 10-17, 1878, Lorentz &
Hieronymus 728 (US); Capillitas, Catamarea, 1875, collector un-
known (ASP); El Candado, Andalgalá, Oct. 2, 1916, Jörgensen
1189 (MBG; UC).

SPECIES Міні OMNINO IGNOTAE

Micromeria pulchella Wedd. Chlor. And. 2: 151. 1857.

Gardoquia salviaefolia Colla Mem. Aecad. Torin. 39: 2. 1836.

Satureia bonariensis Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 300. 1897.

Micromeria bonariensis Fisch. et Mey. Ind. Sem. Hort. Petrop.
10: 56. 1845. А

“hispidula, caulibus erectiusculis, foliis linearibus punctato-
glandulosis integerrimis, verticillastris sexfloris, corollis inclusis.
In Bonaria unde sem. misit Bonpl. Thymus Bonariensis Ten. Ind.
Sem. Hort. Neap. 1839. Corolla in planta culta saepissime
calyptraeformis, non expansa, ex Fisch. Mey. An revera didy-
nama? Ап eadem ас Hedeoma multiflora?” (Benth. in DC.
Prodr. 12: 223. 1848.)

ее * ее,
1927]
EPLING—SOUTH AMERICAN LABIATAE. III 85

INDEX SPECIERUM

New species, varieties, and combinations are printed i in bold face type; synonyms,
{п italics; and previously published valid names in ordinary type.

PAGE

ачты
И аьа E TE E х 82
деде И 83
ТО CV e vo er IST NUS 82
DENEN Or Veo xov a RA Т 47
т’ е RANT TENDUNT T ES 63
о АТАС. то 71
КИЕВ ВАР АИ 59
и р 56
И ео 62
BENE. Lira Fever ТРИ РТ 54
ИР гаи dee 2 72
О Ea е 61
Е ооо. « 56
ПЕНИ Tos Ale Е. 66
раму вает 64
ен 69
о PEPEPEPE И 70
DIEI. c.i per S 53
VIT DUE PEE У а 58
ТА АГТС” 74
ОАТС КТП ОҒ 76
ОИ Ч, 252,222. 84
ОЛБАНИ ie ET 73
CUR Aer c ТЕГІ СОК cue 68
a. Lr МА xs ok) Тет РР 67
Н ОРО 61
е E Ч ENS 60
ИО вно нт 47
ТӘНИ АСС. АМЕ à 80
О oid oed 84
LIGrOU NE S поно: 78
eundo leq каны, v stop pad se 83
ӨШИ aa e ts 83
fiubigenaz са ONES ER re Ne 76
bonia... oo culis neca Те 79
писма, „око а « 84
Micromeria sect. Hesperothymus 50
ос ес ДР 47
COAT OMG O as ОЕ от 53
D MEE IAM а ЎР 47
— Discolores; Ae E A 59
—Hihpticae... v... 9-209 - 53

PAGE
Боя ORT VOTE TELA Se T ETT. 56
а’... ОРС xao +» жа» 71
ЛЕ NS 65
ACUGIONS. СУ 63
Пост rere N 67
argentea. еи 71
ап, ARAS а EA 82
boliviana. оао 80
boliviana var. tarijense.......... 81
bonarlenms....:2. 5 УЬ 84
brevicalyx. iu AAR 82
е ova bs EDO SC EA 59
мер,
Brow subsp. eubrownei....... 51
ре ан 56
ata i Е И 70
Пагу d EMI QR AS 78
discolor; P AERA. 62
elliptica. ... SOEUR 54
TR c PEE PETTY ENTE EE EET TT 70
о 25%, are oe em 83
ТОА аа рУ 72
осо АКТ ТИ s eoa E 61
qut oc АРК 56
i ате r PAra E o SASA 83
EMEN роге 66
се B Dex X xard e 58
И РИО e t e 74
а a hows Pine ов 69
NOMEN GREE Ae quA Sede swa 80
Тор лиле S PUR 75
O саны age ти ж» 55
РР O ық NU V STER 63
И. ОИ 57
о ы ы ыы 70
Iu Па ах 53
MEE ес rA S 76
часе уат, glabrescens....... 78
ООО С; ғаты» 58
eue EUER owes ERI TR 83
ada у ср Ты 65
Ре. 64
о рр РНЕ 69

[Vor. 14, 1927]

86 ANNALS OF THE MISSOURI BOTANICAL GARDEN
PAGE PAGE
АТ 70 Stachys

УЕ Ғы... 74 о 6 Ұ% ба ЛҮ ТӨР 54
РОСС m2 TEE. У 47
АРСО 75 о Ио 51
ПРО E CLE PERLE 73 ор ықы, ыы мере 76
ЭА. Л PD сағ 81 РОТИ P A 76
LLL oa et ERG. 68 NN Ее 67
ЭЛК Viv LZ ТҰР QU ANM. ЕАО 47
о МЧСКЕ Р EET S 61 о EE m E Л КУКЕ 80
о. зул ле нд уч 60 bolivianum var. tarijense......... 81
секао со XA аена 79 о РТР 83
boo МАКСИ 72

Annals

of the
Missouri Botanical Garden

Vol. 14 APRIL, 1927 No. 2

A MONOGRAPH OF THE GENUS PHOLIOTA
IN THE UNITED STATES

L. O. OVERHOLTS

Professor of Botany, Pennsylvania State College,
M ycologist to the Missouri Botanical Garden and Visiting Professor in the Henry Shaw
School of Botany of Washington University

I. Aims AND METHODS

Several years of study of the higher Basidiomycetes have brought
to the writer the conviction that until very recently mycologists
have overlooked many of the important details in the structure |
of these plants. Хо genera are more significant in this respect
than Corticium, Peniophora and Hymenochaete of the Thelephorac-
eae; Flammula, Hypholoma and Pholiota of the Agaricaceae; and
Lycoperdon and Bovista in the Lycoperdaceae. In these and in
many other (perhaps nearly all) genera the microscope reveals
contrasting details of structure that were entirely unknown to the
earlier generations of mycologists. These details have been so
much neglected that it may be truthfully said that the current
conception of species in fungi has lagged considerably behind
that of seed plants. Many of the so-called species іп our myco-
logical literature have been in reality species complexes. While
it must be admitted that specific limitations must always be to
some extent a matter of individual opinion, yet undoubtedly
taxonomists will more nearly agree in these opinions when the
details of structure become a part of the current knowledge about
these plants. The facts concerning them should be set in orderly
array, open to the inspection and criticism of all. Descriptions
alone will not so present them. Much may be said against

ANN. Мо. Bor. Garp., Vor. 14, 1927 (87)

[Vor. 14
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN

running each description, as it were, into а mold or through а
mill, hewing them all down to the same thickness and cutting
them all off to equal length; yet after all it is the only way to
make the plants stand in contrast to each other. How often
one comes upon a descriptive item that he suspects may be of
prime importance, only to find, when seeking a contrast with
another species, that no mention is made of the presence or ab-
sence of this characteristic. Single points of difference do not
usually make a species, to be sure; therefore, all the more reason
for available diagnoses that will enable one to contrast two or
more species on all points. It is the aim of this paper to so con-
trast in particular the microscopic details within one genus of the
Basidiomycetes.

Тһе paper is the outgrowth of a critical study of the species of
the genus Pholiota of the Agaricaceae for the ‘North American
Flora. Тһе drawings illustrating the microscopic details are all
made to the same scale in the different species so that direct
comparisons сап be made. АП were outlined under the camera
lucida from free-hand transverse sections of the gills of dried
plants after softening with aleohol and water, mounting in KOH,
and staining with dilute eosin. Тһе larger drawings of the spores
were made in the same way except under the oil-immersion lens,
yielding а magnifieation of about 1800 diameters. Тһе spore
measurements represent normal variations of what are believed
to be mature spores, a series of usually not less than 10 spores
being measured for each species.

II. HISTORICAL

Only very recently has this genus received critical study at
the hands of American mycologists. In 1908 Peck published!
one of his latest and best monographs, dealing with this genus in
the state of New York. It was followed in 1912 by the work of
E. T. Harper? on the species occurring in the region of the Great
Lakes and to which additions were made in two subsequent
papers. Тһе illustrations published by him constitute the finest

! Bull. М. Y. State Mus. 122: 141-158. 1908.

?'Trans. Wis. Acad. Sci. 17: 470—502. 1912.
3]. c, 17: 1011-1014. 1913; 18: 392-405. 1916.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 89

work of the kind that has appeared in America. Yet his descrip-
tions were in many cases inadequate, though possessing the ad-
vantage that many of them were drawn from the plants
as he found them growing in nature. Yet many were only quo-
tations from the earlier work of Peck. His most important con-
tribution, aside from the illustrations, was the fact, presented for
the first time, that the species may be grouped into phylogenetic
units with rather well-marked characteristics for each group, the
species within these units differing in lesser degree. Тһе impor-
tance of this grouping cannot be ignored yet can be over-em-
phasized to the confusion of one not well acquainted with the
genus. Without access to material of authentic European species
he was forced to rely on published descriptions, and to some
extent he was misled by erroneous identifications by others. In
1918 appeared Kauffman's work! on the gill fungi of Michigan in
which the species were well described but inadequately illustrated.

The contribution that I have attempted to make in the present
paper has been largely in the matter of the details of microscopic
structure of the hymenium. It is a matter of constant surprise
to see how easily species may be separated by this means; and
by a combination of microscopic and macroscopic characters
most of the species take their place with a definiteness that is
remarkable. Even where macroscopic features are in them-
selves characteristic enough to delimit species it is reassuring to
be able to confirm a determination by a set of facts deduced from
an entirely different source.

While this sort of work cannot be said to be а new angle of at-
tack for taxonomie work in the higher Basidiomycetes, yet in-
stances where it has been systematically used in the gill fungi are
remarkably scarce and no instance has come to my attention
where the microscopic characters have been enumerated and il-
lustrated to а sufficient extent in a single genus of that group.
Тһе nearest approach to this has been in the work of Lange оп
the Agarics of Denmark. However, the illustrations there are

1 The Agaricaceae of Michigan. Mich. Geol. & Biol. Surv. Publ. 26: 1-899. 1918.

2 Lange, J. E. Studies in od Agarics of ае I. The genus Mycena.
(Dansk Bot. Arkiv 1*: 1-40. 14); II. Amani Lepiota. Coprinus. (l.c.
2% 1-53. 1915); ІП. ты Lol In aiv (1. с. 27: 1-50. 1917); IV.
Pholiota. Marasmius. Rhodophyllus. (І.с. 21: 1-46. 1921.)

[Vor. 14
90 ANNALS OF THE MISSOURI BOTANICAL GARDEN

woefully inadequate on this point so that the entire effect is not
much better than the rather complete descriptions of microscopic
characters presented by Kauffman.

ПІ. PREPARATION OF THE KEYS

In preparing the key to species I have been at some loss to
know on what points to make the larger divisions. A number of
possibilities present themselves for consideration. Previous writ-
ers have for the most part attempted to follow the scheme out-
lined by Fries, making the first division on the basis of habitat. I
have attempted to use the point of comparative sizes instead. In
any scheme there will always be some overlapping. At first
thought it might appear that in using habitat there can be no
overlapping, for either the plants grow on wood or else they grow
оп the ground. Yet in reality the matter is not so simple. Some
species grow on earth heavily charged with humus or very rotten
wood; some grow from buried wood, and in either case, unless
great care is exercised at all times in the field, the collector will
be at a loss to know how to refer his habitat or if unobservant will
name it wrongly. The weak point about most keys lies in the
failure to take into consideration these overlapping forms, and
when that is properly cared for the exact point on which the
division is made becomes of secondary moment, unless, of course,
one is interested in showing phylogenetic relationships in the
key. But phylogeny assumes or demonstrates similarities, and
the purpose of a key is to show contrasting differences. It is
impossible, therefore, to combine these opposing views in one key
with any high degree of satisfaction.

IV. MICROSCOPIC OBSERVATIONS

Both in the keys and in the descriptive matter I have laid con-
siderable emphasis on the cystidia and the spores. The cystidia
in this genus fall for the most part into three general categories.
These are represented as follows: (1) Text figures 1, 3, 5, 9, 15, ete.,
in which they are enlarged and quite conspicuous, varying from
almost round to fusoid. This type might in fact be broken into
two types, one that does not present a narrowed apex and one
that does. Yet they intergrade to a considerable extent. (2) Text

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 91

figures 40, 43, 46, 83, 87, etc., in which the buried base is enlarged
into а bulb, and the apex, that may be long and cylindrical, pro-
jects prominently beyond the basidia. (3) Text figures 127, 129,
131, 135, 137, 139, etc., in which they are brown in color, par-
ticularly in dried plants, and do not usually project prominently,
sometimes not at all. It has not been deemed expedient to il-
lustrate the fact that these last bodies are colored except in figs.
127, 129, and 135, where they have been darkened slightly. At
least in the key, measurements in diameter of all of these bodies
are measurements through the broadest part, which is usually
imbedded between the basidia. In the matter of spore sizes
rather narrow distinctions have been drawn in the keys at times,
and careful measurements of several representative spores should
be made before conclusions are reached. In general, the smallest
spores, except in cases where spore prints have been obtained,
should be disregarded, especially if they appear somewhat lighter
in color, since such are obviously immature. Spore markings, if
present, can usually be detected with the ordinary high power of
the microscope, especially if а 10 х ог а 12 х ocular be em-
ployed. In such cases, definite indications of roughness on the
part of some spores indicate that they should be classed as rough-
walled. An oil-immersion lens is a convenience in substantiating
roughness where dry-lens examination is not conclusive. Em-
phasis has also been laid on the appearance of a slight degree of
truncateness on one end of the spore in some cases (figs. 2, 8, 14,
95, ete.). This is usually an obscure point at first, but once de-
tected it can be rather readily recognized under the high-power
dry lens. That this truncateness is in reality on the apex of the
spore and does not represent its point of attachment to the sterig-
ma, as would at first appear, has been repeatedly verified.

V. NOMENCLATURE AND SYNONYMY

In the matter of nomenclature and synonymy I have used, for
all my plants, names that have been current in the present gener-
ation of mycologists, without attempting in any case to replace
well-established names with those of an earlier and often a doubt-
ful designation. I have, however, not hesitated to use current
European names in place of names of an American origin when

(Vou, 14
92 ANNALS OF THE MISSOURI BOTANICAL GARDEN

I have been able to satisfy myself of their equivalence. I con-
sider the genus to have been established to all intents and pur-
poses by Fries in the ‘Systema Mycologicum’ in 1821. For the
rest I have followed the International Code in the matter of
citations.

VI. PHYLOGENETIC CONSIDERATIONS

As has been mentioned on a preceding page, the species of
Pholiota fall into several rather well-marked phylogenetic units,
each with а central species around which the others may be
grouped. It may or may not be possible to select the central
species which would represent the ancestral types of the group in
each case, and the evolutionary development within a group may
not have taken place symmetrically but may have been all in
one general direction, but the fact remains that the species within
any one of these groups are more closely related to each other
than they are to the members of any other group. Тһе recog-
nition of these groups is a relatively easy matter, since in most
cases they do not intergrade. Оп the other hand, the boundaries
of the more recently recognized types within these groups are
often quite a difficult matter, since they are not always sharply
delimited. In the matter of marking out these more elementary
species, individual opinion must always play an important part,
and for that reason different investigators may not always come
to the same conclusion.

Since in the presentation that constitutes the bulk of this
article attention is directed to only the fundamental taxonomic
features, I shall here briefly consider these phylogenetie units
within the genus. Relationships are expressed in similarities in
what are believed to be the more significant and deep-seated
characters, and those that are less likely to vary under varying
external conditions. Two of the most important of these are the
spores and the cystidia. For example, the peculiar truncate
spore and the large cystidia, all of the same type and different from
almost all others in the genus, are to me а mark of relationship
between P. dura, P. praecox, P. vermiflua, P. temnophylla, P.
Howeana, and P. Acericola. Itso happens that these species agree
also, at least in a general way, in several other characters, e.g.,

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 93

general form and coloration, habitat, absence of all scaliness, etc.
In the same way there is а small group centering around P. erebia
that shows close agreement in spore characters and in the char-
acteristic cystidia with long projecting necks. Неге again, other
points, less easy to express quantitatively, leave no doubt that
we are dealing with a group of closely related species, and far more
closely related among themselves than to individuals of any other
group. But only about 80 per cent of the species here treated
show similarities by which they can be so grouped at present.
'The remainder must be carried as unattached species, perhaps in
some сазез representing а non-plastie or stabilized condition, at
least temporarily. Some of the forms that I have left unattached
were given more definite placings by Harper, to which I cannot
subscribe. These exceptions have been noted in the tabulation
that follows. Harper recognized twenty such groups or phylo-
genetic units, several of them monotypic and several not desig-
nated by name. After the first dozen have been marked out the
remainder are found to be more problematic. My own inter-
pretation of the situation at present follows.

In the first place, authors since the time of Fries have rather
consistently recognized two large divisions of the genus on the
basis of habitat. These are the sections that have been desig-
nated as the Humigeni and the Truncigeni, since their members
occur on the ground and on wood, respectively. Two or three
species are limited to a muscigenous habitat and have been desig-
nated as the Muscigeni. Within these large divisions the phylo-
genetic units appear as follows:

SECTION I. HUMIGENI

1. The Praecox Group.—With prominent broad cystidia and
spores truncate at the apex. General color ochraceous or paler.
No scales on pileus. й

P. praecoz, P. vermiflua, Р. Howeana, Р. temnophylla, P. Aceri-
cola (also P. dura of Europe).

ile P. Acericola frequently grows on rotten wood and perhaps

1 Items аге to be taken as characteristic of the groups in which they are mentioned.
"Therefore in using this synopsis as a key the plant is to be placed in that group with
which its characters best agree regardless of whether or not these are stated as con-
trasting characters for the groups.

[Vor. 14
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN

never on the bare ground, all of its other characters ally it with
this unit. Harper puts P. Howeana into a separate section with
P. ventricosa (= P. spectabilis), to which I cannot subscribe.

2. The Togularis Group.—Without cystidia. Spores truncate
(except in P. blattaria, at times). Ochraceous or paler in color,
and small in stature, with а median membranous annulus striate
on the upper surface.

P. togularis, P. blattaria, P. rugosa, P. filaris.

Harper includes P. anomala in this group. I have included it
among the unattached species.

3. The Erebia Group.—Spores elongate-elliptic, 10-15 џ long,
not truncate. Cystidia abundant, flask-shaped with long necks.
P. erebia, P. ombrophila, P. subnigra, P. platyphylla.

4. The Caperata Group.—Spores rough. Cystidia none. Pileus
orange-cinnamon or ochraceous-orange. Plants of rather large
stature.

P. caperata, P. Memurphyi.

Harper includes P. Johnsoniana here, but its characters make
it quite distinct in my opinion.

5. The Terrestris Group.—Spores smooth, less than 10 u long.
Cystidia rather indefinite but some sterile bodies present and
recognizable. Pileus scaly with small scales.

P. terrestris, P. angustipes.

This group may not be a natural one but does show some
transition over towards the T'runcigeni. While not known to
grow on wood, the latter species grows around stumps and is likely
attached to buried wood. Р. terrestris may eventually show such
an attachment, and both species bear in the hymenium a very
simple and inconspicuous type of sterile body that resembles to
a degree those of the Adiposa section of the Truncigeni. How-
ever, the group is advanced with some misgivings. P. albivelata
could be forced into the same alliance were it not for the entirely
glabrous and viscid pileus. If it were admitted, P. anomala
might be included also. Р. trachyspora would be included were
it not for its rough spores.

Unattached Species of the Humigent.—(1) P. albivelata. A rather

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 95

anomalous species that might stand as the type of a distinct
group. (2) P. anomala. "Too indefinite to place and may prove
to be referable to an older species. My study of the types would
deny its inclusion in the T'ogularis group as referred by Harper.
(3) P. aurea is difficult to interpret. It may grow attached to
buried wood and belong in the T'runcigeni as would be indicated
by its color. (4) P. duroides and P. Johnsoniana might be re-
garded as constituting a distinct group near the Praecox group but
neither would seem to belong well in that unit. Harper so refers
P. duroides and unites P. Johnsoniana with the Caperata group,
very evidently an erroneous conception.

SECTION II. TRUNCIGENI

6. The Marginata Group.—Spores mostly slightly rough at
maturity. Cystidia present, flask-shaped, with a narrow pro-
jecting neck (fig. 83). Rather small plants, with glabrous, hy-
grophanous pileus, striatulate when moist, watery-brown, drying
to ochraceous or cinnamon.

P. marginata, P. unicolor, P. discolor, P. furcata.

Harper includes P. marginella and P. mutabilis here, but they
differ in having slightly truncate spores and in lacking cystidia.

7. The Mutabilis Group.—As in the Marginata group but spores
slightly truncate, smooth, and cystidia absent.
P. mutabilis, P. marginella.

8. The Cerasina Group.—Reddish or ochraceous-orange species
with glabrous pileus, not hygrophanous. Cystidia none. Spores
smooth or rough.

P. cerasina, P. oregonense, P. rubecula.

9. The Spectabilis Group.—Gills bright-colored (yellow or och-
raceous-orange and remaining so on drying). Pileus dry, scaly.
Spores rough.

P. spectabilis, P. luteofolia, P. aeruginosa, P. curvipes (?).

10. The Adiposa Group.—Spores smooth. Pileus often viscid,
decidedly scaly. Stem usually scaly. With brown cystidia in
the hymenium. Large plants, fleshy and heavy, yellowish colors
predominating.

[Vor. 14
96 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. adiposa, P. aurivella, P. aurivelloides, P. squarrosa, P. squar-
rosoides, P. flammans, P. rigidipes.

I REA united here two groups, the Adiposa group and the
Squarrosa group of Harper, although they can be maintained
distinct on the basis of the different type of scaly covering on the
pileus. Р. limonella, I would prefer to class in the T'uberculosa
group, rather than here as in Harper's synopsis. Р. flammans
departs from the other species of the group in the small stature
and the small spores. Р. rigidipes has all of the characters of
this group except for the sparsely scaly pileus and stem, and the
more slender stature.

11. The Tuberculosa Group.—Spores smooth. Pileus viscid,
and both pileus and stem scaly. No cystidia. Small plants of
bright color, red or orange predominating.

P. tuberculosa, P. lucifera, P. limonella.

P. curvipes might be expected here rather than in the Spectabilis
group, but departs from the characters enumerated above in the
lack of viscidity on the pileus.

12. The Muricata Growp.—Small plants with erect-pointed
dark scales on pileus and stem. Хо cystidia. Spores smooth.
P. muricata, P. erinaceella.

18. The Fulvo-squamosa Group.—Large plants with fibrous eu-
ticle that soon separates into scales. Not viscid. Spores smooth.
Cystidia hyaline if present. Taste and odor of radishes.

P. fulvo-squamosa, P. Schraderi.

In general these two species are somewhat similar but neither
is well known.

Unattached species of the Ттипсідеті.-(1) P. Aegerita. Close to
the Mutabilis group in some respects but the evidence is not clear.
(2) P. albocrenulata. Is really the type of a distinct group, with
its large ellipsoid spores 11-15 и long. (3) P. destruens is a well-
marked type which should be set off by itself or with P. comosa
and P. heteroclita if either of these should be held distinct.

SECTION III. MUSCIGENI

14. The Mycenoides Group.—Very small plants among Poly-
trichum or Sphagnum mosses.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 97

P. mycenoides, P. minima.

There can really be no relationship between the two species
included here. The former has smooth truncate spores 9-11 y.
long, and the latter has spores not truncate, rough-walled, and
6-8.5 u long. The relationships of the former may be with the
Togularis group.

As to the inter-relations of these groups little that is definite
can be hazarded. In some respects the Caperata group might be
considered as the more advanced of the Humigeni. This argu-
ment would utilize the fact that in P. caperata at least there is
present a more or less definite universal veil that leaves at times
а, second annulus-like structure comparable to an imperfect volva,
atits base. For that reason the species is sometimes separated
off in the genus Rozites. Тһе spores аге rough-walled, which,
other things being equal, must probably be considered an ad-
vanced character. Yet otherwise the organization of the hy-
menium is not so complex as in many other species. Yet each of
these groups in the Humigeni presents at least one such mark of
advancement, assuming that the more complex structures in-
dicate advance in proportion.

In the Truncigeni, the Marginata group represents probably
the most advanced condition in one line of development, but is
probably exceeded in another line by the Adiposa group, in some
species of which one finds the most highly organized sporophores
іп the genus. Тһе development of cystidia, the high degree of
coloration, and the presence of an internal medulla in the trama
of the gills (figs. 142, 137, 131, 129, etc.) are marks of advance
over plants with the opposite of these characters.

That either these sections or phylogenetic units can be соп-
sidered as genera is unthinkable. Тһе practice of breaking up
the large genera of the fungi does not commend itself, in general,
as the proper plan of procedure. In a considerable part of such
work that has appeared the limits of genera encroach too closely
on those of species. There is still room and use in the taxonomic
literature for such а group as the Sub-Genus or the Section, and
the confusion that results in attempting to fit new generic names
to old ideas that are well established is lamentable.

[Vor. 14
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN

VII. ACKNOWLEDGMENTS

In the preparation of this paper I have had access to a consider-
able series of European specimens from Romell and from Bresa-
dola and have studied our plants with reference to them. Asa
result, some of my conclusions as to the identity of our species
are at variance with those of my predecessors. Тһе majority of
the specimens with which I have worked are preserved in five
different herbaria: that of the New York Botanical Garden, which
contains by far the most extensive collection; Peck's herbarium
at Albany, in the New York State Museum; the herbarium of the
Missouri Botanical Garden at St. Louis; the herbarium of the
Pennsylvania State College at State College, Pa.; and in my own
herbarium. Му identifications are on the specimens in all of
these places, so that I have not felt it necessary to present here
all of the data required to identify the particular collections.
Consequently, my citation of specimens is limited to data that
concerns only the stations involved, from which the geographical
range may be obtained at a glance.

Acknowledgments for assistance are gladly given to Dr. W. A.
Murrill and the New York Botanical Garden for free access to all
collections of the genus at that institution; to Dr. H. D. House for
the privilege of studying the many collections in the New York
State Museum at Albany; to the Missouri Botanical Garden for
access to the herbarium there; to the officials of the Pennsylvania
State College for their interest and indulgence in the work; to Dr.
R. A. Harper and the Wisconsin Academy of Sciences, Arts, and
Letters for permission to reproduce some of the plates published
by the late E. T. Harper; likewise to Mr. L. W. Brownell of
Paterson, N. J., for original photos presented in plates 16, 17,
20, 21, and 24; to Mr. Burtt Leeper, Salem, Ohio, for plate 18; to
Mr. W. S. Odell for plate 12 (photo by Drayton); to Dr. E. T.
Harper for the photo in plate 11, sent to me a short time prior
to his death; to Mrs. Edna G. Stamy Fox for the final touches
that have contributed so much to the appearance of the text
figures; and to many others who have contributed specimens or
in other ways have aided in the prosecution of the work.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 99

VIII. Taxonomic CONSIDERATIONS
PHOLIOTA

Pholiota Fries, Syst. Myc. 1: 240. 1821.

Plants fleshy, putrescent, solitary to cespitose, geophilous or
xylophilous; pileus glabrous to floccose, fibrillose, or distinctly
scaly; lamellae adnexed to slightly decurrent; stem central,
fleshy, glabrous to fibrillose or scaly, continuous with the flesh
of the pileus; veil present, usually membranous, sometimes fibril-
lose, forming a distinct, though often evanescent, annulus; spores
ochraceous-brown to ferruginous or fuscous, smooth or rough;
cystidia often present.

Type species: Pholiota dura (Bolt.) Fries.

The plants in this genus are analogous to those in Armillaria
of the white-spored series and in Stropharia of the purple-brown-
spored series. At some points the genus grades into Flammula,
due to the early disappearance of the partial veil or the annulus,
and in Flammula the veil fragments may at times persist as an
incomplete annulus.

KEY TO THE SPECIES!

1. Plants at maturity not more than 4 ст. ітояй............................ 2
Plants at maturity 4-15 cm. or more BORI A eee a a ea 29
2. Plants growing on the ground or among шоввев........................... 3
Plants growing on stumps, trunks, or rotting Іорв........................ 14
3. Pileus distinctly fibrillose or squamulose, especially at the center. ........... 4
Pileus entirely glabrous at the center, the margin occasionally with white
Бон Dun De v0l........... селен DES VEM LE n rd REESE 5
4. Stem distinctly scaly below the sheathing veil remnants; gills bright

cinnamon in dried plants; known only from Oregon; spores smooth,
5.5-0/,5 5X 8/0-4.D à... Es ct he pm 20. P. terrestris
Stem not scaly, somewhat fibrillose; gills deep brown in dried plants;
known T from Colorado; spores ое 7-9 х
.19. i feror
5. Plants growing among Былай MOSHER ҒАНА su cit s cu di
Plants growing in wet places among Sphagnum mosses........ 9. к esra
Plants growing оп the ground or оп leaf тоіа............................ 6
. Pileus slimy-viscid; stem heavily white-floccose below the annulus; known
only from Washington and Огероп....................... 22. P. albivelata
Pileus and stem not as above... 2.0%... . . ИЕ ЕСИН 7

e

1 For a supplementary key to the species see p. 169. І should also be noted that
the phylogenetic arrangement on p. 95 can be used as a key to small groups of species,
and that the synoptical key in the text, beginning on p. 104, can be used to the same
end.

[Vor. 14

100 ANNALS OF THE MISSOURI BOTANICAL GARDEN

7.

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—
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к.
e

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сл

ка
-I

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о

Cystidia present and projecting conspicuously from the sides of the gills
КЕСЕК ТЕМЕЛЕР). 2525 eK ee 8
о м ИНАЯ: pk o rn НАЧ oo REA n 11

. Cystidia 15-20 шіп diameter at the broadest part; spores truncate at one

end, 8-105 в lon |I 9

Spores 4.5-6 и Nos: а OU Des voee d e ЕА £P. сот
BEN DT БАИЯ ИРИС 6. P. temnophylla
. Plants small, less than 2 cm. in diameter, uniformly fuscous in color;
known only fróm Washington, ... ACL root ot 18. Р. subnigra
Plants somewhat larger; lighter in color than the above; spores 10-12 u
oe eee Pere Ce ПЕРИ E T 11. P. platyphylla
Plants as in the last but spores up to 15 и long.................. 12. P. erebia

. Stem 1-2.5 em. long; pileus yellowish-red or dark ferruginous and not

лет ео: ИЕР ОЕ УЛОУ ОЕ УТ 8. Р. гидоза
Stem 2-10 cm. long; pileus watery-brown, drying to ochraceous............ 12

. Stem filiform, about 1 mm. thick; pileus not more than 2 em. broad.

Vom ice Е алк 10. P. filaris
Stem not filiform, 2 mm. or more thick; pileus at maturity more than 2

DD AUTORE. s 5 iste He a rel ҚҰЛАУ ОИ T oa ИЕ RAUS PUE + 13

WX lcu cR А ПА А Е И 6. P. blattaria

ONE АЛТ АРНЕ ВОДИТ 7. P. togularis

. Pileus entirely glabrous and hygrophanous,! the margin usually striatulate

V wel UA es es oe a sara ok is coke ec cave АСОИ
Pileus scaly? or densely fibrillose from the first or becoming so in mature
plants; hygrophanous only in P. confragosa, but often viscid............

. Stem distinctly scaly; gills not іогКей....................... 30. P. mutabilis
Stem not at all scaly; gills іогкей............................ 28. P. furcata
Btem not МУ: ie not ен свое оь оао шанаа 16

. Pileus — ey. RU MAU vu) ENT aaro acis VC a DEM a Е 27. P. discolor
PINE BU ҰЙЫ Los en VIEWED CERT ASE be Ur CY ON CR ion Rn Ro ER M «e

. Stem wei "with ЖҮРЕ transverse partitions; known only n Cali-

ПИША on aul PREIS из lesa eo ok ho hin x ЗА ae . P. anomala
Stem stuffed or hollow, Т. internally partitioned; on logs or on Nr dh puer
. Cystidia absent; spores entirely smooth, slightly truncate at ^ end (fig
(| ARCEM СК Ae Phe SNe a A E eed e ^ ы
ee present, flask-shaped, E (fig. 83); spores ig te N
tha usually somewhat roughened when mature..............
А лейк f uel жен: persistent, frit nu pileus never more than
Bom: beondl. ок Н ана аен е с ааа нф е 26. Р. unicolor
Annulus not funnel-shaped, sometimes — as а ring but often
evanescent; pileus often 3-4 cm. or more broad. ........... 25. P. marginata
. Pileus dull cinnamon, dull tawny, or Мотп,............................ 21

Pileus bright-colored, i.e., pinkish-red, ب‎ ochraceous-orange, or
Du ла №, ВИНИТ ИЕ РЕ

11. е. watery-brown when moist, ochraceous as the pileus begins to lose moisture.

? Sometimes young specimens are only fibrillose, the scales appearing as the plant
matures; sometimes the young plants are conspicuously scaly, the scales gelatinifying
with age and often disappearing.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 101

21. ue densely floccose-fibrillose or floccose-squamulose as seen under а
ҒАР ee ped be» tes vox Ar. sn 5300 ee REPRE 7. P. confragosa
Pileus densely covered with small erect spine-like scales....48. P. eri l
22. Spores 3-5.5 и long; dried plants with a yellow тыту арреягапсе.
ЕЕ ХАН ЖЕ Е TL FA TE 88. P. flammans
Spores 6 » or more long; pileus not рауегШепі.......................... 23
93. Рика ium Ғы. 1...2222....... Жел АК eee sensi 24
Pilas do wia OD, aah i i ОА 26
24. Pileus lemon-yellow or bright sulphur-yellow..........................
Pileus with greenish or purplish tinges and not uniformly nor brightly
MIENNE T.V ағы жық У Рр. hire re ka 9 lammula polychroa!
25. Stem concentrically ringed or peronate with white fibrils....... 50. P. lucifera

Stem not peronate, but with scattered recurved yellow scales. .51. P. limonella

26. Gills bright-colored, i.e., ochraceous-orange, bright ferruginous, or
ochraceous-buff, and remaining so in dried ріапёз...................... 27
Gills not bright-colored, rather cinnamon or Бгояп....................... 28

27. Young plants bright-colored and only silky, becoming scaly A -—
when mature; taste mild; spores smooth. os esere io smese . P. curvipes

Young plants dark red and plush-like, becoming somewhat V pc and
pinkish-red when mature; taste bitter; spores minutely roughened (fi

еее IE eI moe TUERI 85. P. Wieofolía

. Pileus drying brown or cinnamon-brown, densely covered with tawny
fibrils or tufted and sub-erect scales (pl. 14, fig. 3); stem not bulbous
. P. muricata

Pileus drying ochraceous-tawny “ dark innate squamules; stem with

58.

bo
oO

CORSO TCHR ө ө ө о Фф ө ә э-э 4€ ө & $4 CWA 4 9» «49 B s RW Фа ж * 4 v t rs

a distinct bulb just at base (pl. 14, fig. 4)................. P. tuberculosa
29. Plants growing on the ground and Dio around old stumps and logs or from
buried wood. i... coc, ОИ а осире ари 30
Plants pint on wood, on sawdust piles, or around old stumps or rotting
logs, sometimes from buried тоой.................................... 42
30. Pileus entirely glabrous or at most only with marginal fibers from the veil... .31
Pileus fibrillose, Яоссове, squamulose, or scaly, at least over the center...... 37
81. Spores more than 10 џ опс.........2..552. EE UL ооочень 32
Spores averaging less than 10 и іопд.................................... 34

32. Cystidia pyriform to subglobose (figs. 1, 3, 5), not projecting conspicu-
ously, about 20 шіп diameter; spores ovoid with a truncate т" y 2)

ЭЛЕЕ СА OQ V waa КҮЛҮ RES DEREN DU US t P e P. vermiflua
Cystidia elongated, projecting prominently beta u) (fig. 43), з и in
diameter; spores elongate-elliptie, not truncate. ........................ 33
33. Pileus 2-5 em. broad; gills 2-4 mm. Бгояй...................... 12. P. erebia
Pileus 4-10 cm. broad; gills 4-6 mm. broad................. TR. и: багайын
34. Plants western; pileus slimy-viscid; stem conspicuously еа
just below the annulus. Noose ala E ое - e ыса

Plants eastern; pileus dry or very slightly viscid in very wet sous
stem not те4отмеісев.......-...2525%555455-42-.-2....52>,59%%2%4%
5. Spores 4—6 и long; annulus membranous, evanescent........... 23. P. duroides

e

1 This species is so frequently m ell-f Я
во I have fons it best to linde it in the key at this neni. No description is
given in the text

[Уох. 14

102 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

оз
с

e
~J

e
Q0

T
>

A
сл

ГМ
>

>
-l

Spores E нің и long; annulus not membranous but forming a cottony roll
nho Ж. ет E Pan T us КРИ КО КИШ" 24. P. Johnsoniana
Spores 55 ЛИН GRA Placa ore dead eee ek ку ГК Л mo ce 86

. Plants growing on the ground in cultivated fields or grassy places, ог in

open grassy woods, or among straw or other waste vegetable matter
carried into the woods; pileus not гарове..................... 2. P. praecoz '
Plants growing on the ground in dense woods; pileus not rugose. .4. P. Howeana
Plants growing on very rich humus around rotting logs; pileus rugose at
tim 8: Р.

Шы, ыы IT RI SCIL ОРИОН Acericola
. Spores 12-17 и long, decidedly rough-walled (fig. 58)..................... 38
Spores 9-12 и long, smooth or nearly во......................... . Р. aurea
Spores 6-9 и long, decidedly os trinis “tt {S| | ENTER 19. к а
Los. 6-0 к long, entirely ооб... осор E a 39
Со $6 plong, с ПАО n e . P. duroides

. Pileus dry or moist, in youth showing fine whitish fibrils Aii a lens;
ос АИ ОС ОКИ ОЛЛО ERE Md 16. P. caperata
Pileus slimy-viscid, енна чре ини ОТ 17. P. Мститрћуі
. Pileus very slightly sq ulose Pod ЛР 24. Р. Johnsoniana
Pileus uniformly squamulose or пеа arl РЕТТІ т 40

. Cystidia present, -— abundant, төне fusoid, projecting sather
46.

о s cdi lo Кыр eae cs Gavaskar ++» ее» Р. Schraderi
Cystidia absent or ROR by small inconspicuous brown bodies in

. Scales of the pileus rather large, scattered; pileus yellow or buff in color

ЖУРОККО ОО ОК гы N аы 45. P. rigidipes

Scales of the pileus —— and dot-like (pl. 23), abundant; pileus brown
Go ND очная 54. P. angustipes
43

. Pileus glabrous or saint OKOE ASEPE et rb bb aed

Pileus more or less distinctly fibrillose or всаіу........................... 48

. Pileus watery-brown, cinnamon, yellowish, or pale tan, or drying some-

what brighter, hygrophanous; cystidia present, sometimes quite rare...... 44
Pileus with more red in its coloration, ochraceous-buff to reddish-yellow
г tawny, concolorous or darker in dried plants; not hygrophanous;

о NN. зы АЗАПТЫ ымды ИЕ Cube pus 46

. Spores truncate at one end (figs. 10, 14), smooth; cystidia 15-20 и in di-

ameter in the broadest part, not with an elongate tip (figs. 9, A
МЕСЕ Vy rain CARRERA LAE ҰЛЫ ESL ice qiia vac Ya мА ТИЕ 8. P. Acericola
Spores not truncate, smooth; cystidia 6-10 шіп diameter in the ое

part, not with an elongate tip (fig. 98)...................... . P. Aegerita
Spores not truncate, rough-walled at least at maturity; cystidia pre и

. Spores rough-walled ыра 103); gills not interveined; plants known only
83. P.

co Қа ИМЕК: CS PETERE cerasina
p th; gills strongly interveined; plants known only from Oregon
О П ае оте Ы 82. Р. oregonense

1927

ГА
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—

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t2

сл
с

л
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сл
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e

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о

]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 103

Бро EL. o 4 2. MERPPCKIETEREEEETOOIETOTET 0o 005 T LU 49
as El 6 x19. РНЕ Ue T roo Pus Vel ec 50
Брого ЭДО Терин. г... нел 89. P. albocrenulata

. Pileus lemon-yellow, zinc-orange, or tawny, with aio mtd
brillose, sub-appressed scales; stem 2-5 mm. thick. ........ P. flammans

Pileus pale cinnamon or cinnamon-buff, with eatnclorodp сы nted
scales (pl. 22); stem 5-12 mm. thick. ................ . P. squarrosoides

. Gills bright-colored, i.e., Ае bright ferruginous, or овога кове,
ИОС ПИН 4 5 4. . y ааг... ds ce ee EE 51
Gills cinnamon to dark ры tawny, or brown 44. See a Vc ИКЕ 53

. Pileus with greenish ti t an early stage = 18);
spores slightly roughened (fig. 1 13) аан eT Б ERN 37. aeruginosa

Pileus pale Ra frequently slightly areolate at the "ш врогев
ЖЕУ НАК OO Е узу oko ec eens cds ЖЫ . P. luteofolia
Pileus apricot-yellow to zinc-orange or tawny, not at all dil BOR Hale shee 52

. Spores rough (fig. 109); pileus 4-15 cm. broad; stem 0.5-3 cm. thick

Spores smooth; pileus 2-5 ст. broad; stem 2-5 mm. thick...... 63.
. Plants with one of the following peer
(a) Spores decidedly rough-walled (fig. 109).............. . spectabilis
(b) gm uniformly covered with minute dot-like ais ‘ol 23)
arcely visible except under a lens............... P. angustipes
(c) Pileus with large white floccose patches or soft white э
о DN de 56. P. destruens
Plants with none of the above сһагасфетв................................ 54
"€. t T T ....:....... QU) aE a e E A A A S 55
Meu ҮШ or slimy-vigtid.1,.. 522205 ае een Chae ks 58
о С ой lacking.. Ао с A marte ы. 56
Cystidia at least in part as brown organs among the basidia............... 57
Cystidia ther hyaline, projecting prominently (fig. 1492). .46. P. Schraderi
ў Рйепв 2-6 em. broad, with small scales; stem 1.5-3 cm. long, 1-5 mm
|o rS ВАН ПА noo LT REB "A E. бонды
Pileus 6-12 cm. broad, with а ее fibrous-scaly covering (pl. 13);
stem 5-8 cm. long, 8-10 mm. thick.................. 66. P. fulvo-squamosa

. Pileus with many large duod often floccose scales (pl. 21); veil forming

a thick persistent floccose annulus; stem conspicuously scaly..44. P. squarrosa

Pileus with scattered appressed inconspicuous scales; veil forming a slight
often evanescent annulus; stem only slightly scaly.......... 45. P. rigidipes
59

. Cystidia none; plants less than 5 ст. broad at maturity....................

Cystidia abundant, hyaline, flask-shaped, projecting; pileus 2-7 c

broad. БЕТТ ә- Flammula nass
Cystidia as brown organs between the База (fig. 127a, 135a); mature

pileus 5-15 cm. оғ more in diameter. ...... 2201 ы соле. Лы Сее

. Stem marked with rings of white fibrils; spores 7-9 и long. ..... е P. lucifera

Stem not so marked; spores 6.5-7.5 и long.................. . P. limonella

: ibd 9-11 » long; known only from Colorado, New Mexico اچ‎ Wy-

ООЛО E E 40. P. aurivelloides
61

1 See footnote, bottom p. 101.

[Уоь. 14
104 ANNALS OF THE MISSOURI BOTANICAL GARDEN

61. Pileus with large appressed, spot-like scales; stem increasingly scaly
downward; pileus tawny or rusty in сојог.................. 41. P. aurivella
Pileus with medium-sized scales, often recurved; stem increasingly scaly
upward; pileus yellow to 2іпс-огапре........................ 42. P. adiposa
І. Spores with a truncate apex (fig. 2) (see also II, p. 117).
A. Cystidia ENT. in the hymenium: largest plants more than 4 cm. broad
(see also B, p. 113).
1. Spores 10-14 n long.

1. Pholiota vermiflua Peck, Rept. N. Y. State Mus. 31: 34.
1879. FL &.
Pileus 2-12 ст. broad, hemispheric to campanulate or plane,
white, cream-color or massicot-yellow, retaining these colors in
rying, dry or moist or
in wet weather almost
viscid, glabrous or the
margin fibrillose from
the veil, the center of-
ten Борон reticulate-
areolate at maturity;
flesh rather thick, often
with a slightly disagree-
able taste, pure white;
gills slightly uncinate to
broadly adnate or sin-
uate, close, 3-10 mm.
broad, white then dark
brown, ochraceous-
tawny to buckthorn-
brown in dried plants;
veil white, membranous,
2%. forming a superior, often

of aie type ді mali oina tho mf ated eyetidial evanescent annulus, or
uu ы ДЫ) Teig do ihe желді
from the үр pe specimens, X 550; 4 and 5, more glo- of the pileus; stem cen-
Бове cystidia not projecting, Х 550. tral, equal or more often
enlarged at the apex and tapering to a somewhat bulbous base,
white or light brown, fibrillose or glabrous, sometimes striate
and pruinose at the apex, solid or with a small hollow, 5-12 cm.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 105

long, 3-15 mm. thick; spores ovoid or elliptic, more or less
truncate at the apex, smooth, 10-14 x 6-8 ш; cystidia present,
subglobose to pyriform with а tapering base, 35-45 Х 20 y,
not prominent, sometimes quite rare.

Habitat: on the ground in cultivated, grassy, or waste places;
not in dense woods.

Distribution: specimens have been examined from Cambridge,
Mass.; Ticonderoga, Rochester, Menands, Bronx Park, and
Searsdale, N. Y.; State College, Pa.; Chapel Hill, N. C.; West
Elkton, Ohio; Greeneastle, Ind.; Lexington, Ky.; St. Louis, Mo.;
Rooks Co., Kan.; Denver, Colo.; Berkeley and San Francisco,
Cal.

Illustration: Harper, Trans. Wis. Acad. Sci. 17: pl. 28 С-Е, 29;
18: pl. 11-13 (as P. dura).

Closely related to P. praecox with which it has often been con-
fused. Тһе larger spores will invariably separate it from that
species. Тһе thin, slender-stemmed, Panaeolus-like form with а
rich brown color to the gills grades over gradually into the thick,
heavy, fleshy form with areolate pileus. "This species is appar-
ently P. dura Bolt. of Europe, often reported from this country,
but the final evidence that the two are the same is yet lacking.
If they are identical then all the earlier descriptions and spore
measurements reported for P. dura are erroneous, and based on
the fleshy forms of P. praecox. Bresadola gives the clue to the
situation in describing P. dura xanthophylla, in which he reports
the spores as 10-12 џ long, and indicates that in the true P. dura
they are the same, although all earlier records report them as ap-
proximately 8-10 y. long; Ricken and Lange, in their recent work
on the Agaricaceae, also give a difference in spore measurements
as indicated above. From these facts I conclude that in all
probability the plants described here as P. vermiflua represent the
true P. dura, and the species 1s distinct in the larger spores. In
P. praecox they rarely reach a length of more than 10 y.

Hebeloma hortense Burt, recently described from Missouri,
probably belongs here as an evelate form. Cooke’s illustration
of P. dura represents that species quite well except for the veil.
See further under P. ргаесох.

2. Spores 8-10.5 u long.

(Vou. 14
106 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. Pholiota praecox (Pers.) Fries, Syst. Мус. 1: 282. 1821.
Agaricus praecox Pers. Comm. Fung. Bavar. 89. 1800. РІ.9.
Pileus 2-14 ст. broad, convex to campanulate or nearly plane,
sometimes umbonate, often fuscous or fuscous-black when very
young, soon whitish or cream-color, often tinged with yellow or
tan, or the center brownish, or in wet weather uniformly tan,
usually ochraceous or tan
in herbarium specimens,
hygrophanous in wet
weather, soft, glabrous, or
at times as though finely
tomentose, dry or moist,
areolate in dry weather or
in large specimens; flesh
white, odor and taste
strongly farinaceous; gills
sinuate-adnate to broadly
adnate or with a very slight
аесот: 6, section of the hy- decurrent tooth, medium-
RPM T nc’ Goa а) аай дабыл close, whitish, becoming
from the hymenium, X 550; 5, у epee, х 1200. brown ог rusty-brown,
АП from Orakels] Hab. 331 honey-yell ow to clay-color
ог snuff-brown in herbarium specimens, 8-12 mm. broad; veil
membranous, forming a white, superior, persistent or evanescent
annulus, or partially adhering to the margin of the pileus; stem
central, equal or somewhat bulbous below, whitish, pruinose-
mealy to slightly fibrillose, squamose, furfuraceous, or becoming
nearly glabrous, often striate above the annulus, stuffed or hollow,
3-15 ст. long, 3-20 mm. thick; spores ovoid, usually with a trun-
cate apex, smooth, deep brown, 8.5-10 x 4.5-6 u; cystidia pres-
ent, flask-shaped or ventricose, sometimes rare, projecting some-
what, 14-18 y. in diameter.

Habitat: on grassy ground, in lawns, fields, etc.; sometimes on
the ground in open woods, or on straw or other litter carried into
woods.

Distribution: specimens have been examined from Boston,
Mass. ; Washington, D. C.; Bronx Park, Whitestone, Lake Placid,
and Syracuse, N. Y.; Philadelphia Co., and State College, Pa.;

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 107

West Elkton, Ohio; Tolland, Lake Eldora, and South Boulder
Cafion, Colo.; Berkeley, Santa Barbara, and Del Monte, Cal.;
Corvallis, Ore.; Seattle, Wash.; Banff, Alberta, Canada; Yakutak
Bay, Aqua Dulce River, and Muir Glacier, Alaska. Also re-
ported from Illinois by Harper, and from Michigan by Kauffman.

Illustrations: Atkinson, Mushrooms, pl. 42; ed. 2, pl. 46; Berke-
ley, Outl. Brit. Fung., pl. 8, f. 1; Bresadola, Fung. Mang. pl. 49;
Cooke, Ill. Brit. Fung. pl. (360) 381; Gillet, Champ. Fr. pl. (292)
524; Hard, Mushrooms, f. 209; Harper, Trans. Wis. Acad. Sci.
17: pl. 27, 28 A, B; Kauffman, Agar. Mich. pl. 59; Murrill,
Mycologia 3: pl. 49, f. 1; Patouillard, Tab. Anal. Fung. f. 112;
Peck, Rept. N. Y. State Bot. 49: pl. 46; Peck, N. Y. State Mus.
Mem. 3: pl. 57.

This is undoubtedly our most common Pholiota and undoubt-
edly grows in every state in the Union. I find it to be abundant
in May and June in Pennsylvania, and it was just as common in
Colorado in June and July at elevations ranging up to nearly
10,000 feet. 'The annulus formed is usually ample and mem-
branous, but rather fragile, and apt to disappear early, or may all
but fail to develop, and may hang to the margin of the pileus
instead. Most writers have remarked of the close relationship
of this species to P. dura, a species that does not occur in America
unless represented by P. vermiflua (which see). Тһе spores are
distinctly smaller than in that species, however. Kauffman's
record (9-13 х 6-7 u) probably includes spores from both species,
while Harper's record of P. dura is undoubtedly this species.

At different times reports have reached the writer of the ex-
istence of an evelate and exannulate form of this species or of
P. dura. These are the plants recently described by Dr. Burt
from St. Louis, Mo., as Hebeloma hortense n. sp. Тһе writer
collected this plant in abundance in the Missouri Botanical
Garden as early as April, 1914. Хо sign of a veil is to be found
and the plant has much of the characteristic habit of the short-
stemmed form of P. praecox or of P. vermiflua. The pileus may
become considerably areolate in dry weather or with age, and the
spores give the larger measurements of those of P. vermiflua, and
have the same characteristic ovoid shape with a truncate apex.
Cystidia similar to those in P. vermiflua have been noted in sec-

[Vor. 14
108 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tions of these plants. It seems quite likely that Hebeloma hor-
lense is а recent variation from P. vermiflua (= P. dura), and
our imperfect system of Agaric classification necessitates its ref-
erence to the former genus.

A sterile form of P. praecox is described by Murrill from the
vicinity of New York City.

3. Pholiota Acericola Peck, Bul. Buffalo Soc. Nat. Sci. 1: 50.
1873. P1 10.

Pileus 2.5—7.5 em. broad, broadly convex or nearly plane, rarely
somewhat umbo-
nate, cream-buff to
clay-color or buck-
thorn-brown when
Y' young and moist,
buff-yellow or Na-
ples-yellow when
old, the center fre-
quently somewhat
darker than the
margin, not strong-
ly hygrophanous
but changing color
from young to old
condition, glab-
rous, clay-color to
tawny in herbar-
ium specimens,
often rugosely re-
tieulated or сог-
rugated; margin
even, sometimes

Figs. 9-14. NJ Acericola: 9, section of the hymenium
showing the peculiar "e: x four-poi nted cystidia Ж and up turned; fles
b), and the spores (е), t spores, Over-
holts d ь, 86); ш 11, “кейеп СА. i ow showing thin, white, taste

ask-shaped cystidia (a verholts Herb. 984); 12, iso- E
cee EA x 550 (Overholts Herb. 3953); 13, spores, and odor farina
Ov ree Herb. 3765); 14, spores, X 1200 (Over- ceous; gills sinuate-

holts Herb. 3 adnate or with a

small decurrent tooth, medium-close, 2.5-7 mm. broad, grayish,

1927] 1 !
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 109

becoming brownish-ferruginous or at some stages with a purplish
cast, the edge floccose-crenulate; veil forming a large, persistent,
or rarely evanescent, membranous, superior or in some instances
nearly median, deflexed, white annulus, sometimes of a rich
brown color and striate on the upper side; stem central, equal ог
thickened at the base, fibrillose-striate to nearly glabrous, white
or whitish, stuffed or hollow, typically with more or less of a
white tomentum and strings of mycelium at the base, 6-11 cm.
long, 4-15 mm. thick; spores ovoid or ovoid-elliptic, with a trun-
cate apex, smooth, dull brown, 8.5-10.5 х 5-6 u; cystidia present
but not abundant, flask-shaped or broadly fusoid but only the
tips projecting, rather conspicuous, the tips rarely two- to three-
forked, 15-20 y. in diameter below.

Habitat: mossy rotted trunks of deciduous trees, perhaps also
on coniferous wood; rarely on the ground around rotting logs or
on leaf mold.

Distribution: specimens have been examined from Bar Harbor,
Me.; Redding, Conn.; Lake Placid, Arkville, Bronx Park, Sand
Lake, Raybrook, Van Cortlandt Park, and East Worchester, N.
Y.; Falls Church, Mountain Lake, Crabbottom, and Blacksburg,
Va.; Pink Bed Valley, N. C.; Auburn, Ala.; State College and
Gray's Run, Lycoming Co., Pa.; Oxford, Ohio; St. Louis, Mo.;
Tolland, Colo.; Santa Cruz, Cal.; also reported from Michigan
by Kauffman.

This is one of our most common species and as here admitted is
found to range from the Atlantic to the Pacific. Тһе distin-
guishing characters are the ochraceous color, the slender habit,
the ample persistent membranous annulus, the truncate spores,
and the white mycelial strands at the base of the stem. Peck’s
original collection was from old maple logs, and I have tried to
limit the species to а woody substratum but it is impossible. I
would like to separate P. sphaleromorpha as a distinct geophilous
species as based on European specimens at New York, but it is
manifestly impossible, at least with our present knowledge of the
plants. Bulliard's original illustration fits our plant remarkably
well. Patouillard (Tab. Anal. Fung. pl. 645) shows а rosy or
purplish tint to the gills of that species very comparable to the
color of the gills in our specimens. А collection from Colorado

[Vor. 14
110 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in my own herbarium (1866) is remarkably similar to authentic
European interpretations of P. sphaleromorpha, but it grew on а
rotten log and can as well be referred to P. Acericola, although the
spores are а trifle broader and only inconspicuously truncate.
Microscopically these plants all agree in their characteristic ovoid
spores with truncate apex, and in the cystidia. In some speci-
mens I find cystidia of peculiar shape with two or three or four
cylindrical projections at the tips, so that except for the lack of
vertical walls in the enlarged base the entire organ is almost an
exact picture of the longitudinally septate basidium of a Tremella
(fig. 9a). This throws an interesting side-light on the origin of
cystidia, as these four projections are undoubtedly a prolifie de-
velopment of the usual four sterigmata of the basidium.

Numerous collections were found at New York to be referred
to the genus Stropharia, as the slight purplish tint of the gills
might indicate. "The habit, especially where the plants are grow-
ing on rich humus, is very much like that of Agaricus silvicola
or А. placomyces. Specimens growing on the ground are likely
to be mistaken for P. praecox which is not usually a woodland
plant. Тһе spores are sometimes slightly wedge-shaped at the
base, giving them an appearance of being obscurely 5-angled as
reported by Kauffman. Several other species show this same
character.

In view of its wide distribution Harper's failure to identify this
species as a member of the flora of the Great Lakes Region is
somewhat surprising.

4. Pholiota Howeana Peck, Rept. N. Y. State Mus. 26: 59.
1874.

Stropharia Howeana Peck, Bul. Buffalo Soc. Nat. Sci. 4: 53.
1873.

Pileus 2.5-8 ст. broad, convex then plane, subumbonate, yel-
lowish or yellowish-brown, sometimes lighter and slightly rugulose
when mature, sometimes darker in the center, dry, even, glabrous;
margin even; context fragile, fleshy, bitter to the taste; gills
sinuate-adnate or with a decurrent tooth when young, often en-
tirely separating and leaving a distinct trace on the stem, whitish,
becoming ferruginous-brown, rather close, eroded and lighter-

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 111

colored on the edge; veil forming a conspicuous, persistent, su-
perior annulus; stem central, equal or slightly thickened at
the base, colored similar to the pileus, glabrous, or the base
white-tomentose, hollow, 5-11 cm. long, 4-7 mm. thick, some-
times with white mycelial

cords at the base; spores

ovoid or ovoid-elliptie, trun- x
cate at the apex, smooth, 22 d.
dilute brown, 8-10 X4.5-5.5u;

cystidia present but quite rare,

broadly flask-shaped, project-

ing, hyaline.

Habitat: on the ground in Са а tn a on of
woods and bushy places, often зе gla ee 16 pu All
among grass.

Distribution: reported, in addition to the type locality, Center,
N. Y., from Michigan by Kauffman.

I must admit my inability to distinguish this species among the
several hundred collections I have examined, in both fresh and
dried condition. Harper likens it to P. sphaleromorpha, and
Patouillard shows that species with a rosy or purplish tint to the
gills as would be indicated for Peck's plant by its original in-
clusion in the genus Stropharia. Undoubtedly, if it is a woods
form, and I have met with it, I have referred the specimens to the
terrestrial form of P. Acericola, but unless the color of the gills is
well marked I hardly see how the species can be admitted.
P. sphaleromorpha, as І have remarked in another place, is
extremely similar to the terrestrial form of P. Acericola. Му first
sections of the type specimens of P. Howeana yielded no
cystidia, but I have since found them to be present but quite
rare. When present they are very similar to those in P.
Acericola, but somewhat less narrowed at the neck. Kauffman
reports the species as occurring in open grassy fields, while Peck
says it occurs in woods and bushy places. "The five-angled form
of the spores stressed by Kauffman is not more marked in the
types of P. Howeana than in P. Acericola, but the spores may be
slightly broader in proportion to their length than in that species.
I have so far seen no characteristics that would set it off from

[Vor. 14
112 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. praecox if it grows in grassy open places, nor from the ter-
restrial form of P. Acericola if it grows in deep woods.

5. Pholiota temnophylla (Peck) басс. Syll. Fung. 5: 740. 1887.

Agaricus temnophyllus Peck, Rept. №. Y. State Cab. 23: 90.
1873.

Pileus 2.5-5 em. broad, hemispheric becoming convex, och-
raceous-yellow, cinnamon-buff to ochraceous tawny in dried
plants, smooth, glab-
rous, dry; gills ob-
liquely sinuate-adnate,
medium-close, 4-8 mm.
broad, brownish-ferru-
ginous; veil membran-
ous, white, forming a
distinct, superior, per-
sistent annulus; stem
central, equal, white,

F 7-19. P. temnophylla: 17, кена taken at
абы places along zu vm wing various glabrous, hollow, 5-10
types of ат: (64); 550: 1 18, "брови X 550; ет. long, 4-8 mm.

19, spores, x 1200. АП 5 са "type в ecimens. :
UP ае thick; spores elliptic

to ovoid, one end usually slightly truncate, smooth, 9-10.5 x
6-7 ш; cystidia present but rather rare, flask-shaped or fusoid,
hyaline, 17-21 и in diameter, projecting prominently.

Habitat: grassy ground by roadside.

Distribution: known only from the type locality, Sand Lake,
N. Y., except as reported by Harper from Illinois and by Kauff-
man in Michigan.

Illustration: Harper, Trans. Wis. Acad. Sci. 17: pl. 38, A.

I have been unable to detect this species in my own collecting
grounds or in the herbarium material examined. It is evidently
too close to P. praecox to be recognized from dried plants unless
the broader apex of the cystidia will be an aid. I did not locate
the cystidia in my first sections of the type material but a later
examination showed their presence, but they are not abundant.
Harper's specimens were identified by Peck. "The obliquely and
strongly truncate proximal edge of the gills is stressed as an im-
portant character.

1927
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 113

B. Cystidia absent from the hymenium; largest plants not more than 4 ст. broad.
Spores 6-9 y long.
a. On wood.
See II B 1b, Nos. 29 and 80 (P. mutabilis and P. marginella).
b. On the ground.

6. Pholiota blattaria Fries, Syst. Myc. 1: 246. 1821.

Pileus 1-4 cm. broad, convex
to plane, obtuse or more often
somewhat umbonate, clay-color
to ferruginous or hazel, ochra-
ceous or cinnamon-buff on dry-
ing, hygrophanous, glabrous,
the margin striate or striatulate
when moist, often becoming
somewhat rugose on parting
with the moisture; flesh con-
colorous, odor none, taste mild;
gills rounded behind, soon be-
coming free, 2-4 mm. broad,
close, ventricose, clay-color to
cinnamon; veil forming a sub-
persistent superior or median
annulus, often striate on the
upper side; stem central, equal
or tapering upward, covere
with small white fibrils either
entirely or only at the base, 24, spores of same, X 550; 25-26, various
whitish or slightly brownish, types of spores from same collection, Х550.
hollow, 2.5-5 cm. long, 2-4 mm. thick; spores ovoid or narrow-
ovoid, sometimes with a slightly truncate apex, 6.5-9 X 4—5 u;
cystidia none.

Habitat: on the ground in wooded or grassy places.

Distribution: specimens have been examined from Long Island,
N. Y.; Oxford, Ohio; and St. Louis and Meramec Highlands, Mo.

Most European writers assign spore measurements of 6-8 y. in
length to P. blattaria, and Peck's basis for separating P. rugosa
from this species was on the larger spores that are 8-10 y long
as in P. togularis Bull. (non Fries). "Тһе only specimens of this

[Vor. 14
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN

group with the shorter spores that I have seen are those collected
on Long Island by Dodge and Seaver, a collection by Trelease
and one by myself from Missouri, and one from Ohio.

2. Spores 9-11 и long.

7. Pholiota togularis (Bull) Quel. Champ. Jura Vosges, 92.
1872. Рі, 15.

Agaricus togularis Bull. Herb. Fr. pl. 595, f. 2. 1793.

Pileus 1-4 em. broad, campanulate or convex, becoming nearly
plane, watery-brown when fresh, hygrophanous, becoming och-
raceous on drying, warm-buff
or ochraceous-buff in dried
specimens, the margin striatu-
late when moist, glabrous, even
or perhaps rugose at times; gills
sinuate and narrowly attached,
: medium-close or subdistant,

Figs. 27-30. P. togularis: 27, section Yellow becoming pale ferrugin-
of hymenium showing База, x 550; ous, often ochraceous-buff in
Ali from collection іп New York State dried specimens; veil forming
30, spores, Х 1200, from рт at 8 conspicuous, persistent,
New York from Falls Church, Virginia, median annulus, striate on the
upper side; stem central, equal,
yellow at the top and brownish at the base or entirely brown-
ish, slightly fibrillose, hollow, 2.5-10 em. long, 2-5 mm. thick;
spores narrow-ovoid ог ovoid, {типса{е at the apex, smooth,
9-10.5 X 4-6 u; cystidia none.

Habitat: on the ground in woods or pastures.

Distribution: specimens have been examined from Falls Church,
Va., and Stow, Mass.; also reported by Harper (as P. blattaria).

Illustrations: Bulliard, Herb. Fr. pl. 595, f. 2; Boudier, Ic. Мус.
pl. 101; Cooke, Ш. Brit. Fung. pl. 350 (379); Gillet, Champ. Fr.
pl. 289 (528); Harper, Trans. Wis. Acad. Sci. 17: pl. 59 (as P. blat-
taria).

This species seems to be well separated from P. rugosa by the
larger stature throughout and the ochraceous or buff colors on
drying. Аз here described the species is not P. togularis of Fries
and perhaps Ricken, but compares well with the original il-

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 115

lustration of Bulliard. In this sense both Boudier and Patouillard
have correctly described our plants, the plate given by the former
being particularly good. P. blattaria as illustrated by Harper is
more typical than any of those mentioned, however.

8. Pholiota rugosa Peck, Rept. N. Y. State Mus. 50: 102.
1897. Pl. 15.

Pileus 8-25 mm. broad, broadly conic or campanulate to convex
or plane, sometimes umbonate, yellowish-red or dark ferruginous,
cinnamon or tawny in herbarium specimens, hygrophanous,
glabrous, slightly rugose in the
center; margin striate, often up- que
turned in age; flesh very thin, ( "m
concolorous, odor and taste not ( )
characteristic; gills at first ad- «Ры;
nate, becoming adnexed or free
with age, medium-close, usually
minutely denticulate on the edge,

Figs.31-33. P.rugosa: 31, section of

1-3 mm. broad, yellowish white, i rein showing basidia; 32. spores,
becoming ferruginous or brownish- ссе ибне, м» All from
ferruginous, amber-brown in drie
plants; veil forming a white, persistent, membranous, median
annulus, striate on the upper side; stem central, equal or taper-
ing upward, yellowish above, brownish or blackish-brown below,
finely floccose below the annulus, pruinose or mealy above, hol-
low, 1-2 cm. long, 1.5-4 mm. thick; spores elongate-elliptic,
slightly truncate at one end, smooth, 8-11 x 4-5 ш; cystidia
none.

Habitat: on the ground in open woods; also in greenhouses.

Distribution: specimens have been examined from Adirondack
Mts. and North Elba, N. Y.; State College, Pa.; also reported by
Kauffman from Michigan.

While closely related to P. togularis Bull., the stature is smaller
and the color is dark reddish-brown or féifusinous and remains so
іп herbarium specimens. P. blattaria is distinct in the smaller
spores. I find the plant occurring on the ground in greenhouses.

9. Pholiota mycenoides Fries, Syst. Мус. 1: 246. 1821.

[Vor. 14
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Pileus 0.5-2 ст. broad, at first convex, at maturity plane,
rather thin and membranous, rusty-brown when moist, buck-
thorn-brown to ochraceous-tawny in her-
barium specimens, hygrophanous, drying
first at the center and becoming ochraceous
to deep cream-color, glabrous; margin
widely striate and sometimes white-fibrillose
from the cobwebby veil; flesh concolorous,
odor none, taste mild or sub-farinaceous;
gills adnate or somewhat sinuate, and be-
coming nearly free, sometimes uncinate,
medium-close or slightly distant, rusty-
brown, 2-3 mm. broad; veil forming a con-
os spicuous, nearly median, membranous an-
Pv wd Mia nulus, often striate on the upper side; stem
hymenium showing basi- central, equal or tapering upward, slender,
БЫ); % е M, "x D ox pallid to brown, glabrous or nearly so, hol-
АП trom Е нед» low, 410 qui long, 1-8: ий. thick: pores
— ovoid or broadly ovoid. the apex truncate,
smooth, 9-11.5 x 6-7.5 u; cystidia none.

Habitat :among mosses, especially Sphagnum, in swampy places.

Distribution: specimens have been examined from Stow, Mass.;
also listed by Kauffman from Michigan on the basis of Beards-
lee’s report.

Illustrations: Boudier, Ic. Мус. pl. 102; Cooke, Ш. Brit. Fung.
pl. 503 (405) В

Apparently а very rare species. Тһе very slender plants, with
long stem, narrow pileus, median annulus, and the truncate spores
point to a relationship with the P. togularis group. Тһе habitat
is taken, by European writers, to be a characteristic feature of
the species. We have in this country a plant somewhat similar
to this species, but with a stem that is distinctly peronate with
white fibrils and spores broadly elliptie without sign of truncate
apex.

10. Pholiota filaris (Fries) Peck, Bul. N. Y. State Mus. 122:
144. 1908.
Pholiota togularis filaris Fries, Ic. Hym. 2:2. 1877.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 117

Pileus 0.5-2 ста. broad, campanulate or convex to plane, och-
raceous when fresh, tawny or cinnamon-rufous in dried plants,
glabrous, dry; margin striate, the striae visible in dried plants;
flesh very thin; gills adnate, medium-close, yellow, becoming pale
ferruginous, tawny or cin-
namon-rufous in dried
plants, 1-2 mm. broad;
veil forming а distinct,
persistent, median or su-
к thou gh distant an» Figs. 37-39. P. filaris: 37, section of hymen-
nulus, striate on the up- ium showing раза, X 550; 38, spores, X 550;
per side; stem central, 2) Nome Х 200, All оной at
equal, glabrous or slightly
fibrillose, pallid to brown, hollow, 2-5 cm. long, about 1 mm.
thick; spores elongate-ellipsoid or elongate elliptic, often in-
equilateral, truncate at the apex, smooth, brown, 9-10.5 х
4,5-5 ш; cystidia none.

Habitat: on the ground in woods or pastures.

Distribution: specimens have been examined from Catskill
Mts., М. Y.; Washington (state).

There seems to be room for a species in the P. togularis group
with the following salient features: small size, slender habit, with
thread-like stem, and large spores 9-10.5 u long. However, it
may intergrade too much ‘nto P. togularis, though the habit
seems distinet enough. I have followed Peck in recognizing the
species.

II. Spores not with truncate apex.
A. Growing on the ground or among moss (see also B, p. 132).
1. Plants with large spores 10-15 р long (see also p. 125).
a. Plants 1-5 cm. broad at maturity (see also p. 120).

11. Pholiota platyphylla Kauffman, Papers Mich. Acad. Sci. 1:
145. 1921.

“ Pileus submembranous, 1-3 (4) em. broad, convex, obtuse or
obsoletely subumbonate, hygrophanous, 'tawny' (Ridg.) disk
‘mars brown,’ fading to ‘antimony-yellow’ or ‘warm-buff,’ ob-
scurely striatulate on margin when moist, glabrous; flesh very
thin on the incurved margin, concolor. Gills broadly adnate,
decurrent by tooth, very broad, ventricose, close to almost sub-

[Vor. 14
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN

distant, thin, at the very first pallid, soon ‘clay color’ to *buck-
thorn-brown' (Ridg.). Stem 3-5 (6) em. long, 2-4.5 mm. thick,
equal, or slightly tapering upwards, subflexuous in age, stuffed,
concentrically white-zoned from the delicate veil, zones termin-
ating above middle
of stem in a flaring,
membranous, whitish
annulus, white-scurfy
above annulus, be-
coming at length silky
or glabrous below,
brownish within and
without under the veil
a Бі. rad (a), and зрог 8 (bY X 550; 41, а ci remnants, fuscescent.
cystidium, Х 550; 42, NL Х 1200. One spore
aires e ш faint minute roughness. All from type Spores subellipsoid,
specim but narrower toward
one zin inequilateral in one view, smooth, 10-12 x 5-6.5 y,
pale rusty brown, cystidia scattered on sides of gills, 60-70 x
9-11 o, narrowly lanceolate above the slender pedicel, hyaline;
sterile cells similar but narrower and crowded."

Habitat: on wet moss under pine and spruce along stream.

Distribution: known only from the type locality, Tolland, Colo.

I quote above the original description as given by Kauffman.
I have seen only a fragment of the type collection but it appears
to me to be too close to P. erebia, although in my mounts the
spores run slightly smaller and the cystidia are much less numer-
ous.

12. Pholiota erebia Fries, Syst. Myc. 1: 246. 1821. FL. il.

Pileus 1-5 ст. broad, convex becoming plane or slightly de-
pressed, rarely slightly umbonate, at first brown or blackish-
brown, drying out to umber, then ochraceous-tawny or buck-
thorn-brown, or slightly darker in herbarium plants, somewhat
viscid when moist, glabrous or with a few white fibers on the
margin, even or somewhat reticulate; margin slightly striatulate
at times; gills adnate or slightly decurrent, medium-close or
slightly distant, 2-4 mm. broad, pallid or grayish, becoming
rusty-brown, ochraceous-tawny to cinnamon in dried plants; veil

1927]
OVERHOLTS—THE GENUS PHOLIOTA ІМ THE UNITED STATES 119

forming a thin persistent, white, membranous, superior annulus;
stem central, equal, brownish below, white and pruinose above
the annulus, fibrillose or glabrous below, or at times slightly
squamulose at the

base, solid, 2.5-7 cm.

long, 3-8 mm. thick;

spores elongate-ellip-

tie or elongate-ovoid,

smooth, 11-15 x 5.5-

7.5 ш; cystidia pres-

ent, usually abun-

dant, hyaline, 7-12 в, 24

broad at the enlarged

base, projecting 30- % 45
40 џ.

Habitat: on the Figs. 43-45. Р. erebia: 43, section of hymeniu
ground by roadsides eia y а aiii d (a), and spores (b), Х 550; 44,

i pores, X 1200 (both from type spe n P.
and in woods. орной! 45, isolated cystidia from collection at New

Distribution: spec- York from Bronx Park by Murrill, Х 550.
imens have been examined from Waltham, Mass.; Redding,
Conn.; Bronx Park, West Park, North Greenbush, and Boltons
Landing, N. Y.; Ridgewood, N. J.; St. Louis, Mo.; also reported
from Wisconsin by Harper.

This species grades into P. subnigra, on the one hand, and into
P. ombrophila, on the other, апа cannot be sharply distinguished
from them. In young specimens the pileus is brown or blackish-
brown as shown by Peck's drawings on the type sheet at Albany,
although the dried plants are distinctly cinnamon in color as they
are in most other collections, but occasionally an immature plant
retains the blackish color on crying as have the type specimens of
P. subnigra, a western species, and as have the specimens at New
York collected by Seaver in Bronx Park in 1913. Most collec-
tions that are referable here rather than to P. ombrophila are
smaller plants that have narrow gills that are not decurrent, and
the opposite of these characters is applicable to P. ombrophila.
I am of the opinion, however, that we have one variable species
which should include P. aggericola, P. subnigra Murrill, P. wash-
ingtonensis Murrill, P. ombrophila, and P. erebia. Ricken states

[Vor. 14
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in the body of his text that the last two named are not distinct,
but later he evidently changed his mind and added P. ombrophila
in the Appendix.

13. Pholiota subnigra Murrill, Mycologia 4: 258. 1912.

Pileus 1.3 cm. broad, convex, slightly umbonate, uniformly
fuscous or dusky-drab except оп the margin where а hoary pu-
bescence remains from the
veil, otherwise glabrous,
slightly viscid; gills sinu-
ate-adnate, ventricose,
medium-close, becoming
fulvous, the edge whitish,
2 mm. broad; veil forming
a persistent white super-
| i ior membranous annulus;
е 46507, Р. nies, 48, ain Sf stem central, equal, pallid,
X 550; 47, spores, X 1200. АП тот typespeci- rough with short soft
— whitish conie scales point-
ing upward, solid, 2 em. long, 2.5 mm. thick; spores somewhat
elongate-ellipsoid, smooth, 10-13 x 5-6 u; cystidia abundant,
projecting, conspicuous, flask-shaped.

Habitat: attached to а small buried root.

Distribution: known only from the type locality, Seattle, Wash.

b. Plants 2-15 cm. broad at maturity.
aa. Spores smooth (see also p. 123).

14. Pholiota ombrophila Fries, Hym. Eur. 216. 1874.

Pileus 3-10 em. broad, campanulate-convex to nearly plane,
dull brown, close to wood-brown or verona-brown, russet or light
cinnamon in herbarium specimens, at first with а few white
floccose fibers or scales on the margin, soon glabrous, hygrophan-
ous or subviscid when moist; margin even or faintly striate; flesh
white or somewhat colored, taste mild, odor none; gills usually
decurrent, medium-close or slightly distant, 4-6 mm. broad, clay-
color to rusty-brown, the margin white-crenulate in fresh plants;
veil forming a median or superior, membranous, broad, con-
spicuous, often striate, persistent annulus; stem central, equal or

1927]
OVERHOLTS—THE GENUS PHOLIOTA ІМ THE UNITED STATES 121

enlarged below, whitish or brownish, often dark below, stuffed
or hollow, 4-8 em. long, 4-15 mm. thick; spores cylindric-elliptic
or fusoid-elliptic,
smooth, 11-15 x 5-
7v; cystidia abun-
dant, conspicuous,
projecting 20-40 u,
flask-shaped.

Habitat: in grassy
places in woods or
wooded pastures.

Distribution: spec-
imens have been ex-
amined from Oneida,
Bronx Park, and Lake
Placid, N. Y.; Seattle,
Wash

22
Illustrations: Fries, () ce M

Іс. Hym. pl. 103, f. 2;

PEZ T"
Harper, Trans. Wis. HE SN

Acad. Sci. 17: p l. 31. 48-53. P.ombrophila: =, рт of hymenium
This species repre- Бе cystidia, X 550; 49, sp X 550; 50, spores,
i x 1200 All from specimens e^ sd York from Bresa-
sents the height of dola. 51, section of hymenium showing cystidia,
development in a 550; 52, spores X 550; 53, poni x 1200. АП Ех
type specimens of P. was ashing

т

£

LI
i EN 7

group in which have

been described in addition, P. subnigra Murrill, P. aggericola
Peck, P. indecens Peck, P. washingtonensis Murrill,and P. erebia
Fries. It cannot be sharply separated from either the first or
the last of these and the otaers must be referred to synonymy.
Plants usually retain more of the characteristic umber coloration
of the pileus than in P. erebia, they are often larger, and the
gills are more distinctly decurrent. Sometimes herbarium speci-
mens show the yellowish-brown color usually present in mature
dried plants of P. erebia. If Harper's spore measurements
(8-9 u long) are correct, his plants cannot belong to this species.

15. Pholiota aurea (Mattuschka) Fries, Syst. Мус. 1: 241.
1821. РІ, 12.

[Vor. 14
122 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Agaricus aureus Mattuschka, En. Stirp. Sil. Herbor. 351. 1779.
Pileus fleshy, 10-15 cm. broad, convex to plane, often upturned
іп age, subumbonate, firm, surface dry, pinkish-buff to cinnamon-
buff or ochraceous-buff, or the center more cinnamon, covered
with a fine but distinct concolorous scurf that may separate into
distinct granulose points much as in Lepiota granulosa; margin
even, thin, appendiculate; flesh thick,

о compact, yellowish; gills rounded at
Wana ihe base, rather close, 0.7-2 cm.

broad, ochraceous-buff to ochrace-

INA ous-tawny; veil forming a firm,
large, pendulous annulus, lemon-

color on the upper side, dark buff

Figs. 54-56. Р. aurea: 54, section beneath, eventually disappearing;
Я reda 56. eie, х stem central, tapering upward from
x 1200. All from Overholts Herb. ап enlarged base, below the annulus
No. 8364. :

granular scurfy as the pileus, above
glabrous or nearly so, concolorous or lighter than the pileus,
stuffed or solid, 10-18 em. long, 2-3 ст. thick, inserted 2-5 em.
into the ground; spores ferruginous in mass, golden-brown under
the microscope, narrow ovoid to elongate-ovoid, smooth or per-
haps sparingly warted in age, 9-12 х 4-6 џ; cystidia none.

Habitat: on the ground under bushes.

Distribution: known only from British Columbia.

The specimens on which my record of this species is based are
unique in the character of the covering of pileus and stem, having
somewhat, though not altogether, the appearance at times of the
granulose covering of the stem and pileus of Lepiota granulosa,
as is emphasized in Ricken's description of this species.

In KOH solution or in glycerine the spores have a strong tend-
ency to cohere in twos and fours. In most cases they are un-
doubtedly smooth but I have seen a few that seem to be sparingly
marked with prominent echinulate points. А6 best they are un-
favorable objects for study in the particular specimens I have
examined.

In his emended description of this species Ricken emphasizes
the peculiar covering of pileus and stem and the smooth spores as
characters separating P. aurea from P. spectabilis. Also the
spores are larger and somewhat different in shape.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 123

In preparing the paragraph descriptive of this species I have
relied largely on the excellent notes sent with the specimens by
Mr. W. S. Odell of the Dominion Experimental Farms, Ottawa,
Canada.

bb. Spores rough.

16. Pholiota caperata (Pers.) Fries, Syst. Мус. 1: 241. 1821.
РІ, 13.

Agaricus caperatus Persoon, Syn. Fung. 273. 1801.

Pileus 5-15 em. broad, hemispheric or ovate, then campanulate
or convex to nearly plane, ochraceous-buff to ochraceous-tawny,
sometimes blackish at the center in dried plants, dry or moist,
at first covered with appressed white fibrils that give the pileus
а gray appearance and may be more or less arranged in scales,
finally more glabrous, especially on the margin, and then likely
to be considerably wrinkled or qoran: margin even, at first
incurved; flesh white, taste mi
gills sinuate-adnate or adnate
slightly decurrent, medium-close, A Fa
5-15 mm. broad, at first whitish, ] EXE
then clay-color, finally ferrugin-
ous, becoming cinnamon in dried
plants, usually variegated with
close, narrow, light and dark E 57-58. Р. caperata: 57, section

menium showing asidia and
cross-bands and appearing trans- белен X 550; 58, spores, X 1200. АП
versely wrinkled, the edge floc- {290 Overholts Herb. No. 5241.
cose-crenate; veil forming a persistent, superior or almost
median, white, deflexed, merabranous annulus, sometimes striate
on the upper side; stem central, equal, white or slightly yellowish,
floccose-tomentose above the annulus, glabrous or slightly
fibrillose below, sometimes with a definite annular ring just at
the base—the remains of the universal veil—solid, 6-15 ст. long,
7-25 mm. thick; spores broadly and inequilaterally elliptic,
echinulate, 11-19 x 7-11 u; cystidia none.

Habitat: on the ground in woods.

Distribution: specimens have been examined from Bar Harbor,

e.; Ontario, Canada; Stow, Mass.; Chappaqua, Adirondack
Mts., West Ft. Ann, and Gansevoort, N. Y.; Lamar, Stone Valley,

[Vor. 14
124 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and Reitz Gap, Pa.; also reported from Michigan by Kauffman
and by Harper.

Illustration: Bresadola, Fung. Mang. pl. 48; Fl. Dan. pl. 1675;
Gillet, Champ. Fr. pl. 520 (518); Harper, Trans. Wis. Acad. Sci.
17: pl. 24; Hard, Mushrooms, f. 212; Peck, Rept. N. Y. State Bot.
54: pl. 73, f. 1-6; Ricken, Blátterp. pl. 55, f. 2.

Among the true Pholiotas this species is а characteristic one by
reason of the floccose fibers on the young pileus, the bright color,
the conspicuous transverse banding on the gills of both fresh and
dried plants, and the large, elliptie or lemon-shaped, rough spores.
In most instances the so-called volva at the base of the stem is
not visible in dried plants, and the genus Rozites, to which the
species is often referred, is scarcely tenable. "The nature of this
basal ring is the same here as in certain large species of Coprinus,
and in the light of recent morphological studies is the remnant of
the universal veil in which, to greater or less degree, most fleshy
agarics are at first enclosed. The species represents an inter-
mediate condition, therefore, in regard to veil development,
between those forms in which a definite volva results, and those
іп which the enclosing fibers are too scanty to leave such а trace.

P. Mcmurphyi, described from California, probably represents
the Pacific Coast form of this species.

In age, plants of this species present an often misleading simi-
larity to Hypholoma rugocephalum, in which the pileus colors are
almost identical and the spores strikingly similar.

17. Pholiota Mcmurphyi Murrill, Mycologia 4: 260. 1912.

Pileus 4-8 em. broad, convex to nearly plane, rather thick and
fleshy, greenish-yellow at the margin, orange-cinnamon at the
center, ochraceous-orange to tawny when dry, slimy-viscid,
glabrous; flesh white, without characteristic taste or odor; gills
adnate or slightly sinuate, close, soon brownish, 4-8 mm. broad;
veil forming an inconspicuous, fibrillose, superior, torn annulus;
stem central, equal, yellowish-white, solid, below the annulus
rough with several conspicuous ridges, 4—6 cm. long, 1-2 cm.
thick; spores ovoid or elliptic, quite rough, 13-17 (-21) x 7-
7.5 (-10) в; cystidia none.

Habitat: on the ground among leaves in oak woods.

1927]
OVERHOLTS—THE GENUS ?HOLIOTA IN THE UNITED STATES 125

Distribution: known only from the type locality, Searsville
Lake, Cal.

This species is close to, if not identical with, P. caperata, with
which it agrees microscopically. There is present on the gills of
dried plants to some extent the peculiar appearance of cross-
bands, as characteristic of P. caperata, but they are not nearly so
well marked and indeed are scarcely noticeable in one of the two
mature specimens in the type collection. The peronate stem is
different also, and the annulus is of a more delicate texture and
more evanescent.

2. Plants with small spores ир to 9 и long
a. Spores rough-walled (see also p. 1 26).

18. Pholiota minima Peck, Rept. N. Y. State Mus. 41: 65.
1888.
Pileus 3-8 mm. broad, hemispheric or campanulate, umbonate,

brown when moist, pale buff or yel- OQ
lowish-white when dry, brown in « Kow
herbarium

glabrous, margin striatulate when
moist; gills adnexed, subdistant, fer-
ruginous, cinnamon in dried plants, |
1-2 mm. broad; veil forming ап Figs. 59-61. inima: 50
evanescent, median or superior an- 6548 $50 80" a n ahoni С
nulus; stem central, equal, concolor- spores, x 1200. АШ from type
ous with the pileus, shining, glab- 9Pecmens.
rous, solid, 1-2.5 cm. long, 1-2 mm. thick; spores ovoid or elliptic,
minutely asperulate, 6-8.5 x 3.5-4.5 u; cystidia none.

Habitat: among hair-cap mosses.

Distribution: specimens herve been examined from the Catskill
Mts., N. Y.; Auburn, Ala

Тһе minute roughness of the spores is likely to be unnoticed
under the usual 4-mm. objective, but comes out well under the
oil-immersion lens.

19. Pholiota trachyspora Clements & Clements, Crypt. Form.
Colo. No. 373. 1906.
Pileus 2.5-6 em. broad, convex or plane, deep golden- or coffee-

[Уог. 14
126 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

brown to dark melleous, entirely covered with matted floccose
fibrils of the same color or these

е) &, collected into squamules particu-
eo OSP AS) larly at the center of the pileus,
95%” mÁ ГА
2% j

ЗА агу; gills adnate to adnexed, deep
HOF) brown, medium-close, 3-6 mm. broad ;
98 ^ veil forming a persistent or evanes-
Figs. 62-64. P. trachyspora: 62, cent, superior annulus; stem central,
9 9607 gp eod pow equal, fibrillose, umber-brown, 4-8
spores, X 1200. Allfromtypespec- em. long, 4-10 mm. thick; spores
broadly ovoid or subglobose, dark
under the microscope, decidedly rough-walled, 7-9 x 5-6 в;
cystidia попе.

Habitat: on the ground in woods.

Distribution: known only from the type locality, Sugar Loaf
Park, Colo.

The above description is drawn from the dried specimens as
distributed (1. с.) by Clements. A collection from Tennessee in
the herbarium of the New York Botanical Garden bears strong
resemblance to this species, but is not admitted at present.
The plants are decidedly dark-colored in the dried state. The
subglobose spores are unlike anything else in the genus Pholiota.
A photograph accompanying the exsiccati specimens shows a well-
formed annulus in mature plants. In some respects the plants
resemble species of Cortinarius.

b. Spores smooth.
aa. Plants 1-3 ст. in diameter; from the Pacific Coast only (see also
p. 128). |

20. Pholiota terrestris Overholts, N. Am. Fl. 10: 268. 1924.

Pileus 1-3 em. broad, convex, uniform sayal-brown to cinna-
mon-brown in dried plants, dry, squamulose with appressed,
dark-colored, fibrillose scales or the margin fibrillose only; gills
adnate or slightly decurrent, medium-close, 2-4 mm. broad,
bright cinnamon; veil ample, membranous, not forming a distinct
annulus but sometimes adhering considerably to the margin of
the pileus; stem central, equal, pallid above, brown below, the
apex floccose, distinctly scaly below the sheathing veil remnants

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 127

on the stem, 3-6 ста. long, 2-5 mm. thick; spores oblong-ellipsoid,
smooth, 5.5-6.5 x 3.5-4.5 и; cystidia of several inconspicuous
types, none projecting strongly, some
brown or with а brown mass within.

Habitat: on a lawn.

Distribution: known only from the
type locality, Corvallis, Ore.

'The species approaches P. angustipes
of the East but the spores are somewhat
smaller, the stem more scaly, and the Figs. 65-67. P. terrestris; 5,
cystidia more conspicuous. These lat- section of hymenium showi
ter organs are typically clavate-fusoid, inconspicuous eystidia, X 550;
with a slightly projecting tip and a dis- 66, spores, Х 550; 67, spores, х
tinct ball-like content that has а strong in n. E from type MID
affinity for the eosin stain, so that sec-
tions left in it for half an hour show these sterile structures ex-
tremely well. P. terrigena of Europe is similar, but according
to specimens from Bresadola has spores 8-9 u long and entirely
lacks cystidia. Harper's plants referred to P. terrigena seem to
belong here and he gives the smaller spore record of this species.

21. Pholiota anomala Peck, Bul. Torr. Bot. Club 22: 202.
1895.

Pileus 1.5-2.5 cm. broad, at first hemispheric or sub-conical,
then convex, broccoli-brown when moist, pale yellow or cream-
color when dry, warm-buff in dried
plants, hygrophanous, glabrous; gills
adnate or decurrent, medium-close
or slightly distant, 3-4 mm. broad,
pale, becoming brownish-ferruginous,
ochraceous-orange to cinnamon in
dried plants; veil forming a slight,

Figs finally evanescent, annulus; stem
sectio meni р
narrow cystidium-like body, X 550; central, equal, fibrillose or glabrous,
69, spores, Х 550; whitish or brownish, hollow with

1200. All from co-type specimens Se
from the McClatehie Herb. at New irregular transverse partitions or the
York.

cavities filled with a cottony tomen-
tum, 3-6 cm. long, 2-6 mm. thick; spores oblong-ellipsoid, smooth,

[Vor 14
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN

very dilutely colored, 7.5-10 X 4-5 u; cystidia present, or so rare
as to pass unnoticed, not conspicuous, clavate and attenuate,
sometimes to a long whip-like point that projects 10-15 » beyond
the База, 40-50 х 5-7 y.

Habitat: on sticks and leaves on the ground.

Distribution: known only from Pasadena, Cal.

The type specimens at Albany are in a far better state of pre-
servation than those in a collection from the McClatchie Herba-
rium at New York. The gills in mature plants are strongly
decurrent and the annulus is nearly or quite gone. Sections of
the stem show the peculiar-chambered internal structure in good
shape. Тһе organs described above as cystidia may prove to be
quite different in their origin from true cystidia, and probably
their importance should not be stressed. Тһе species has not
been recognized since its discovery and its affinities are in doubt.

bb. Mature plants more than 3 cm. in diameter.

22. Pholiota albivelata Murrill, Mycologia 4: 260. 1912.

Pileus 2-6 ст. broad, convex to plane, sometimes somewhat
umbonate, isabelline tinted with rose, resembling the color of some
specles of Gomphidius, the
umbo slightly darker, honey-
yellow to light clay-color in
herbarium specimens, slimy-
viscid, glabrous; gills adnate
or slightly sinuate, ventri-
cose, medium-close or slight-
ly subdistant, becoming ful-
vous, the edge white-crenate;
veil forming a large erect or
pendant, superior or median,
oe ie showing different афа of а persistent annulus, pure
0; r^ joe d X 550; ы pores, X white on the lower side, the
rounded body near the > apex х 550. АП upper side brown from the
тош Overbolts — spores and striate from the
gills; stem NIE. equal, usually heavily white-floccose just be-
low the annulus, pruinose or slightly Носсозе above, becoming
subglabrous and rarely yellowish toward the base, solid or hol-

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 129

low, 5-8 em. long, 4-10 mm. thick; spores ovoid to narrow-ellip-
soid, smooth, slightly apiculate at one end, 7.5-9 X 4.5-5.5 u;
cystidia abundant, not projecting conspieuously, small, hyaline
with a rounded conspicuous dark-staining body in the apex,
sometimes pointed, but more often obtuse, 30-35 Х 7.5-9 y.

Habitat: on the ground in woods.

Distribution: specimens have been examined from Seattle,
Wash.; Salem, Newport, and Glen Brook, Ore.

This species is fairly well represented at New York. The heavy
white tomentum on the stem just below the annulus is a marked
character in most specimens. Тһе cystidia with their colored

contents are different from those of any other species, and while
they do not project conspicuously, yet through their abundance
they are not likely to be overlooked.

23. Pholiota duroides Peck, Bul. N. Y. State Mus. 122: 148.
1908.

Pileus 2.5-9 ста. broad, convex becoming nearly plane, dry or
moist but not hygrophanous, creamy-white to ochraceous-buff,
chamois, cinnamon-buff or nearly
ochraceous-orange, retaining these
colors on drying, glabrous or
slightly appressed-squamose ог
with spot-like scales or depres-
sions in the center, margin even;
flesh white, taste mild; gills ad-
nexed or sinuate-adnate, some-
times with a decurrent tooth,
close, narrow, 2-5 mm. broad,
whitish becoming brown or rusty-
brown, with а white-crerulate
edge, cinnamon-buff or snuff-
brown when mature; veil forming
а superior, white, membranous, _ Figs. 74-76. P. duroides: 74, section

, of hymenium showing the еттер
pendant or rolled, subpersistent ous cystidia (a), and nd spores (b x
annulus, often striate on the up- 75, spores, , X m type

й specimens. 76, pean ne of pe
per side; stem central, equal or in 75, > Х 550. From Отко Нал,
о. 7479.

enlarged below, glabrous, ог at
times fibrillose-scaly below, whitish, stuffed or hollow, 3-10 cm.

[Vor. 14
130 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

long, 4-15 mm. thick; spores ellipsoid or ovoid, smooth, 4-6 x
3-4.5 u; eystidia present, obtuse, not projecting strongly yet
rather conspicuous in point of size, often mucronate-tipped,
8-11 шіп diameter.

Habitat: on the ground in waste places, especially in open
woods.

Distribution: specimens have been examined from Stony Brook
and Easton, Mass.; Redding, Conn.; Mount Vernon, Bronx Park,
and Syracuse, N. Y.; Delaware Water Gap and Stone Valley, Pa.;
White Post, Mountain Lake, and Blacksburg, Va.; Unaka Springs,
Tenn.; St. Louis, Mo.

The species is well represented in the herbarium of the New
York Botanieal Garden. Many of the collections had been re-
ferred to the genus Stropharia, due possibly to the peculiar color
of the gills. In fact the plant scarcely has the appearance of a
Pholiota, and dried specimens sometimes resemble thin dried
plants of Russula foetens or В. pectinata in color, though some are
distinctly colored, and in one collection from Blacksburg, Va. the
color is such a distinct reddish-tan that it is close to ochraceous-
orange of Ridgway. Тһе annulus varies considerably but is
typically pendant as compared with the cottony roll of the an-
nulus of P. Johnsoniana.

Considerable confusion has resulted from attempts, initiated
by Peck, to draw a parallel between this species and P. dura, to
which it is not closely related. "The very narrow gills, between
buffy-brown and Dresden-brown in color, and the small spores
make this a distinct species. It may be advisable to separate
from it the darker-colored forms with pileus with a spotted ap-
pearance, but that course seems scarcely justified at present.
Harper describes (Trans. Wis. Acad. Sci. 17: 476. pl. 25. 1912)
unnamed collections of what are in all probability this species
from Wisconsin.

24. Pholiota Johnsoniana (Peck) Atkinson, Mushrooms, 153.
1900.

Agaricus Johnsonianus Peck, Rept. N. Y. State Cab. 23: 98.
872.

Pileus 3-10 ст. broad, convex or nearly plane, thick at the

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 131

center and thin on the margin, yellowish or ochraceous, or yellow
in the center and white on the margin, more or less cinnamon-
buff in dried specimens, glabrous or at times with small ap-
pressed squamules in
the center, dry; mar-
gin thin and some-
times striatulate;
flesh white, flavor
agreeable; gills ad-
nate or sinuate-ad-
nate, close, 2-5 mm.
broad, whitish then
rusty-brown, snuff-
brown to bister in
dried specimens; veil 1

forming а thick;* Figs. 77-82. P. Johnsoniana: 77, section of hymen-

жағы AE а ib:
ENS D ус 122

А TS: Қ

эзе

Sa

white, persistent an- showing om e елы um (а), A S. 550; 79,

б врогев, Х 550; врогев, Х 1200. rom type
nulus; stem central, specimeas. 81, section of hymenium with cysts (a),
equal, glabrous, licht: and spores (b), X 550; 82, spores, X 1200. АП from

colored, solid, slightly Overholts Herb. No. 3946.

striate at the top, 7-10 cm. long, 0.8-1.5 em. thick; spores ovoid
or ellipsoid, smooth, 5-8.5 х 3-4 u; cystidia scarcely noteworthy
in some plants and quite conspicuous in others, some imbedded
and blunt, others projecting, usually pointed.

Habitat: on grassy ground in pastures or in leaf mold in woods
in late summer.

Distribution: specimens have been examined from Redding,
Conn.; Knowersville and Bronx Park, N. Y.; State College, Ра.;
also ыраса from North Czrolina by Anson and from Michi-
gan by Kauffman.

Illustrations: Atkinson, Mushrooms, pl. 49 (44), f. 149 (145);
Murrill, Mycologia 7: pl. 193, f. 10; Peck, Rept. N. Y. State
Cab. 23: pl. 3, f. 4-6.

Atkinson and Kauffman stress the turbinate shape of the pileus
as a character for recognizing the species. In the one collection
of fresh plants I have seen this character was not particularly
pronounced, and in the type specimens is not evident in their
present condition although shown in two of the three illustrations

[Vor. 14
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN

published by Peck. Тһе species has some of the aspects of P.
praecoz, but has smaller spores, lacks the cystidia of that plant,
and the annulus persists as a distinct cottony roll on the stem.
The cluster of stellate crystals mentioned by Kauffman as oc-
curring among the basidia are apparent in the type specimens only
on the edge of the gills. They are needle-shaped and arranged in
a stellate manner. The closest relative of this plant is undoubt-
edly P. duroides, and on the basis of the type material at Albany
I have been unable to cite a single distinguishing character.
Microscopically they are closely alike. The spores of P. duroides
are constantly less than 6.5 џ long; those of P. Johnsoniana vary
between 5 and 8.5 y. and are slightly more irregular in shape. Тһе
cystidia of P. duroides, types, are not conspicuous but are present
as slightly projecting mucronate-tipped organs much larger than
the База. In P. Johnsoniana, types, they are perhaps even less
conspicuous. In other collections referable to the one or the
other of these two species they are usually more conspicuous than
in either of these, and readily distinguishable in crushed prepara-
tions of the gills. Some of them do not project and would seem
to be the old post-mature basidia.

As interpreted by recent writers, P. Johnsoniana does not have
quite the appearance of P. duroides. The latter most often has the
habit of a Russula with short and comparatively thick stem, while
P. Johnsoniana has more the appearance of a slender-stemmed
Agaricus such as A. silvicola. The gills of P. duroides are
very narrow, while those of P. Johnsoniana are considerably
broader. The annulus of the latter is a cottony roll on the stem,
while that of P. duroides is membranous and soon largely evan-
escent.

B. Growing on wood.
1. Pileus not ана. scaly (see aW n p. 142).

a. Spo ugh-walled; prominent flask-shaped cystidia present in the
рб (fig. 88) land also b, р. 187).

25. Pholiota marginata (Batsch) Fries, Epicr. Syst. Myc. 169.
1836-38. РІ. 14.

Agaricus marginatus Batsch, Elench. Fung. 65. 1783.

Pileus 1-7(-8) em. broad, convex to plane or slightly depressed,
sometimes somewhat umbilicate, more rarely slightly umbonate,

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 133

argus-brown, cinnamon-brown, or Sudan-brown when moist,
warm-buff to ochraceous-orange when dry, usually of the latter
colors in dried plants, hygrophanous, glabrous; margin usually
striatulate when moist, often extending somewhat beyond the
gills; context fleshy, thin, concolorous with the pileus, taste and
odor farinaceous; gills sin-
uate-adnate to adnate or
slightly decurrent, me-
dium-close to slightly dis-
tant, 2-7 mm. broad, light
yellowish-brown to buck-
thorn-brown; veil forming
a fugacious or subpersist-
ent annulus; stem central,
nearly equal or somewhat
swollen just at the apex,
fibrillose, pruinose at the
top and with a white to-
mentum at the base, con-
colorous or lighter than Figs. 83-80. P. marginata: 83, section of hy-
the pileus, hollow, 2-8 cm. menium showing cystidia (a), апа spores (b),
long, 2-6 (710) mm. thick; spores, x 1200; 8 i кошш M
spores elliptic or ovoid, Ded eur ирки, 5 1200. Allfrom
often smooth when young, EUNT |

with a rough wall when mature, 7-10 x 4.5-6 ш; cystidia present,
but usually not abundant, flask-shaped with а long apex that
projects 15-25 и, hyaline, 50-80 X 10-15 y.

Habitat: on rotting wood, either exposed or buried, of either
deciduous or coniferous trees; often on sawdust.

Distribution: specimens have been examined from Natlick,
Mass.; Van Cortlandt Park, Ithaca, Karner, and North Green-
bush, N. Y.; Auburn, Ala.; State College and Bear Meadows, PA
Oxford, Ohio; Falling Springs, Ill.; Creve Coeur Lake, Wicks,
Meramec Highlands, and St. Louis, Mo.; Minnesota; Vancouver
Island; also reported from Michigan by Kauffman.

Illustrations: Atkinson, Mushrooms, f. 143 (147); Cooke, Ill.
Brit. Fungi, pl. 372 (403); Hard, Mushrooms, f. 215; Harper,
Trans. Wis. Acad. Sci. 17: pl. 64, 55; Ricken, Blatterp. pl. 56, f. 7.

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[Vor. 14
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Under this name I have listed one of our most common species
on old logs in woods from spring to fall. It differs from P. mar-
ginella in the larger, rough spores, and in the presence of cystidia.
P. discolor differs largely only in the viscid pileus and the brighter
colors on drying. I have been at much loss as to how the species
may differ from P. unicolor, as they seem very closely related if
indeed there is a separating character of enough constancy to
enable one to distinguish the one from the other. On the Pacific
coast there is an apparently common plant which dries a bright
color, has a persistent, well-developed, superior membranous an-
nulus, and the spores are less roughened or altogether smooth.
Also, the stem is slender, scarcely more than 1-2 mm. thick.
This plant I am calling P. unicolor. The same type is also found
in the East where these characters seem to be less constant but
more or less distinguishable. In P. marginata the stem is 2-4
mm. or more thick, has an evanescent and fibrillose, rather than
membranous, annulus, and dried plants are more often of a dull
brown color. I feel that the two are, however, hardly worthy of
separate specific rank. This difference is essentially that of
Kauffman, I believe, although he admits plants of somewhat
larger size to P. unicolor.

Agaricus (Pholiota) autumnalis Peck is
а synonym.

26. Pholiota unicolor (Vahl) Fries, Epicr.
Syst. Мус. 170. 1836-38.

Agaricus unicolor Vahl, Fl. Dan. 655: 7.
1792.

Pileus 0.5-2 cm. broad, campanulate
to conic-campanulate or somewhat con-
vex, often sharply umbonate, cinnamon-
buff to ochraceous-buff or ochraceous-
orange, herbarium specimens cinnamon

Figs. 87-88. P. unicolor: to ochraceous-tawny, glabrous, hygro-
87, section of hymenium phanous, margin striate or fluted; flesh
cid prr or dern thin, somewhat farinaceous to the taste;
X 1200. All from specimens gills squarely adnate, often becoming
somewhat free with the expansion of

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 135

the pileus, rather close, sub-triangular in shape, more or less
tawny or ferruginous; veil forming a persistent median or
superior, upright, funnel-shaped annulus; stem central, equal,
yellowish-brown, decidedly floccose-mealy above the annulus,
fibrillose below or the base white-tomentose, hollow, 2.5-5 ст.
long, 2-4 mm. thick; spores ovoid or elliptie, smooth or slightly
rough when mature, 7.5-10 x 5-6 u; cystidia rare, projecting,
flask-shaped, hyaline, pointed.

Habitat: on rotten wood of coniferous or deciduous trees.

Distribution: specimens have been examined from Piscataquis
Co., Me.; Redding, Conn.; 3ronx Park and Palisades, N. Y.;
Auburn, Ala.; Corvallis and Mill City, Ore.; Seattle, Wash.;
Palo Alto, Cal.; also recorded by Kauffman from Michigan.

Illustrations: Bulliard, Herb. Fr. pl. 530, f. 2 (as Ag. xylophilus);
Cooke, Ill. Brit. Fungi, pl. 356 (404), f. b.

The species is closely related to P. marginata, from which the
persistent membranous annu us, the less roughened spores, the
thinner stem, and the bright colors of dried plants may separate
it. See also under P. marginata.

27. Pholiota discolor Peck, Bul. Buffalo Soc. Nat. Sci. 1: 50.

1873. Pl. 15.

Pileus 1-3 (-5) ст. broad, convex then expanded or slightly

depressed, cinnamon-rufous,

bright ochraceous-yellow when

dry, cinnamon-buff, ochraceous-

orange, or cinnamon in dried С) qı

plants, hygrophanous, smooth,

viscid, margin striatulate when

moist, even when dry; gills ad- ЖЕ;

nate or with a decurrent tooth,

medium-close or somewhat dis- 96

tant, 1-8 mm. broad, pallid then _

pale ferruginous, mikado-brown Sacs i ЗВ уана aoa б

or russet in dried plants; veil 550; 90, spores, Х 550; 91, spores X 1200.
3 Е . . All from type specimens.

forming a subpersistent, dis-

tinct annulus; stem central, terete, equal, pallid, fibrillose-striate,

hollow, 2.5-8 em. long, 1.5-5 mm. thick; spores elliptic, rough,

[Vor. 14
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN

dilute brown under the microscope, 7-9 x 4-6 u; cystidia present
but sometimes rare, projecting 20-40 д or more, flask-shaped,
40-70 х 10-12 u.

Habitat: on old logs and on rotten wood.

Distribution: specimens have been examined from Greig and
Gansevoort, N. Y.; Edgewater, N. J.; Washington, D. C.; Auburn
and Montgomery, Ala.; Creve Coeur and St. Louis, Mo.; also
reported from Michigan by Harper and by Kauffman.

Illustrations: Harper, Trans. Wis. Acad. Sci. 17: pl. 61B.

While very closely related to P. marginata it seems possible to
distinguish P. discolor by the viscid pileus and the brighter colors
in both fresh and dried plants. Microscopically, the two species
are very similar, with spores and cystidia in general the same.
Cystidia, however, are quite rare at times and may be difficult to
locate but I have never failed to find them. I have noted a
tendency for the spores of P. discolor to be quite dark in color in
sectioned material mounted in KOH—so much so as to give а de-
cidedly darker color to the edge of the basidial layer as seen under
the microscope. І am convinced this is an important diagnostic
character for this species. Furthermore, this darkening rather
effectually obscures the rough character of the spore wall so that
spores do not appear so rough-walled as
in P. marginata. 'The character of the
viscid pileus is confirmed in the notes of
several collections, particularly at New
York and at St. Louis. Flammula uni-
color is a quite similar species in the South.

28. Pholiota furcata Overholts, N. Am.
Fl. 10: 272. 1924.

Pileus 1-3 ст. broad, convex to plane,
somewhat gibbous at times, reddish-
brown when moist, ochraceous when dry,
ل‎ cinnamon-buff in dried plants, hygro-

Figs. 92-93. P. furcata: phanous, glabrous; margin even, at first
ы: moon cystidium (a), and Іпейгуей; flesh concolorous, odor попе;
A res (b Ұсақ 38, sporos gills slightly decurrent, close, ochraceous

| to dull einnamon, 2-3 mm. broad, con-

OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 137

spicuously forked and connected by veins so as to appear some-
what poroid; veil forming а superior, distinct but somewhat evan-
escent annulus; stem central, equal, dark watery-brown, Носсове-
pruinose above the annulus, white-fibrillose below, solid or spongy,
1.5-2.5 ст. long, 2-3 mm. thick; spores ovoid or ellipsoid, some-
what rough at maturity, 7.5-9.5 X 4.5-6 ш; cystidia present and
fairly abundant, hyaline, flas«-shaped, ending in a long projecting
tip.

Habitat: on old mossy logs.

Distribution: known only from Van Cortlandt Park, N. Y.
City.

This is the only species of the genus that has forked gills. It
may be an anomalous condition that will not be found again, yet
the type specimens appear otherwise to be entirely normal. The
relationships are with the P. marginata-discolor group, as shown
by the slightly roughened spores and the flask-shaped cystidia.

b. Spores smooth (see also c, p. 141).

29. Pholiota marginella Peck, Rept. N. Y. State Mus. 51: 289.
1898.

Pileus 1-4 cm. broad, convex becoming nearly plane, buck-
thorn-brown or yellowish-red when young or moist, whitish or
yellowish-buff when dry, warm-buff or cinnamon-buff in dried
plants, hygrophanous or at times sub-
viscid, glabrous; young margin striatu-
late, slightly silky, with whitish fibrils;
gills sinuate-adnexed or sinus.te-uncinate,
easily separating, medium-close, 1-4 mm.
broad, minutely eroded on the edge,
whitish becoming dark ferruginous; veil Figs.94-95. P.marginella:
forming a slight or well-developed fuga- 92, section of m

d В owing spores (a), X 550;
cious annulus; stem central, equal, fibril- 95, spores, x 1200. All from
lose below, pruinose above the annulus, YP specimens.
stuffed or hollow, whitish or pallid, sometimes with a white
tomentum at the base, 8-10 cm. long, 1-6 mm. thick; spores
ellipsoid or ovoid, smooth, slightly truncate at one end, brown,
6-8 (-9) x 3.5-4.5 u; cystidia none.

Habitat: decaying wood or on sawdust piles.

[Vor. 14
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: specimens have been examined from North
Conway, N. H.; North Elba, Bronx Park, and Staten Island,
N. Y.; Trenton, N. J.; State College, Bear Meadows, and Stone
Valley, Pa.; Oxford, Ohio; Tolland, Colo; Seattle, Wash.; Yaku-
tak Bay, Alaska.

Illustrations: Peck, Rept. N. Y. State Mus. 51: pl. B, f. 12-20.

Though apt to be confused on external appearances with P.
marginala and P. discolor this species is decidedly distinct in the
smaller, smooth spores and in the absence of cystidia. It is
much more closely related to P. mutabilis from which it scarcely
differs except in a lack of scaliness on the stem. А favorite haunt
for it is an old sawdust pile, and most of the collections examined
have been from such a habitat. Оп drying the pileus is typically
much lighter than in P. marginata but not so bright as in P. dis-
color. The color is a hue entirely distinct from either of these,
and by it alone the species can be recognized. Т found it to be
the most abundant species in the vicinity of Tolland, Colorado,
in 1913 and 1914, and at least а dozen collections are preserved
in my herbarium from that locality.

30. Pholiota mutabilis (Schaeff.) Fries, Epicr. Syst. Мус. 169.
1836-38.
Agaricus mutabilis Schaeff. Fung. Bavar.
() С? 4: Ind. 6. 1774
91 Pileus 1.5-3 em. broad, convex to plane,
QS cinnamon when moist,paler when dry,

ochraceous-buff in dried plants, hygro-

© phanous, glabrous; gills adnate or slightly
“ОСУ decurrent, medium-close, 2-4 mm. broad,
Q pallid then cinnamon; veil forming a

white or dark, superior, evanescent or
persistent annulus; stem central, equal,
concolorous with the pileus, decidedly

Bas SS scaly below the ring, pruinose above,
Figs. 96-97. Р. mutabilis: stuffed then hollow, 3-7 cm. long, 3—5
ае air. p pen mm. thick; spores ovoid or elliptic, slightly
Х 1200. All from — truncate at one end, smooth, 6—7.5 «4-5 u;

mens at New York da.
Bresadola. cystidia none.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 139

Habitat: on stumps and logs.

Distribution: specimens have been examined from Cleveland
and Oxford, Ohio; Gunnison River, Colo.

Illustrations: Batsch, Elench. Fung. pl. 38, f. 208; Bresadola,
Fung. Mang. pl. 51; Cooke, Ill. Brit. Fungi, pl. 355 (402); Fries,
Sverig. Atl. Svamp. pl. 47; Lanzi, Fung. Roma, pl. 76, f. 3;
Schaeffer, Ic. Fung. pl. 9.

Apparently the species is much more common in Europe than
in America. With us it is quite rare and was reported by neither
Harper nor Kauffman. Its external appearance would place it
near the P. marginata group, among which it is unique in having
a distinctly scaly stem, and entirely smooth, slightly truncate and
much smaller spores. The sem is scaly with scattered scales up
to the annulus, in which character it is distinct from P. marginella.

31. Pholiota Aegerita (Brig.) Fries, Epicr. Syst. Myc. 164.
1836-38.

Agaricus Aegerita Brig. Funghi Litogr. Napol. pl. 1. 1824.

Pileus 3-13 ст. broad, sub-hemispherie to convex and then
expanded, avellaneous to chamois,
cinnamon in dried plants, dry, glab-
rous, rugulose on the margin when
young; flesh white, firm; gills adnate
or becoming nearly free, sometimes
slightly decurrent in lines on the
stem, close or medium-close, 4-8
mm. broad, dark brown; veil form-
ing а median-superior, persistent,
conspicuous annulus; stem central,
nearly equal, brownish, more or less i
white-fibrillose, solid, 4-15 сти. long,
4-10 mm. thick; spores ovoid or York from Portugal by Bresadola

: and Torrend.

more often elongate-ovoid or elon-
gate-elliptic, smooth, 9-11 x 4.5-6 ш; cystidia present but not
conspicuous, projecting somewhat, hyaline, 6-10 » in diameter,
some pointed at the apex.

Habitat: on recently felled trunks or from wounds in deciduous
trees.

ГУ от. 14
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: Michigan.

Illustrations: Bresadola, Fungi Mang. pl. 50; Briganti, Hist.
Fung. Nap. pl. 32; Cooke, Ill. Brit. Fung. pl. 365, 453.

I have seen no American specimens of this species. It is ге-
ported by Kauffman as occurring on debris in low grounds in
Michigan. It is a wood-inhabiting form, and specimens at New
York from Bresadola and others collected in Europe by Murrill
agree quite well with Cooke's illustration which Kauffman says
represents aberrant specimens. Тһе colors are similar to P.
praecor but а deeper tan and the pileus may become areolate,
particularly іп the variety strobiloidea. It was evidently this
last character that led Professor Atkinson to concur in desig-
nating Harper's plants as this species. It is the only character
that his plants have in common with this form of P. Aegerita,
and the plants are undoubtedly P. aeruginosa Peck, which seems
to be more common westward.

Only occasionally is the truncate apex of the spore visible to
such a marked degree as in P. Acericola, but traces of it are oc-
casionally seen. Тһе cystidia seem to present a well-marked
characteristie, being rather numerous and projecting only slightly,
reminding one of the conditions shown by Dr. Burt for Stereum
ochraceoflavum.

32. Pholiota oregonense Murrill, Mycologia 4: 262. 1912.
Hypodendrum oregonense Murrill, My-
cologia 4: 261. 1912
Pileus apparently 5 cm. or more broad
when mature, convex, obtuse, thick and
fleshy, dry, smooth, glabrous, ochraceous-
buff to ochraceous-tawny and retaining
these colors when dried; margin strongly
incurved; flesh thin, cremeous, with an
еды nutty ог amygdaline taste in
dried plants; gills adnate, medium-distant
т to distant, yellow or yellowish-brown,
gs. 100-101. . oregon- ы к я
ense: 100, к мейе 4 hymen- becoming darker, strongly interveined,
e, the edges irregular; veil forming a su-
(b), X 550; Ay spores, X perior or nearly apical, irregular, yellow-

1200. АП from type speci- ‘
mens. ype spee- ish-white annulus; stem central ог excen-

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 141

trie, terete or compressed, equal or enlarged upward or down-
ward, yellowish above, fulvcus below, with small scattered uni-
colorous subfloccose evanescent scales pointing upward, solid,
6-10 em. long, 8-20 mm. thick; spores ovoid or elliptic, smooth,
7.5-10 x 3-5 u; cystidia none or not noteworthy.

Habitat: on decayed spot in trunk of living willow.

Distribution: known only from the type locality, Glen Brook,
Ore.

In outward aspects this species is very similar to P. spectabilis.
There can be no question, however, that the spores in the types
are absolutely smooth and frequently more pear-shaped than
ovoid. The specimens are not mature, and it is possible but I
think not at all probable that the spores may become rough-
walled at maturity. Тһе glls are distant as the plants stand,
and the roofs of the gill cavities, at least near the stem, are re-
ticulated with rather conspicuous veins. The gills are not so
bright-colored as is usual in P. spectabilis. Further study alone
can decide аз to the status o? the species.

c. Spores rough-walled; no cystidia in the hymenium.

33. Pholiota cerasina Peck, Bul. Buffalo Soc. Nat. Sci. 1: 50.
1873.

Plants cespitose, 5-12 em. broad, convex to plane, cinnamon-
color to tawny, perhaps lighter at times, dry or somewhat hygro-
phanous, glabrous or nearly #0; margin even; context fleshy, rather
thin, bitter to the taste, wish an amygdaline odor that is best
noticed in young plants; gills m
medium-close or slightly distant, з
sinuate to adnate ог slightly de-
current, yellow, becoming сіппа-
mon or ferruginous, finally prui-
nose from the spores, 5-12 mm.
broad; veil present, forming an Figs. 102-103. ыр лы
early evanescent, spore-stained ssim di hymenium with spores p^
annulus; stem central or more е Лоя spores X 1200. All from
often excentrie, equal or enlarged
below, concolorous with the pileus, fibrillose, at least at the apex,
solid or stuffed, 5-15 cm. long, 5-12 mm. thick; spores elliptic

[Vor. 14
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN

to ovoid, slightly roughened, brown, 6-9 x 4.5-5.5 ш; cystidia
none.

Habitat: on dead wood of deciduous trees.

Distribution: specimens have been examined from Bar Harbor,
Me.; Smuggler's Notch, Vt.; Sterling and Mexico, N. Y.

Тһе species is scarcely distinct from P. spectabilis, and I have
admitted it here only on the basis of Peck's contention that the
pileus is hygrophanous—an unusual condition for plants of this
type. It cannot be considered distinct in the amygdaline taste,
as P. spectabilis may be either bitter or amygdaline. I have re-
ferred here but a very few collections that seem to match the
types exactly. Other collections are in Peck’s herbarium so re-
ferred, but they depart from the original conception of the species
in being decidedly fibrillose and I have referred them to P. specta-
bilis. И there is а hygrophanous Pholiota with glabrous pileus
in this section, the species will stand; otherwise not. Peck re-
ported it as extremely rare.

34. Pholiota rubecula Banning, Rept. N. Y. State Mus. 44: 70
(182). 1891.

Pileus 5-6 сіп. broad, convex to plane, reddish-brown, dark
reddish-brown, or blackish in dried specimens, glabrous, dry;
gills adnate or slightly decurrent, 4—6 mm. broad, tawny when
dry, rather close; veil not apparent; stem excentric, equal or
tapering and rooting at the base, floccose above, glabrous below,
solid, 3-6 ст. long, 4-7 mm. thick; spores ovoid or elliptic,
8-9.5 X 4-6 u, rough when mature; cystidia none.

Habitat: old stumps and trees.

Distribution: known only from the type locality, Baltimore, Md.

This species has not been recognized since its original publi-
cation by Peck. The types still exist at Albany. There are
two specimens accompanied by а water-color sketch by Miss
Banning. Тһе sketch shows a pileus orange-cinnamon, ochre-
red, or tawny, and gills and stem similar but lighter. Тһе color
is about that of the pileus of Lactarius lactifluus. The specimens
show no trace of veil or annulus. The rough spores ally it with
the P. spectabilis group, М + which its affinities are іп doubt.

2. Pileus distinctly scaly at matu
a. Spores rough-walled (see Pe % р. 148).

927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 143

35. Pholiota luteofolia Peck, Rept. N. Y. State Mus. 27: 94.
1875.

Pileus 2-6 ст. broad, согуех, very young specimens dark red
or reddish-brown, becoming pinkish-
red or yellowish-red when mature,
ochraceous-buff or ochraceous-tawny
in dried plants, dry, appressed fibril-
lose-squamulose and sometimes areo-
late in the center, fibrillcse on the
margin; context fleshy, thin, typically
lavender in fresh plants, bitter; gills
adnate or uncinate, sometirnes becom-
ing sinuate-adnate, medium-close or
somewhat distant, 3-8 mm. broad,
yellow becoming bright ferruginous,
mostly ochraceous-buff or ochraceous- سڪ‎
orange in dried specimens; veil form- т A өф iy ba dio
ing a slight, fugacious, spore-stained spores (a), X 550; 105, spores,
annulus; stem central or somewhat ex- ferb. No. 3741. 106, More
centrie, equal or enlarged downward, mou, n showing т
concolorous with the pileus, fibrillose, 107, spores, x 1200. Ай from
solid, 3-9 em. long, 3-10 :mm. thick; type specimens.
spores ellipsoid to elliptie, slightly Tough. 6-9.5 x 3.5-4.5 u;
cystidia none or not notewcrthy.

Habitat: dead wood of deciduous trees.

Distribution: specimens have been examined from Forestburgh,
West Ft. Ann, and Bronx Park, N. Y.; West Elkton, Ohio;
Pacific, St. Louis, and Creve Coeur Lake, Mo.; also reported from
Michigan by Kauffman anc. from Illinois by Harper.

Illustrations: Harper, Trans. Wis. Acad. Sci. 17: pl. 48.

This species is most closely related to P. aeruginosa, and further
studies may result in rearranging the dividing line between the
two species, as some of the collections of the former species ex-
amined are without notes concerning salient characters necessary
in the identification of dried plants. P. luteofolia seems always
to lack the green hue of P. aeruginosa and does not become so
rimose-areolate and the scales appear to be more innate. It isa
smaller plant than P. spectcbilis, and the gills are much brighter-

[Vor. 14
144 ANNALS OF THE MISSOURI BOTANICAL GARDEN

colored in dried plants. From P. cerasina it differs in the dis-
tinctly squamulose pileus. The color of mature plants is typic-
ally а pinkish-red of about the color of Lepiota rubrotincta, but
may fade to more pallid or change to yellowish-red in age. Young
specimens are darker in color.

36. Pholiota spectabilis Fries, Elench. Fung. 28. 1828.
РЬ 1617,
Pileus 4-15 em. broad, convex becoming nearly plane, buff-
yellow to apricot-orange or zinc-orange, becoming at times slightly
more brownish (tawny) in dried plants,
© dry or moist, finely silky or in some very
19 young plants practical glabrous at
times, to distinctly fibrillose or rivulose

<> or in mature plants squamulose; margin
even; context yellow, taste bitter or
22 amygdaline; gills adnexed to adnate or
with decurrent teeth or lines, medium-

ИХ close, 3-8 mm. broad, yellow becoming
ю ferruginous, yellow-ochre to ochraceous-

biis 108. section | IT orange or tawny in dried specimens; veil
menium with spores (a), X forming a distinct, superior or apical,
rires type «ҒАН. of Spore-stained, persistent or subpersistent
+ lutea. 110, spores, X annulus, sometimes striate on the upper

w York from Bresadola side; stem central or nearly so, nearly
from Helland. equal to decidedly ventricose or bulbous-
enlarged at the base, yellow or tawny, yellow and floccose above
the annulus, fibrillose or furfuraceous below, 3-15 em. long, 0.5-3
ст. thick, solid; spores elliptic, rough, 7-9 x 4.5-6 ш; cystidia
none.

Habitat: on stumps and trunks of deciduous or rarely coniferous
trees, or growing from buried wood.

Distribution: specimens have been examined from Toronto,
Canada; St. Andrews, New Brunswick; Newfane, Vt.; Pittsfield,
Mass.; Monmouth Co., Paterson, and Forked River, М. J.;
Frederick, Md.; Washington, D. C.; Montgomery and Bir-
mingham, Ala.; Claryville, Mexico, West Albany, Bronx Park,
and Floodwood, N. Y.; St. Louis and Creve Coeur Lake, Mo.;

1927
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 145

Berkeley, Cal.; Corvallis, Огз.; Seattle, Wash.; also reported by
Kauffman from Michigan.

Illustrations: Bernard, Champ. Roch. pl. 55, f. 1; Cooke, Ill.
Brit. Fung. pl. 352 (394); Егев, Ic. Hym. pl. 102; Gillet, Champ.
Fr. pl. 527 (529); Harper, Trans. Wis. Acad. Sci. 17: pl. 44, 50;
Kauffman, Agar. Mich. pl. 61; Murrill, Mycologia 1: pl. 7, f. 4.

As here admitted, the species is characterized by the usually
cespitose habit, the uniform buff-yellow color of pileus and stem,
the appressed and usually rainute fibrils or fibrillose squamules
covering the entire pileus at maturity, the subpersistent, nearly
apical annulus, and the rough spores 7.5-9 p long. I have seen
but a single collection in which the young pileus is practically or
entirely glabrous, and older specimens in the same collection
show the fibrils quite distine;ly. In other collections the squam-
ules are large enough to be readily visible to the unaided eye. In
every collection examined the annulus has persisted in one or
more mature specimens and frequently is well developed. The
color of the gills is fairly bright but not quite so bright as in P.
aeruginosa or Flammula pulchrifolia.

In some collections, involving dried plants, I get a distinctly
amygdaline taste to the context, although Peck records a bitter
taste and Ricken says it is hitter (brennend-bitter) in European
plants. In one young collection of dried plants I have obtained
a distinctly unpleasant bitter taste from the somewhat worm-
eaten context, although the specimens are certainly of the same
species as other collections in which the taste is amygdaline.
Plants of still other collections are practically tasteless. Most
curious of all, in one collection sent from Alabama and described
as having an “intensely bitter" taste in the fresh plants I am
able to detect an extremely pleasant amygdaline taste in the
dried plants.

Nearly always the plants are found around old stumps or in
similar situations where buried wood is present, to which they are
probably always attached, although this point needs investiga-
tion.

That P. lutea Peck is in reality this species I think there can
be no question, after studying the abundant collections at New
York, the European literature, and such European specimens as

[Vor. 14
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN

are available. Аз it usually occurs in this country, P. lutea is
less robust than the European plant. If we take Fries’ illustra-
tion of this plant as typical of the European species, then our
plants mostly differ in not having quite as conspicuous squamules
as there shown, even in our most squamulose specimens. І be-
lieve, however, that there is no other European illustration that
shows so squamulose a pileus as does Fries’. Cooke’s illustration
(pl. 352) approaches it most closely but here the pileus is shown
with appressed rivulose squamules formed by the separation of
the long fibers of the cuticle. This type is represented in America,
at least on the Pacific Coast. Опе collection at New York from
Berkeley, Cal., 1912, shows this condition almost exactly. In
the same category belongs the type collection of P. ventricosa
Earle, also described from California, but here the plants are not
mature enough to illustrate the character as well as in the other
California collection. 16 is also worthy of note that this last col-
lection has stems that are taper-pointed at the base as shown for
example in Gillet (Champ. Fr. pl. 529), although the other col-
lection agrees with Fries’ figure and Cooke’s illustration in being
obtuse or enlarged at the base. АП of these are quite robust and
must be considered to be the best-developed type of P. spectabilis.
But there occurs also, on both our eastern and our western coasts,
plants (the collections show only young specimens) in which the
stem is fully robust enough to be listed as rather typical P. specta-
bilis, with the obtuse or enlarged stem base. These have uni-
formly, however, the pileus surface of our typical eastern P.
lutea. I refer here to specimens at New York from Seattle,
Wash., by Parker in 1891, and particularly to a collection from
Vermont by Burlingham in 1906. From this condition, the trans-
ition to the typical P. lutea is easy, where the stem is scarcely
bulbous at the base, the plants are medium in stature, and the
pileus varies from slightly fibrillose to squamulose.

The question may well be asked, ‘‘Does the typical P. lutea
occur as such in Europe?" The answer must be in the affirm-
ative, on the basis of an illustration of P. spectabilis published by
Bernard in Champ. Rochelle, pl. 55, f. 1, where our plant is quite
well shown. If Peck had based his species on the types of plants
illustrated in this reference and represented by one collection at

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 147

New York from Alabama by Earle, another from Bronx Park by
Murrill in 1911, and two or three others, there would be more
basis for keeping it distinct. For this type represents one ex-
treme of development, the California specimens the other, and
P. lutea as described by Peck is intermediate between the two.
Moreover, specimens of P. spectabilis at New York from Bresadola
show scarcely a trace of fibrils on the pileus and in stature compare
well with our plants. There is a specimen at Albany, from New
Brunswick, on birch, that nearly exactly matches Bresadola's
specimens.

I must conclude, therefore, that P. spectabilis in Europe and
America is about equally variable, ranging from medium-sized
plants with cylindric or enlarged stems and nearly glabrous to
fibrillose-squamulose pileus, to large robust forms with ventricose
stems and pileus most often heavily fibrillose and becoming
rivulose but sometimes only slightly fibrillose.

37. Pholiota aeruginosa Peck, Rept. N. Y. State Mus. 43: 35
(81). 1890. РІ. 18.
Pileus 2-10 сш. broad, convex, greenish becoming tinged
with yellow or brown, drab
to cinnamon or ochraceous-
buff in dried specimens, dry,
at first glabrous, usuallv
soon more or less areolate
with each areola surmounted
by one to three fibrillose
scales, or sometimes com-
pletely squamulose without
areolae; flesh with a green
tinge, yellowish іп dried
plants; gills adnate or sinu- „f
ate-adnate, easily separating, 113, spores, Х 1200. All from "Overholté
Herb. No. 8081. 114, spores, Х 1200 (from
3-7 mm. broad, pale-ochra- type specimens); 115, spores, X 1200 (from
ceous when young, becoming Sigo. s Herb. . 8773, spores probably
ss mature).
ochraceous-orange ог apri-
cot-buff on drying; veil leaving only a slight fibrous, lacerated
annulus, or entirely evanescent; stem central or excentric, equal

Figs. 111-115. P. aeruginosa: Hs section
i о 550;

[Vor. 14
148 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

or nearly so, glabrous or slightly fibrillose, sometimes sulcate-
striate, colored like the pileus, solid, 3-8 ст. long, 4-10 mm.
thick; spores ellipsoid or elliptic, rarely entirely smooth, typically
slightly echinulate when mature, 6-8 x 3.5-4.5 u; cystidia none.

Habitat: on decaying wood, probably mostly of coniferous trees.

Distribution: specimens have been examined from Trexler-
town and Aaronsburg, Pa.; Staten Island, and West Ft. Ann,
N. Y.; Womble, Ark.; Priest River, Idaho; Seattle, Wash.

In Peck’s herbarium this species is represented by the type
specimen from Pennsylvania and a collection from West Ft. Ann. у
New York. Another Pennsylvania collection is at New York
City, by Mellvaine. Dr. Murrill has already referred his Flam-
mula viridans from the Pacific Coast to this species. Тһе species
seems to be more common westward and exhibits more variation,
even in the same collection, as regards scaliness and the areolae
on the pileus. The plants described by Harper on Atkinson's
determination as P. Aegerita belong here, I judge. Pholiota
Aegerita of Europe is entirely dissimilar in every respect except
the areolate pileus. Excellent specimens of that species are at
New York collected by Murrill in Europe.

It is difficult to separate this species in most aspects from Flam-
mula pulchrifolia, and the writer is of the opinion that they may
represent the same plants.

The markings on the spore walls are discernible only under
close scrutiny with the high power, but are very evident under the
oil-immersion lens, in all but one collection I have examined. In
that one they are mostly smooth even with the higher magni-
fication, but a few are distinctly though sparingly roughened (fig.
115).

Pholiota luteofolia is closely related but the present species
seems distinct in the green tints of young plants, the areolate
pileus of mature ones, and in the more fibrillose scales.

b. Spores smooth.
1. Spores 3.5-5.5 џ long (see also 2, p. 149).

38. Pholiota flammans (Batsch) Fries, Syst. Myc. 1: 244.
1821. РЬ 10.
Agaricus flammans Batsch, Elench. Fung. 30. 1783.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 149

Pileus 2-5 (-8) em. broad, convex to plane, sometimes um-
bonate, lemon-yellow or tawny-yellow, zinc-orange or tawny in
dried plants, dry, adorned with yel-
low, superficial, floccose-fibrillose scales
that may in large part disappear with
age; flesh thin, yellow; gills adnate or
very slightly uncinate, medium-close,
2-5 mm. broad, yellow or ferruginous,
snuff-brown in dried plants or young
specimens may retain their yellow
color; veil lemon-yellow, fugacious;
stem central, equal, with yellow, re-
curved, floccose scales, or scarcely more

than densely yellow-floecose up to the
annulus, stuffed or hollow, yellow, 2-7 Ж Бы
сш. long, 2-5 mm. thick; spores ob- us

long, smooth, 3-5.5 х 2-3 д; cystidia Figs. 116-119. P. flammane
abundant, flask-shaped or clavate- 116, section of hymenium show-
fusoid, brown or hyaline, projecting "92098 (0), and pore (p)
slightly, 30-40 x 6-12 y. section of hymenium showing

Habitat: on dead wood cf both de- lied, X 550; 119, isolated
ciduous and coniferous trees. Eder сес EL.

Distribution: specimens have been d т om Overholts Herb.
examined from Lake Placid, Osceola, x
Greig, and Fourth Lake (Herkimer Co.), N. Y.; Glen Brook,
Ore.; also reported from Michigan by Kauffman and by Harper.

Illustrations: Fries, Ic. Hym. pl. 104, f. 1; Cooke, Ill. Brit.
Fungi, pl. 396 (368); Harper, Trans. Wis. Acad. Sci. 17: pl. 41C;
Ricken, Blatterpilze, pl. 55, f. 5.

In one collection at New York, from a birch stump, the col-
lector’s notes say “viscid.”

P. flammans is distinct (rom all related species in the very
small, oblong spores, measuring 3-5 u long. It is indeed a well-
marked species with bright colors and soft floccose scales, and
while generally considered £s occurring only on coniferous wood,
yet a collection at New York is very evidently on the wood of
birch. Тһе species is rare, and I have seen no American speci-
mens more than 4 ст. broad.

2. Spores between 9.5 and 15 и long (see also 3, p. 152).

[Vor. 14
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN

39. Pholiota albocrenulata Peck, Bul. Buffalo Soc. Nat. Sci. 1:
. 1873. РІ. 19.
Pileus 2.5-12 em. broad, conic-campanulate to broadly convex
or nearly plane, sometimes
umbonate, uniformly yellow-
ish-brown to sayal-brown or
tawny, the darker of these
colors in dried plants, quite
viscid, with rather large
scales that are appressed and
sub-gelatinous when wet, but
become suberect, Носсове,
and lighter in color on dry-
ing, easily separable and
sometimes disappearing in
old plants; gills sinuate-ad-
nate or slightly decurrent,
medium-close or in large
specimens rather distant,
3-13 mm. broad, grayish be-
coming ferruginous, the edge
distinctly white-crenulate
and remaining so at least in
part in dried plants; veil
forming a torn fugacious an-
nulus or partly appendicu-
late to the margin of the
pileus; stem central, equal
or somewhat enlarged below,
Figs. 120-120. Р. albocrenulata: 120, Sparingly or abundantly
section of hymenium showing central medulla squamose with fibrillose
X 1200. АП from type specimens. 123, scales up to the annulus,
кердең tpe ium downs the oeeasionel pallid or brown below, white
125, spores, X 1200; 126, section of edge of and furfuraceous at the apex,
dge the crenulate appearance, X 550. АП 5-11 cm. long, 0.5-2.5 cm.
» thick; spores broadly-elliptic

or fusoid-elliptie, smooth, brown, 11-14 x 6-8 ш; cystidia ap-
parent in fresh specimens as occasional clavate projecting hyaline

49

` 1927]
OVERHOLTS—THE GENUS ?HOLIOTA IN THE UNITED STATES 151

bodies with one or rarely two long slender apical points, rather
difficult to locate in sections of dried plants.

Habitat: at base of trees or on prostrate trunks, especially of
sugar-maple; one collection said to have been from a hemlock
stump.

Distribution: specimens have been examined from Hebron,
N. H.; Adirondack Mts. and Lake Pleasant, N. Y.; Cadillac,
Mich.; Unaka Springs, Tenn.; Bear Meadows, Center Co., Pa.

Illustrations: Harper, Trans. Wis. Acad. Sci. 17: pl. 42, 28.

The dark color of the dried pileus recalls P. aurivella and the
general appearance is similar to that of the European P. sub-
squarrosa. From both of these the species is amply distinct in
the large fusoid-elliptic spores. In fact no other similar species
of this section, except P. aurivelloides, has spores at all comparable
in size to these, and in that species they are quite different in
shape. Тһе plants usually occur singly. Тһе white crenulations
on the edges of the gills are due to the presence there of tufts of
radiating clavate cells as shown in my illustration.

In most collections of
this species the plants are
slender and small in stat-
ure, rarely more than 7
em. broad. Professor Har-
per illustrates, and I have
also found, a large form
with pileus 10-12 cm.
broad, but otherwise with
the characteristies of
Peck's types.

40. Pholiota aurivelloides
Overholts, sp. nov.
Pileus 5-8 cm. broad, =

hemispheric ог broadly Te. 1c HN В. жалады: 22; section
n of hymenium showing an imbe cystidium
маре an late to сону, (a), spores (b), and central medulla (c), х 550;
ferruginous to tawny, 1, врогев, X 1200. АП {тота type quisi
: 9, section of hymenium showing im
lig hter colored on the cyitidia (а), and central medulla (b), х 550; 130,
margin, sometimes carob- spores, X А rom specimens іп New
. ave ] Ycrk State Museum, from New Mexico, by
brown in dried plants, Cockerell.

н [ Уот. 14
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN

probably viscid, with а few scattered, spot-like or appressed
scales; flesh yellow, rather thick; gills sinuate-adnate or with а
decurrent tooth, medium-close or slightly distant, 7-12 mm.
broad, whitish then ochraceous-tawny or russet; veil forming а
thin, superior, persistent or somewhat evanescent, floccose or
submembranous annulus; stem central, equal, yellowish or
brownish, more or less scaly, the scales sometimes somewhat
gelatinous, solid, 4-8 сш. long, 5-10 mm. thick; spores oblong-
ellipsoid, smooth, deep-brown, 9-11 X 6 u; cystidia numerous or
rare, brown, not projecting, 25-35 x 6-8 u.

Habitat: on dead trees or from wounds in living Alnus, бай, or
Betula.

Distribution: specimens have been examined from Ohio Creek,
Colo.; Pecos, N. Mex.; Copperton, Wyo.

'The name here used was applied by Peck in his herbarium to
the Ohio Creek collection made by Bartholomew, Aug. 24, 1899,
and here used as the type collection. It seems sufficiently distinct
from P. aurivella in the very broad gills and the larger spores that
are of a much darker color and with a heavier wall. Their shape
is broadly oblong-ellipsoid, while those of P. aurivella are narrowly
oblong-ellipsoid or ellipsoid. Тһе brown sterile organs in the
hymenium are of a somewhat different type also. I have tried
to consider these plants as P. aurivella but they seem quite dif-
ferent from the plants, various as they are, that have passed
under that name.

3. Spores 6-9 u or rarely longer.
aa. Cystidia t and rath as brown or hyaline,

imbedded or projecting organs (see also bb, p. 159).
aaa. Pileus viscid (see also bbb, p. 156).

41. Pholiota aurivella (Batsch) Fries, Syst. Myc. 1:242. 1821.
Pl. 24.

Agaricus aurivellus Batsch, Elench. Fung. f. 115. 1783.
Pileus 4-13 cm. broad, campanulate to convex, often broadly
umbonate, when young more or less uniformly ochraceous-orange
to tawny, when mature becoming more uniformly tawny, at first
covered with large appressed spot-like scales which may in large
part disappear and when wet may become more or less gelatinous,
viscid; flesh yellow; gills sinuate-adnate or adnate, close, dark

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 153

rusty-brown when mature; veil forming a superior, torn, spore-
stained, partly evanes-
cent annulus; stem cen-
tral or excentric, equal
or tapering upward, dry,
yellowish or yellowish-
brown, Носсове above
the annulus, fibrillose
below and increasingly
scaly or shreddy down-
wards with fibrillose 4 ВС,
scales that may become Lone! lac EE 202 |
recurved, solid, 5-8 ст. № АНА ==
long, 5-15 mm. thick;
spores exactly and con- Figs. 131-133. P. aurivella: 131, section of
stantly oblong-ellipsoid, hymenium showing the imbedded cystidia (а),
smooth, 7-9.5 X 4-5 р; ора of hymentum from anothor gill showing
eystidia present, often pedis at New York from Colorado, b
rather rare, brown, Bethel 133, spores, X 1200, from specimen at
sometimes projecting New York from Italy by Bresadola.

and rather conspicuously sharp-pointed, sometimes imbedded and
blunt, 6-8 y in diameter.

Habitat: on trunks of living (rarely dead) deciduous or con-
iferous trees.

Distribution: specimens have been examined from Colorado;
Corvallis, Oregon; San Francisco, Cal.; also reported by Harper
from River Forest, Ill.

Illustrations: Batsch, Elench. Fung. pl. 22, f. 114, 115; Cooke,
Ill. Brit. Fungi, pl. 390 (351); Harper, Trans. Wis. Acad. Sci. 17:
pl. 38, 89.

For the distinguishing characters of this species as contrasted
with P. adiposa see the notes under the latter. The sterile organs
in the hymenium of plants referred to this species present wide
variations as will be seen from the drawings. In specimens at
New York from Bresadola there is an occasional lance-shaped
colored organ with much resemblance to the setae of some fungi.
In but one American colleczion have I seen these present—that
collected by Gilbert in Oregon, on willow, in 1915. "The col-

pO QE
Ame

жалын

(ШІП!
RA

NS
А?
^

(>
b VS
IA Ge,
ALE,

№

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D

KS
UNS
А,

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ыты

E

Хе.

[Уоь. 14
154 ANNALS OF THE MISSOURI BOTANICAL GARDEN
lection is preserved at New York. Неге these seta-like bodies
are quite numerous and conspicuous as shown in the illustration.
Other collections show a more inconspicuous type of sterile organ
and always entirely imbedded. Such collections may represent
the type of another species but can scarcely be P. adiposa, the
most closely related species. Тһе species as here described is
admittedly unsatisfactory and consists of collections somewhat
similar to P. adiposa but yet certainly distinct from it and the
next species. Harper's photograph is of particular excellence.

42. Pholiota adiposa Fries, Syst. Myc. 1: 242. 1821. РІ. 20, 21.
| Pileus 3-16 ст. broad, hemispheric

® Eo to convex or plane, antimony-yellow
Sr m=O to zine-orange or finally the center
Cy ES somewhat tawny, decorated with
rather medium-sized squamules of а

darker color than the rest of the
pileus, and drying down to small
dark spots on а tawny base, large
thick specimens sometimes areolate
in dry weather, in wet weather the
pileus occasionally glabrous, all colors
well retained in dried plants, viscid
or glutinous or dry in dry weather;
flesh thin or rather thick, white or
light yellow, taste not marked; gills
adnate ог sinuate-adnate, rather
close, 4-10 mm. broad, grayish-
brown then yellow or rusty-brown,
honey-yellow to tawny in dried
specimens; veil yellow, forming a
slight, floccose, evanescent annulus;
stem central or excentric, terete,

бү?

Б
л an sim um
Зы =

Figs. 135-138. P. peer 135,
section of hymenium showi the

imbedded cystidia (a), spores (b),
and central er (с), Х 550; 136,
spores, X 1200. All from Overholts
No. 6191. x Y e s
spores (b), and pens Dg medulla (c)
X 550; 138, spores, x 1200. All
с Overholts Herb. No. 6042

equal or nearly so, viscid (dry in dry
weather), yellow or tawny, with few
or many erect or somewhat recurved
yellow or tawny scales, or sometimes
only fibrillose, solid or stuffed, rarely

1927] à
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 155

with a small hollow, 4-12 cra. long, 5-20 mm. thick; spores ellip-
soid or oblong-ellipsoid, smooth, 7-9(-11) x 4-5 u; cystidia pres-
ent, not conspicuous, projecting only slightly if at all, dark brown,
clavate, 20-42 x 5-10 y.

Habitat: stumps ad АЕ Е of deciduous trees.

Distribution: specimens have been examined from Cambridge,
Mass.; Lake Placid, Bronx Park, and Wells, N. Y.; Forked River,
N. J.; State College, Musser Gap, Bear Meadows, and Westport,
Ра.; Greencastle and Putnam Co., Ind.; Edgmont and Falling
Springs, Ill.; Oxford, Ohio; St. Louis, Mo.; Fort Dodge, Iowa;
Plumas Co., Cal.; and Seattle, Wash.

Illustrations: Atkinson, Mushrooms, pl. 47 (43); Berkeley, Outl.
Brit. Fungi, pl. 8, f. 2; Cooke, Ill. Brit. Fungi, pl. 395 (853);
Hard, Mushrooms, f. 211; Mycologia 1: pl. 7, f. 1-2. 1909;
Peck, Rept. N. Y. State Bot. 49: pl. 46.

The pileus in both fresh and dried condition is typically golden-
yellow or lemon-yellow as contrasted with the prevailing tawny
or rusty color of P. aurivella, its closest relative. Тһе latter
species has larger and more spot-like scales and the stem is short
and thick and increasingly scaly
downward, while in P. adiposa
the stem is equally scaly up to
the ring or increasingly from the
base upward. Ricken states that
the stem of P. aurivella is not
viscid as it is in P. adiposa.

43. Pholiota squarrosoides Peck,
Rept. N. Y. State Mus. 31: 33.
1879. PK 22.

Pileus 2.5-10 ст. broad, sub-
globose to convex or nearly plane,
light-colored, typically cinna- i35 "Section of hymenium. showing
mon-buff or cinnamon in dried cystidia (a), and "xen (b), X 550;
plants, viscid, covered with erect, holts Herb. No. 3770. 141, section of
pointed, cinnamon or tawny scales spores (b), x 580; 142, section м
that give color to the pileus, қуам кетім central medulla

А к қ ue, X 550. АП from type вресі-
sometimes disappearing on the mens. ii

[Vor. 14
156 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

margin; flesh white or slightly yellowish, gills sinuate-adnate,
medium-close, 4-7 mm. broad, whitish becoming brownish-fer-
ruginous, cinnamon or ochraceous-tawny in dried plants; veil
forming a floccose, persistent or evanescent annulus; stem central,
equal, brownish and with recurved light cinnamon or tawny scales
below the annulus, white and smooth above, solid or stuffed, 5-15
em. long, 5-12 mm. thick; spores ellipsoid or oblong-ellipsoid,
smooth, 4-6 x 3-4 u; cystidia present, variable, of two general
types: (a) hyaline, pointed at the apex, projecting slightly, and
(b) brown, obtuse or pointed, sometimes projecting, both types
measuring 25-35 x 8-12 y.

Habitat: on stumps and trunks of deciduous trees.

Distribution: specimens have been examined from Maine;
Redding, Conn. ; Vaughns, Catskill Mts., Canandaigua, and North
Elba, N. Y.; Bear Meadows, Pa.; also reported from Michigan by
Harper and by Kauffman.

Illustrations: Hard, Mushrooms, f. 216; Harper, Trans. Wis.
Acad. Sci. 17: pl. 36, 37; Peck, Ann. Rept. N. Y. State Mus. 54:
pl. ?8, f. 6-14; White, Hymen. Conn. pl. 21.

Тһе scales on the pileus appear Носсозе under a lens, but they
are rather harsh to the touch in dried plants. In young plants
they appear more or less erect, particularly over the center of the
pileus, but on the margin they may be only recurved or in dried
plants practically appressed. From P. squarrosa the species
differs in the viscid pileus, the smaller size, the more cespitose
habit, the squarrose center of the more highly scaly pileus, and
the smaller spores. The spore difference seems insignificant as
here stated but when spores of the two species are compared side
by side under the microscope it is very striking. Only rarely do
the spores of P. squarrosoides reach a length of 6 u, the wall is
thin, and the shape more apt to be broadly ellipsoid, while in P.
squarrosa, 6 u is about the minimum length, the spore wall is
heavier or darker, and the shape more oblong-ellipsoid. The
cystidia are very comparable in the two species. Small specimens
that have become tawny in drying have a strong resemblance to
specimens of P. subsquarrosa at New York from Bresadola, and
microscopically the two are quite similar.

bbb. Pileus dry.

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 157

44. Pholiota squarrosa Fries, Syst. Мус. 1:243. 1821. РІ.21.

Pileus 3-10 em. broad, can: panulate to convex or plane, yellow-
ish or yellowish-brown, antimony-yellow to tawny in dried speci-
mens, covered with rather large, recurved, tawny or yellowish
scales, dry; flesh yellowish, taste mild; gills sinuate-adnate and
often somewhat decurrent, medium-close, 3-6 mm. broad, pallid
then ferruginous, in
dried plants varying
from honey-yellow to
tawny-olive or tawny ;
veil forming a thick,
persistent, floccose
annulus often striate
on the upper surface;
stem central, equal,
pallid, yellow or
brown, with conspic-
uous recurved scales `
up to the annulus, л ее
solid, 5-12 em. long, аа 19, өле арои ы N O58 еме
5-12 mm. thick;
spores oblong or ellipsdid! smooth, 6-8 x 3.5—4.5 ш; cystidia
present, variable, of two general types: (a) hyaline, abundant,
pointed at the apex, projecting, and (b) brown, blunt or truncate
at the apex, mostly projecting, both types 25-35 x 7-14 y.

Habitat: on dead trunks ог stumps of various trees, both de-
ciduous and evergreen.

Distribution: specimens have been examined from Piscataquis
Co., Me.; Stocktenridge, Мазз.; Yama Farms and North Elba,
М. Y.; Washington, D. С.; Sate College, Pa.; Jack Brook, Colo.;
also reported from Michigan and Wisconsin by Harper.

Illustrations: Bernard, Champ. Roch. pl. 19, f. 1; Cooke, Ill.
Brit. Fungi, pl. 391 (367); Hard, Mushrooms, f. 217; Низзеу, Ш.
Brit. Мус. 1: pl. 8; Harper, Trans. Wis. Acad. Sci. 17: pl. 35;
Patouillard, Tab. Anal. Fung. f. 340; Peck, Rept. N. Y. State
Mus. 55: pl. 79, f. 1-7.

For detailed data as to the differences between P. squarrosa and
P. squarrosoides, see the latter species.

А [У от. 14
158 ANNALS OF THE MISSOURI BOTANICAL GARDEN

45. Pholiota rigidipes Peck, Bul. N. Y. State Mus. 157: 31.
912.

Pileus 4-8 cm. broad, broadly convex, sometimes slightly ог
broadly umbonate, pale yellow or buff, buff-yellow to ochraceous-
buff or ochraceous-orange in dried plants, squamulose with scat-
tered, appressed, slightly darker-colored, fibrillose scales that are
more prominent in the center though never conspicuous and par-
tially disappear in
mature plants, dry;
flesh white, tinged
yellow next the gills,
distinctly yellow in
dried plants; taste
mild; gills sinuate-
adnate or adnate,
medium-close, 3-7
mm. broad, cinna-
mon or ochraceous-
™ .. tawny and retaining
№... дығы км (а) p "m Sun these colors in dry-
Е medulla (f), Х 550; 148, spores, Х 1200. All ing; veil forming a
from type specimens. ‘

slight, often evanes-
cent annulus; stem central, equal, pallid or yellowish and fibril-
lose-squamulose or becoming nearly glabrous below the annulus,
white and pruinose at the top, stuffed or hollow, 5-9 cm. long,
4—6 mm. thick; spores oblong or oblong-ellipsoid, smooth, 6.5-8.5
х 3.5-4.5 ш; cystidia present, not always abundant, brown, some
projecting, 25-40 x 6-8 y.

Habitat: in woods, probably always on buried or exposed wood.

Distribution: specimens have been examined from Constable-
ville and Utica, N. Y.; Forked River, N. J.; Creve Coeur Lake,

о.

The species is well represented at Albany by the type collection
which shows it to be a medium-sized plant with sparsely scaly
pileus and fibrillose-scaly slender stem of the P. adiposa group.
In spite of Peck's original note that it occurs on the ground among
fallen leaves, I am of the opinion that it must have been attached
to wood, perhaps buried wood. "The aspect is much like that of

a“.
о

927]
OVERHOLTS—THE GENUS FHOLIOTA IN THE UNITED STATES 159

а Flammula. А second collection, perhaps referable here, is
labeled P. terrigena and was collected by Atkinson, October, 1902.
This collection has dirt attac ned to the basal part of the stem, but
may have been also from buried wood. Іп both collections the
flesh is distinctly yellow, close to lemon-yellow. The plant is
not at all related to P. angustipes, as the scales are larger,
more fibrillose, and scattered, and the colors are distinctly
brighter.

46. Pholiota Schraderi (Peck) Overholts, N. Am. Fl. 10: 271.
1924.

Stropharia Schraderi Peck, Bull. Torr. Bot. Club 32: 80. 1905.

Pileus 5-8 ст. broad, convex to nearly plane, pallid when
young, ochraceous-buff when mature, dry, fibrillose, squamulose,
ог rimose-squamulose оп “һе
disk; context white, with taste
of radishes; lamellae adnzste,
close, thin, whitish then brown;
annulus small, lacerate, white,
sometimes evanescent; stem
central, subequal, squamulose |
and concolorous with the pileus
below, white and mealy above,

А Figs. а ж” P. Schraderi: 149,
solid, 2-4 cm. long, 8-12 mm. Өй... of hymenium showin idi

ng суз
thick; spores ovoid, not apicu- $ г J, and spores [o X 550; i ns

t
late, smooth, 6-8(-9) х 4-6 и; m type specimen

cystidia rather abundant, hyaline, fusoid, projecting rather prom-
inently; also irregular organs as though post-mature basidia im-
bedded in the hymenium.
Habitat: in sandy soil about stumps.
Distribution: known only from the type locality, Washington,
C

'The plants are considerably like P. fulvo-squamosa but the stem
is less prominently scaly, the spores are not at all apiculate, and
cystidia are rather abundant and conspicuous. From P. specta-
bilis it differs in the smooth spores and the cystidia.

bb. Cystidia none.
aaa. Pileus 1--3 ст. in diameter (see also bbb, p. 162).

[Vor. 14
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN

47. Pholiota confragosa Fries, Еріст. Syst. Мус. 169. 1836-
38. РІ, 22, 23.
Pileus 1-2.5 ст. broad, convex to nearly plane, cinnamon-
rufous when moist, tawny when dry, nearly uniform warm-buff
or cinnamon-buff in dried plants, densely and finely floccose-
squamulose or floccose-fibrillose under a lens, in age becoming
somewhat denuded at times but never entirely so, hygrophanous,
margin striate when moist; context fleshy-fragile, pallid, odor
c and taste not marked; gills
ds C adnate or slightly decurrent,
sometimes white-crenulate
on the edge, medium-close,
1-3 mm. broad, rufous to
cinnamon-brown; veil form-
ing а superior, membranous
annulus, erect and sub-rigid
for a time, finally more an-
nulate and in rare cases all
Figs. 151-152. P. confragosa: 151, section but disappearing; stem cen-
uel оба eno the slavateorlence- tral, equal or enlarged just
550; 152, spores, X 1200. All from Overholts at base, concolorous with the
Herb. No. 8061. f
pileus or paler, markedly
fibrillose below the annulus and sometimes white-tomentose or
strigose at the base, floccose and sometimes striate above, 2-5
cm. long, 1.5-5 mm. thick; spores ellipsoid or broadly ellipsoid,
smooth, not truncate, dilutely colored under the microscope,
6-8 Х 4-5 u; cystidia none except for radiating tufts on the edge
of the gill.

Habitat: rotten mossy trunks of deciduous trees.

Distribution: specimens have been examined from Sebec Lake,
Me.; Mt. Mansfield, Vt.; Stockbridge, Mass.; Lake Placid, Adi-
rondack Mts., and Fourth Lake (Herkimer Co.), М. Y.; also re-
ported from Michigan by both Harper and Kauffman.

Illustrations: Fries, Іс. Hym. pl. 105, f. 2; Harper, Trans. Wis.
Acad. Sci. 17: pl. 41, f. D, E.

When once learned this species is among the easiest to recognize
by the peculiar constant color, especially of dried plants, and the
matted fibers on the pileus, resembling as seen under a lens, the
pileus fibers in P. caperata or P. comosa as seen without a lens.

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1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 161

48. Pholiota erinaceella Peck, Rept. N. Y. State Mus. 30: 70.
1878.

Pileus 0.5-2.5 ст. broad, hemispherie then convex or nearly
plane, tawny-brown, not chenging color in drying, dry, densely
covered with minute, erect, pyramidal, spine-like, or granular,
tawny scales; gills adnexed S
or adnate, medium-close or go Jo
subdistant when young, 1-2 Ors
mm. broad, pallid then cin- 29

namon-brown; veil forming NH

a slight, superior, floccose,
central, equal, tawny, with Figs. 153-155. P. erinaceella: 153, section

evanescent annulus; stem 155
of edge of gills showing clavate sterile cells,
numerous Яоссозе scales or х 550; 154, spores, Х 550; 155, spores, Х

granules below the annulus, 200. All from Overholts Herb. No. 3751
smooth above, stuffed or hollow, 1.5-2.5 cm. long, 1-1.5 mm.
thick; spores ellipsoid or naviculoid, smooth, very dilute brown
under the microscope, 6-8 < 4-5 u; cystidia none on the sides
of the gills, protruding abundantly from the edge of the gills.

Habitat: on dead wood of deciduous trees.

Distribution: specimens have been examined from Lake
Pleasant and Boreas, N. Y.; St. Louis, Mo.; also reported by
Harper from Michigan.

Illustrations: Harper, Trans. Wis. Acad. Sci. 17: pl. 51.

The species is well marked by the small stature, the covering
of spine-like scales on both pileus and stem, and the tufts of
cystidia on the edges of the gills. Agaricus (Pholiota) detersibilis
Peck is a synonym.

49. Pholiota muricata Fries, Syst. Myc. 1:244. 1821. РІ. 14.

Pileus 1-3 ст. broad, convex to plane, obtuse or often slightly
umbilicate-depressed, golden-brown or tawny-yellowish when
fresh, brown ог cinnamon-brown in dried plants, covered by a
dense cuticle of short tawny fibrils or fibrillose-tufted scales, or
granulose-squarrose at the center, dry; flesh thin, yellowish, taste
none, odor mild; gills sinusite-adnate, often nearly free in age,
medium-close or slightly distant, light yellow then cinnamon-
brown or rusty-brown, the edge white-crenate, 3-4 mm. broad;

[Vor. 14
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN

veil forming an indistinct superior evanescent annulus ; stem
central, equal, yellowish but with few, rusty-brown, fibrillose,
sub-erect scales or becoming nearly
05%. glabrous, stuffed then hollow, 3-7 cm.
N long, 2-5 mm. thick, sometimes with
SS Migs X] а bright yellow mycelium at the base;
|; PA spores ellipsoid, smooth, 6-8 x 3.5-4 u;
bre е. cystidia none.
NA DE B biet Habitat: on rotting logs of deciduous
156, section of hymenium show- trees.
spores x 1200 All fog Distribution: specimens have been
men at New York from Michi- examined from Ann Arbor, Michigan;
m also reported from Illinois by Harper.
Illustrations: Harper, Trans. Wis. Acad. Sci. 17: pl. 52, 58.
The species as here understood is based on plants collected by
Kauffman in Michigan. There is a resemblance to P. curvipes
but the colors are not so bright and the gills in particular are not
bright ochraceous-orange as in that species. The scales on the
pileus are also different. Р. erinaceella may not be distinct but is
а smaller plant, with more spine-like scales and more brown in
color.

3

bbb. Pileus 2.6-7 cm. broad.

50. Pholiota lucifera (Lasch) Fries, Epicr. Syst. Myc. 167.
1836-38.

Agaricus luciferus Lasch, Linnaea 3: 408. 1828.

Pileus 2-5 em. broad, convex or
plane, uniformly yellow, ochra-
ceous-buff or ochraceous-orange in
dried plants or the center more
tawny, viscid, decorated with small
yellowish or tawny squamules;
flesh yellow; gills adnate, close,
2-4 mm. broad, yellow then bright
rusty-brown, white-crenulate on di z
the edge when young; veil forming X 550; 159, spores, x 1200. АП from
a superior evanescent annulus; specimens at New York from Bresadola.
stem central, equal or slightly thickened at the base, solid, yel-

1927]
OVERHOLTS—THE GENUS ?HOLIOTA IN THE UNITED STATES 163

lowish above, brownish below, peronate-fibrillose, 2-5 cm. long,
3-8 mm. thick; spores oblong-ellipsoid or bean-shaped, smooth,
7-9 x 4-5 u; cystidia none.

Habitat: on dead trunks and limbs.

Distribution: reported by Kauffman from Michigan.

Illustrations: Bresadola, Fung. Trid. 1: pl. 85; Ricken, Blat-
terp. pl. 54, f. 1

European authors agree in describing the plant as with a per-
onate stem, i.e., with a distinct sheath that is at first continuous
as the veil over the pileus. Bresadola represents it as somewhat
glutinous or at least forming glutinous scales. In this character
it seems to be different from P. limonella. I have seen no
American specimens of this plant.

51. Pholiota limonella Peck, Rept. N. Y. State Mus. 31: 33.
1879

Pileus 2.5-5 em. broad, convex or nearly plane, sometimes um-
bonate, lemon-yellow when fresh, retaining the color in the dried
plants or becoming slightly tawny, with scattered, reflexed or
suberect, fibrillose, reddish or tawny CX
scales, viscid; flesh thin, yellow; gills or ae O

sinuate-adnate or slightly adnexed, > y
close, 2-4 mm. broad, whitish, be- 0 pos
coming ferruginous, honey-yellow or ч. (ЛА ҮІІІ
cinnamon-buff in dried plants; veil AN ОИ
forming a floccose, evanescent, yel- c = SS =
low annulus; stem central, equal, Figs. 160-161. P. limonella: 160,
yellowish, with scattered recurved section кы ак etri пы
yellow scales, smooth above the an- баа м (e) X 550; 161, spores, К:
nulus, solid, 3-7 ста. long, 3-5 mm. ! 200. АШ from type specimens.
thick; spores ellipsoid or ovoid, Senn. deep brown, 6.5-7.5 X
4.5-5 u; cystidia none; trama with a distinct central medulla.

Habitat: prostrate trunks of beech and birch.

Distribution: specimens have been examined only from Grif-
fins, N. Y.

Although this species migh: be compared to P. flammans yet the
colors are different, the spores are larger, sterile organs are absent
from the hymenium, and the plant lacks the yellow pulverulent

[Vor. 14
164 ANNALS OF THE MISSOURI BOTANICAL GARDEN

appearance of dried plants of that species. It may not be distinct
from P. lucifera.

52. Pholiota tuberculosa Fries, Syst. Мус. 1: 244. 1821.
РІ. 14.
Pileus 2-6 ст. broad, convex to plane, rarely depressed, obtuse,
more or less ochraceous-orange, ochra-
ceous-tawny in herbarium specimens,
dry, glabrous when young, soon break-
ing up into appressed or erect innate
squamules; flesh yellow, thin, taste
mild, odor none; gills emarginate or
becoming free, broad, yellow then
tawny, the edge serrate; veil forming
& superior, often evanescent annulus;
stem central, with a prominent bulb
from Europe by just at base and rooting below, fibrillose
or slightly scaly, incurved, hollow, yel-
lowish, 1.5-3 cm. long, 1-5 mm. thick; spores ellipsoid or oblong-
ellipsoid, smooth, 6-8.5 х 4-5 д; cystidia none.
Habitat: on dead wood of deciduous trees.
Distribution: reported by Harper from Michigan.
Illustrations: Cooke, Ill. Brit. Fung. pl. 398a (370); Fries, Ic.
Hym. pl. 104, f. 2; Harper, Trans. Wis. Acad. Sci. 17: pl. 41, A,
B

Bresadola.

I have seen no plants referable to this species. Harper's speci-
mens seem to meet the requirements, although the spore measure-
ments (3 X 5-6 u) are somewhat smaller than I find in European
plants.

53. Pholiota curvipes Fr. Epicr. Syst. Мус. 168. 1836-38.

Pileus 2-5 em. broad, convex to plane, ochraceous-orange,
more tawny in age and in herbarium specimens, at first innately
floccose or densely silky-floccose, іп age breaking up into small
fibrillose scales, even, dry; margin even, sometimes appendiculate
from the veil; flesh yellow, thin, taste mild, odor none; gills
adnate, medium-close to slightly distant, bright ochraceous-
orange when mature and in dried specimens, the edge white or

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 165

yellow, conspicuously floccose-crenate, 3-6 mm. broad; veil form-
ing a superior, soon evanescent, radiate-floccose annulus, or an-
nulus none; stem central, equal or

tapering крт; ochraceous-orange ог
ochraceous-tawny, clear yellow at apex, C
decidedly floccose-fibrillose, hollow, 2-5 4.
em. long, 2-5 mm. thick; spores ellip-

soid or oblong-ellipsoid, өкіне 7-8.5 СУ,
X 4-5 u; cystidia none.

Habitat: on dead wood of deciduous 104, Бе. =й. ‘ide ok
trees. put (а), : 550; 165, spores,

qm А 4 Íl from specimen at

Distribution: specimens have been Sor T from Sebec Lake,
examined from Sebec Lake, Ме.; Os- “amne.
ceola, N. Y.; and Creve Coeur, Mo.; also reported from Ann
Arbor, Mich., by Kauffman.

Illustrations: Fries, Ic. Hym. pl. 104, f. 3; Cooke, Ш. Brit.
Fungi, pl. 398 (370) b.

Among the similar-appearing species in the genus Flammula
and the related species in Fholiota it is extremely difficult to set
up specific limitations that will hold. Kauffman seems certainly
to have had my plant, although descriptions are hardly adequate
for the determination of species in this group. In the several
hundred collections of Pholicta at the New York Botanical Garden
I found but one that I think is undoubtedly this species as illus-
trated by Fries. Burt has made two collections in Missouri that
agree in all respects with my conception of this species. The cap
is nearly uniformly ochracecus-orange, the gills of about the same
color near the margin of th» pileus, but darker and more sordid
toward the stem, and the stam is practically concolorous through-
out with the pileus. In the dried plant the scales on the pileus
are appressed and very slightly darker than the cuticle. Another
collection agreeing even move closely with Fries’ illustration is in
the herbarium at Albany and a portion in the writer’s herbarium.
It was collected in New York by Peck. While the colors here
would indicate a relationship with P. limonella, yet the stem there
is distinctly clothed with recurved floccose scales and the gills
are more sinuate or even adnexed as they are also in P. flammans,
which species differs further in having much smaller spores. The

[Vor. 14
166 ANNALS OF THE MISSOURI BOTANICAL GARDEN

affinities of the species appear to me to be more with these enu-
merated than with P. muricata and P. erinaceella. Р. lucifera
seems to differ in having а lemon-yellow viscid pileus and a scaly
stem. It is reported on the wood of coniferous trees.

Without seeing the plants described and figured under this
name by Harper I cannot admit their identity. Тһе scales and
the gills are both described as tawny while in my plants they are
much brighter in color.

54. Pholiota angustipes Peck, Rept. N. Y. State Mus. 30: 40.
1878. PL 38
Plants cespitose, 2.5-7 em. broad, hemispheric becoming convex
or nearly plane, brown or grayish-brown, becoming ochraceous-
brown or subalutaceous, drying between avellaneous and cin-
namon-buff or somewhat ochraceous-tawny, slightly viscid when
moist, squamulose with minute, dot-
like appressed scales; context fleshy,
thin, yellowish or whitish, taste un-
pleasant; gills sinuate-adnate to adnate
or slightly decurrent, medium-close,
= 3-6 mm. broad, whitish or creamy-
Figs. 166-167. P. angustipes: Yellow, becoming tawny-brown, but
166, section of hymenium н 5. cinnamon-buff or cinnamon in dried
(a), and spores (b), x 550; 167, plants; veil forming a slight, usually
ей т ох 1200. All from type evanescent annulus; stem central, equal
or tapering downward, whitish to avel-
laneous, slightly squamose or fibrillose, stuffed or hollow, 3-7.5
ста. long, 4-12 mm. thick; spores ellipsoid, smooth, dilute-brown
under the microscope, 6-8 x 3-5 u; cystidia none or scarcely
noteworthy as small collapsed basidium-like bodies, brown in
color, and occurring with the basidia.

Habitat: in pastures or open woods, commonly near or around
old stumps.

Distribution: specimens have been examined from Schenevus,
Staten Island, and Menands, N. Y.; West Elkton, Ohio; also re-
ported by Harper from Madison, Wis.

Illustration: Harper, Trans. Wis. Acad. Sci. 17: pl. 34.

Тһе species is a well-marked one in the pallid to brown color-

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 167

ation and the minute scales that thickly cover the pileus and to
the unaided eye give the appearance of small dots. Yet in one
ample collection from Ohio some specimens are decidedly squa-
mulose, while other younger ones are entirely glabrous. This
same collection has а reddish hue to the center of the pileus that
is not mentioned in the notes on the fresh specimens. The sterile
organs in the hymenium are quite inconspicuous but show up
much more strongly when the sections are cleared in glycerine.
No distinct annulus is formed and the species may fit better in
Flammula than in Pholiota. Harper's photograph is excellent,
showing extremely well the dot-like scales on the pileus.

ccc. Pileus 6-15 cm. in diameter.

55. Pholiota fulvo-squamosa Peck, Bul. Torr. Bot. Club 30:
95. 1903. РІ. 13.

Pileus fleshy, 6-12 cm. broad, rather thin, convex becoming
nearly plane, dry, covered with a tawny fibrous cuticle of brown-
ish fibrillose scales, the lighter-colored flesh showing up when the
fibers separate into scales, sometimes concentrically cracked about
the disk; flesh white, becoming brownish where cut, with taste
and odor of radishes; gills rather

narrow, close, adnate or joined QO
to a slight collar around the 6 )
stem, whitish becoming pinkish-
cinnamon then dark cinnamon
with a white-crenulate edge; an-
nulus ample, membranous, per- , Ка е фл un Р. , fibsquamosa:
sistent, scaly below, the woper (a), Х 550; 169, spores, x 1200. All
surface striate; stem central, "9?" “Overtolts Herb. No. 934.
equal, stuffed or hollow, covered below the ring with numerous,
erect, subfloccose, tawny sca.es, slightly floccose above the an-
nulus, 5-8 cm. long, 8-10 mm. thick; spores elliptic or ellipsoid,
rather strongly apiculate at the base, dark ferruginous-brown,
smooth, 6-8 х 3.5-4.5 u; cystidia none or not noteworthy.
Habitat: about the bases of trees or attached to buried wood.
Distribution: Michigan.
Illustration: Harper, Trans. Wis. Acad. Sci. 17: pl. 60.
I have seen no specimens cf this plant other than the type at

[Vor. 14
168 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Albany. At first glance the dried plant looks much like an
Agaricus, similar to A. subrufescens. Тһе gills are quite narrow
as in P. duroides. The spores are quite dark and very similar to
those of Agaricus but no Agaricus has so pronounced a scaliness
on the stem. Harper notes the absence of the radish odor and the
color change of the flesh and gives excellent figures of the plant.

56. Pholiota destruens Brond. Plant Crypt. de l'Agen, pl. 6.
1828-30.

Pileus 6-15 ст. broad when mature, heavy-fleshy, convex to
expanded, sometimes umbonate, pallid or more often cervine or
wood-brown, the cuticle sometimes weathering off to a white
color, subviscid, with scattered, large, white, floccose patches or
squamules that are rather adnate
and sometimes imbricate and may
disappear, or with soft white scales;
margin frequently rivulose and with
white fibrils or fibrillose scales; flesh
white, thick, odor not marked, taste
mild or saponaceous; gills adnate to
sinuate, close or crowded, at first
white, finally deep cinnamo n, 4-14
mm. broad; veil white, soon break-
б ing, forming an evanescent, white,
floccose-tomentose annulus; stem
central or excentric, equal or enlarged
downward, white floccose-tomentose above the annulus, with a
few large white squamules or indistinctly and broadly peronate
from the veil fibers, white to wood-brown, 5-15 cm. long, 1-6
em. thick, solid; spores ovoid or ellipsoid, smooth, 7.5-9.5 x
4—6 u; cystidia none.

Habitat: on stumps and trunks of Populus and other deciduous
trees.

Distribution: specimens have been examined from Charlotte,
Pittsford, Knowersville, and Utica, N. Y.; Oxford, Ohio; River
Forest, Ш.; Indianapolis, Ind.; St. Louis, Mo.; Kansas; Denver
and Boulder, Colo.; Priest River, Idaho; Sequim, Wash.; also
reported by Harper from Frankfort, Mich. (as P. heterocliia and
P. comosa).

from Kansas by Bartholomew.

1927]
OVERHOLTS—THE GENUS FHOLIOTA IN THE UNITED STATES 169

Illustrations: Bresadola, Fung. Trid. pl. $4; Hard, Mushrooms,
f. 214; Harper, Trans. Wis. Acad. Sci. 17: pl. 45-47; Kalchbrenner,
Ic. Hym. Hung. pl. 18, f. 1 (as P. comosa).

Bresadola considers P. сотоза and P. heteroclita as synonyms
of this species, and he has been followed in this country by Kauff-
man. Harper, on the other hand, describes and illustrates P.
comosa from Michigan and Illinois as covered with small white
innate fibrils that become separated into small appressed scales,
while his P. heteroclita seems to compare better with the plants I
have referred to P. destruens. At best these plants are all very
closely related and specific limitations are largely a matter of
individual opinion.

Notes on a collection made aear St. Louis by Glatfelter describe
the taste as sweet at first, becoming bitter.

SUPPLEMENTARY КЕҮ TO THE SPECIES
1. Spores rough at maturity (either distinctly so under the high power of the
microscope or requiring the use of the oil-immersion lens)................ 2
meres smooth at all stages. . 1.: 21) QE ает Диа 9
. Wood-inhabiting;! fresh plants bright-colored, i.e., some shade of yellow,
green, or red; gills frequently bright-colored; not viscid; spores 6-10 ш
ИСУ а not noteworthy ЗООНУ оодо: 3
Wood-inhabiting;! gills not bright-colored but cinnamon; spores mostly
6-10 и long; cystidia present though not always abundant, flask-
shaped With а projecting пері. fi. ae oso v's bo oe 6
Ground-inhabiting, in woods; spores variable, but in all species except
ohe mote than 10u long.. A A O оо
Among Polytrichum moss on the ground; very small plants, less than 1
em. broad; stom 1-2 mm. fhich. ЕКЕ... P. minima
. Pileus with distinct shades of greer. or ashy-green when young or on being
handled; flesh green-tinged; gills bright-colored (yellow to ochraceous-
orange) and remaining so in dried plants; pileus scaly with fibrillose
s M ИИА Е ро Vo НЫ у. P. aeruginosa
Planta погон as above. l.. lice S bv as ls ohne Ea
. Pileus 2-5 ст. broad, "РЯ pinkish-red in color, squamulose with іп-
nate scales (i.e., not formed by the separating of a fibrillose cuticle on
the pileus); flesh usually pale ‘ауепдег; gills bright-colored in dried
anig......... 9... ТУ F. ТТІ
Plants not entirely as аһсте:........... ағы, г ЛЕ.
5. Pileus 4-15 ст. broad at жана dry, glabrous or fibrillose, or if some-
what squamulose then the scales definitely formed by the separation
of a fibrillose cuticle; taste bitter or amygdaline............... P. spectabilis

t2

©

>

1 Plants found growing in clusters from a common point, apparently on the ground,
are likely to be attached to buried wood, but this point should be carefully investi-
gated when making the collection.

[Vor. 14
170 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Pileus hygrophanous, glabrous, otherwise as in P. spectabilis....... P. cerasina
Plants hygrophanous (never viscid), watery-brown and striatulate on the
margin when moist, желеді оп п drying, ennamon to tawny in her-
barium specimens, not at a I evanescent
or leaving only a band on the stem. МҮТУМТ ТТГ ТУГЕ? P. marginata
Plants as above except pileus Mae annulus sometimes persistent, and
dried plants cinnamon-buff to сіппатоп....................... P. discolor
Plants as in P. marginata except pens membranous, persistent, con-
о OTE NTT ETT ONT COR P. unicolor
Plants as in P. marginata except gills distinctly forking............. P. furcata
. Spores broadly-ovoid to subglobose, 7-9 X 5-6 и, distinctly verrucose;
pileus entirely covered with matted floccose fibrils, or these collected
into squamules at the center of the pileus; plants with somewhat the
NEN а р, РВС P. trachyspora
Spores ellipsoid or elliptic, more than 12 и long, distinctly verrucose......... 8
Plants slimy-viscid, glabrous; gills not conspicuously banded with al-
ternating light and dark transverse bands; plants western....P. Mcmurphyi
Pileus scarcely moist or sub-viscid, young plants with a coating of hoary
fibrils; gills conspicuously banded with transverse light and dark bands,

e

-1

9o

particularly in dried plants; plants eastern.................... іа

9. Plants growing оп the ground! and with either pileus or stem or both sald. .10
Plants growing on the ground! or among moss and with neither pileus nor

stem scaly, or at most only floccose or вош?у.......................... 17

Plants growing on wood! and with either pileus or stem or both scaly or with
ИКОН I ҚАЙРАҚ. E мт TRENT O

Plants growing on wood! and with neither pileus nor stem scaly............ 52
с а ос оО РЕ ыу қ ады 13
Stem not scaly ог at most only strongly floccose ог scurfy................. 11
11. Spores constantly 6 џи or less long; annulus membranous, disappearing
ИРИ т ОИУ: РЕСУРСТУ ЦИ У P. duroides
А considerable number of spores in each mount measuring as much as 8
or 9 и long; annulus a persistent cottony roll on the stem................ 12
Spores 9-12 и long; annulus membranous, disappearing. ............. P. aurea
12. Pileus brown, ochraceous-brown, or ochraceous-tawny, minutely scaly
all over; annulus slight, еуалевсепі......................... P. angustipes
Pileus Orne to pallid, minutely scaly only at the center; annulus a
persistent cottony roll on the stem ....................... . Johnsoniana
13. Sterile organs of conspicuous form or color entirely lacking from the hy-

enium; pileus and stem covered with a pipi bean that soon
separates into soft silky ды, on both pileus and stem. ................. 14

Sterile organs present in the hymenium, sometimes use Е but some-

times with the aspect of over-mature basidia, and brown in color and

not projecting; plants not otherwise as аһоче.......................... 15
14. Spores 6-8 и long, rather strongly apiculate................ г. fulvo-squamosa
Spores 8-9 и long, not арісшізде................................ P. terrigena

! Plants found growing in clusters from a common point, apparently on the ground,
are likely to be attached to buried wood, but this point should be carefully investi-
gated

1927]
OVERHOLTS—THE GENUS PHOLIOTA IN UNITED STATES 171

15. Scales of the pileus scattered, appressed, sometimes all but disappearing,
rather large; cystidia as brown sterile organs, some imbedded but many
projecting, of definite form; pileus yellow or buff.............. P. rigidipes

Scales of the pileus numerous, small, and appearing as black dots to the
unaided eye, or medium-sized апа more conspicuous; cystidia not as
b

QTD ior СЕ ЗИ | Е vaa EA M A T 16
Scales of the pileus none; cystidia abundant, projecting, flask-shaped. P. subnigra
16. Cystidia occasional, not of definite form but mostly as inconspicuous

brown organs comparable to post-mature basidia, but usually not pro-
jecting; pileus pale brown or pinkish-cinnamon, 3-7 ст. broad.. P. angustipes

Cystidia abundant, projecting, sharp-pointed, many hyaline or with
rounded brownish globules inside; plants dark brown, 1-3 em. broad

VU RP НИРО CU. JE |o opp NEP CENE е P. terrestris
Cystidia rather meme bi eh јанан fusoid, hyaline; plants 5-8 cm. or
more broad when matu ochrae “уу Een TSE P. Schraderi
17. Spores ovoid or pla > ith a gai apex and base more or less
obtuse or rounded though sometimes apiculate........................ 18
Spores variously shaped, not truncate at the apex but sometimes apiculate
кРІЗІеЗме.............-- ара тата bes exor e Wn vete e 5% 27

18. Spores narrow-ovoid, 4—5 и broad, or else annulus median, conspicuous,
striate on the upper side; cyst dia none; slender plants not more than

А от: broad..........-- ЛТ а. смея 19
Plants not entirely as above. о и... чае, 23

19. Plants growing in wet places among Sphagnum moss.......... P. mycenoides
Zune not as above. ......1.. DO Du ue ento err qn 20

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Plants yellowish-red or dark ferruginous and retaining these colors in

herbarium en stem 1-32: 5 и... . Tugosa
21. 00-9 p lotig....... S. о о... P. blattaria
Spores 9-10.5 и koi VUES PEE TUNER Ux. ий 22

22. Stem 2 mm. or more thick; mediv m-slender plants more than 1 em. broad
И а роль, БОСА ooo e P. togularis

Stem about 1 mm. thick; very slender plants less than 1 em. broad... . P. filaris
23. Plants growing on the ground in cultivated fields or grassy places, or in
open grassy woods, or among straw or other waste vegetable matter

Plants growing on the ground in lense woods, or growing on wood.......... 26

24. Spores 10-14 и long; cystidia scarcely projecting, the apex broadly pet
ed; реи white, soon areolatei . O ио ein sickens vermiflua

Plants not entirely as above; spores 8-10.5 и long; cystidia Фа
and projecting conspicuously if ргевепб................................ 25

25. Spores 4-6 и broad; pileus white or tinged with yellow or tan; gills narrow
or medium-broal.....5..1..4.-2 o RERO ERR . praecox

Spores 6-7 и broad; pileus ochraceous-yellow; gills very broad. . . . P. temnophylla

26. Plants growing on wood or on rich humus; pileus rugose at times. P. Acericola
Plants growing on the ground; pileus never rugose................ P. Howeana
27. Spores up to9 u 08g. occ bee deprenos EL RERO a reo челов абе 28

[Уоь. 14

172 ANNALS OF THE MISSOURI BOTANICAL GARDEN

>

оз
ы

wo
с

w

о

о:

e

о
©

. Plants growing among Polytrichum moss, small, less than 1 cm. broad;

spores finely verrucose - the oil-immersion lens............. P. minima
ссы а M EN Li. oo rcs Eq 2^ ere ry core. Pee 29

. Plants small or bi shh less than 4 cm. broad; cystidia none or not
ось АЕ ИИ Л К ТГ AN ERS EET PR TEE 0

Plants larger; cystidia various, sometimes absent...................... . 31
. Annulus median, er. striate on E upper surface....P. Бана
Annulus superior, еуапесепі.................................. P. anomala
. Plants western; Арғы кеней Stem conspicuously white-tomentoes

NE DN MO CIE caus cioe ae S ac So n E EUCH P. albivelata
Plants eastern; dea dry; stem not white-tomentose. .................... 32
All spores 6 р or less long; cystidia rather abundant but not conspicuous,

mostly with a mucronate tip; annulus membranous, disappearing. © durotdes
Some spores in every mount аз much as 8 or 9 и long, mostly 5-9 и; cys-

tidia quite rare and scarcely — or abundant and NE

Р. J

annulus a cottony roll on the вбет........................ ohnsoniana
. Plants small, less than 2 cm. D NOLENTES NEUSS ЦЕН P. subnigra
а ша № ПАРА ИОВ 34

. Plants 2-5 ст. broad; gills adnate or slightly decurrent; stem fibrillose

см Хх адақ мады сары rapaces crea MES roa ыы P. erebia
Plants as above, but stem concentrically white-zoned from the veil. P. platyphylla
Plants 4-10 cm. broad; gills distinctly decurrent................ Р. ombrophila
Plants 10-15 cm. broad; gills 7-20 mm. broad, rounded behind......... P. aurea
. Spores 9 и or less long, or if somewhat longer, then pileus glabrous.......... 36
Spores 9-11 и long, oblong-ellipsoid; brown sterile organs present in the

hymenium; plants western; pileus scaly.................. P. aurivelloides
Spores 11-14 и long, [еее sed ora or fusoid-elliptic; brown sterile

organs absent from the hymenium; plants eastern; pileus scaly . P. albocrenulata
. Stem only scaly and spores with a truncate apex; small plants up to З cm

bro no sterile organs in the hymenium.................. P. mutabilis
У not ontiroly т... ИОС АОИ ОА 37
от. СААР ЛК A У ААРОН 38
И еВ р aio ok Sas ООН К 39

. Pileus pallid to cinnamon when fresh, viscid, densely scaly with erect or
sub-erect, pointed, concolorous scales; stem with concolorous —
рыу РИМНЕН Ao aac fee ali ғазы тақа uarrosoides

Pileus lemon-yellow to tawny or fiery-yellow when fresh, dry, "iih fibril
lose, superficial, sulphur-yellow scales; the stem with yellow Яоссозе
scales or scarcely more than yellow flocci.................... P. flammans

. Plants pallid to brown, grayish-brown, or ochraceous-brown; pileus
squamulose, with abundant minute dot-like appressed scales, never
areolate, 3-7 cm. broad; stem only slightly, if at all, scaly; no conspicu-
ous projecting sterile organs! іп the hymenium; spores 6-8 ш lon
Қалы не EU S d tn Е ЕИО dno wis wide E NUN PR P. angustipes

Plants ochraceous-buff to ochraceous-tawny; pileus glabrous; gills some-
what distant, strongly interveined; stem 8-20 mm. thick, with small
subfloccose evanescent scales; spores 7.5-9.5 и long.......... P. oregonense

+ But there are usually present more inconspicuous bodies, not projecting, and
he appearance of old basidia, irregular in shape. "These are apt to show up

particularly well in permanent sections in glycerine.

1927

]
OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 173

Plants not entirely as in either of the аһохе.............................. 40
40. Hymenium with numerous steril2 brown organs, imbedded or projecting;
all species bright-colored, yellow, reddish, or tawny, and with bo
pileus and stem scaly, and piles more dun 8с С АСАР ори 41
Hymenium with distinct projecing hyaline fuscid cystidia; pileus ap-
pressed-squamose and often areolate; stem squamulose......... P. Schraderi
Hymenium without sterile brown organs or cystidia except perhaps on the
edges of the gills; other characters уагідМе............................ 4
41. e with large appressed spot-like scales; stem increasingly scaly down-
Па... 1. 0 7 Р.а —
Plants р, ТР ТОРТТОРУ е BRE
оа а ООО cM I РИО eM о Р; Ж
у т фе ГУУ re | РРР Е НИ ст
43. Pileus with appressed scales; stem only inconspicuously scaly, 4-6 mm.
с АЕ T ТЕЕ CET s: - . P. rigidipes
Pileus ا‎ recurved scales; stem conspicuously scaly with recurved scales
and 5-12 mm. thick. E 2 5 4 SIE CCA P. squarrosa
44. Large әнің 6-15 ст. broad, white to раШа ог wood-brown, with large

ТМ
л

ТМ
-4

ны
Qo

т
e

сл
bo

сл
(59

5
white floccose spots or patches on the pileus; spores 7.5-9.5 u long. . P. destruens
Large species 6-12 ст. broad, wih fibrillose scales; plants growing at the
base of trees or attached to buried wood; spores 6-8 и long, strongly

| Ю.............. CER А оо P. fulvo-squamosa
Pete not entirely as above... о Ы оосо evan ae 45
БОШ pildus and stem sealy. d.: рауы ek eee rede he ede peas 46
Peut oniy sealy..........1..13:00: о о оос 49

. Pileus deep rich brown or golden brown to бампу........................ 47
Pileus brighter-colored, lemon-yellow to ochraceous-orange................ 48

. Both pileus and stem densely covered with a sheath of small erect, conical,
Pe

Stipertioial scales............10 осо а, rinaceella
Pileus covered with a dense coating of soft fibrillose scales that are erect,

if at all, only in the center of the pileus; stem sparingly scaly or only

Ghetiogs, not sheathed. . ... оао ene rn P. muricata

. Stem with a distinct bulb just at base; not sheathed but scaly...P. tuberculosa
Stem not with a bulb at the base, but sheathed and scaly.......... Р. luci
Stem not with a bulb at the base; not sheathed but scaly.......... P. limonella

. Pileus 1-2.5 ст. broad, hygrophanous; gills cinnamon; annulus —€—

регин, menibrapous. .1..] 5o eek ENR a e eos P. confragosa
Pileus 2-10 em. broad; 4” bright ochraceous-orange; annulus fibrous-
lacerate andeyaánescent. . 1..1... TTE TERRE Ie epe atro e Be deals

. Pillows то. ТР ood. 1G 2277 Mew P. lucifera
Pileus diy.. 22... 2.1.45 КЕЛШЫШ 52 СУР 51
Pileus soon areolate and often with greenish tints; edge of gills жеу

4244.24... 2. А S о ДОКТ {поза
Pileus not areolate, never greenish; edge of gills conspicuously don d
érenate.......24. eects © fee ods ds Sok os ONERE ri . curvipes

. Pileus Носсове or silky-floccose, | 2-5 cm. broad; gills, pileus, and stem

bright ochraceogs-ot ge. 1. Д е . curvipes
Plante not entirely aa BOTE о PB eek var 552226221222. 53

[Vor. 14
174 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Plants at maturity more than 4 cm. Моай.............................. 56
54. Spores with a truncate apex, 6-8 u long; cystidia none; veil evanescent

Spores not truncate, 7.5-9.5 и long; flask-shaped cystidia present and
projecting, though not ADORE 22,................,,............... 55
55. Pileus viscid; annulus вирегвінбепб............................. P. discolor
Pileus dry; annulus at first funnel shaped, persistent and —Ü
А О QR E dori ci C SOR v» rs o ic ia С) nicolor
Pileus dry; annulus usually persistent but not funnel-shaped. . . . P. бонсай
56. Pileus ochraceous to tan; annulus long-persistent, membranous............ 57
Pileus bright-colored, oobracsous-lnff to tawny; eagle a soon evanescent;
gills rather distant and the gill cavities strongly veined........ Р. oregonense
57. Cystidia rather abundant, flask-shaped, strongly Динне pileus often
rugose; on rotten wood or on Һатив....................... . Acericola
Cystidia present and rather abundant but projecting only slightly and
more fusoid than flask-shaped; on recently felled logs or from wounds
"4а ОЕ ПЕН P. Aegerita

SYNONYMS AND DOUBTFUL AND EXCLUDED SPECIES

Pholiota aggericola Peck is P. erebia of this paper. Зее p. 119.

Pholiota appendiculata Peck, Bul. N. Y. State Mus. 94: 33.
1905. = P. ornella Peck, which is referable to Flammula polychroa
Berk.

Pholiota autumnalis Peck is P. marginata (Batsch) Fries. See
p. 134.

Pholiota dactyliota (Berk. & Mont.) Басе. Syll. Fung. 5: 750.
1887 (Agaricus dactyliotus Berk. & Mont.; Mont. Syll. Crypt.
115. 1856). Described from plants collected in Ohio by Sul-
livant, and said to be similar to P. squarrosa.

Pholiota comosa Fries is here regarded as a form of P. destruens
Brand. See p. 169.

Pholiota detersibilis Peck is P. erinaceella Peck. Зее p. 161.

Pholiota dura (Bolt. Quel. Champ. Jura Vosg. 91. 1872
(Agaricus durus Bolt. Hist. Fung. pl. 67, f. 1. 1788). A species
reported in every list of fleshy fungi issued in America. All col-
lections examined belong as well under P. vermiflua Peck which
will probably be found to be a synonym of P. dura. Europeans
are far from an agreement as to the limits of the species and
Peck’s name can be used for the present. See p. 105.

Pholiota heteroclita Fries is here regarded as not distinct from
P. destruens Fries. See p. 169.

Pholiota hormophora (Mont.) басс. Syll. Fung. 5: 754. 1887

1927]
OVERHOLTS—THE GENUS PHOLIOTA ІМ THE UNITED STATES 175

(Agaricus hormophorus Mont. Syll. Crypt. 116. 1856). Col-
lected by Sullivant in Ohio and described by Montagne. Said to
resemble P. tuberculosa Fries and is described as having a bulbous
enlargement at the base of the stipe.

Pholiota indecens Peck is Р. ombrophila Fries. See p. 121.

Pholiota lutea Peck is here treated as а form of P. spectabilis
Fries. See p. 145.

Pholiota luxurians (Fries) Gill. Champ. Fr. 439. 1876. Re-
ported by Harper from the Great Lakes region. The specimen
seems to be related to P. aeruginosa Peck.

Pholiota mollicula Banning in Peck, Rept. N. Y. State Mus.
44:182 (70). 1891. Originally described from Maryland, grow-
ing on the roots of trees.

Pholiota ornella Peck, Bul. №. Y. State Mus. 122: 151. 1908
(Agaricus ornellus Peck, Rept. №. Y. State Mus. 34: 42. 1883).
Not distinct from Flammulo polychroa Berk.

Pholiota radicosa (Bull. Quel. Champ. Jura Vosg. 92. 1872
(Agaricus radicosus Bul. Herb. Fr. pl. 160. 1783). Reported
from the Pacific coast by Harkness and Moore but I have seen
no specimens of this highly characteristic species.

Pholiota sabulosa Peck, Bul. Torr. Bot. Club 23: 414. 1896.
Described as growing in sandy soil in Alabama. The spores are
rough-walled, 8.5-9.5 x 5-5y. Flask-shaped cystidia project
from between the basidia. }Зо+һ of these characters ally the plant
to the P. marginata complex in which there is already an over-
abundance of described species. Specimens in the Underwood
Herbarium at New York show the plant to have been growing
from humus-charged earth, and it is probably to be regarded as a
form of P. marginata or P. discolor.

Pholiota speciosa Clements, Bot. Surv. Neb. 2:41. 1893. The
description is inadequate fcr the recognition of the species. И
the spore measurements (5 X 4.5 y) are correctly recorded it
would fall in the neighborhood of P. duroides, with which it seems
to have other characters in common.

Pholiota sphaleromorpha (Bull.) Quel. Champ. Jura Vosg. 91.
1872 (Agaricus sphaleromorphus Bull. Herb. Fr. pl. 540. 1791).
Harper is of the opinion tha; P. Howeana Peck is referable to this
species. At all events it is very similar and there are now too

ГУ оз. 14
176 ANNALS OF THE MISSOURI BOTANICAL GARDEN

many species described with the peculiar truncate spores, the
prominent cystidia, and other characters common to this group.
See p. 109, 111. қ

Pholiota subsquarrosa (Fries) Sace. Syll. Fung. 5: 750. 1887.
Reported by McIlvaine. I have seen no specimens so referable,
and Mellvaine records that the plants seem different from the
European species. See p. 151, 156.

Pholiota terrigena (Fries) басс. Syll. Fung. 5: 737. 1887. Тһе
species has been reported from the United States but I have seen
по material that corresponds to specimens from Bresadola. Тһе
plants so recorded should be compared with P. terrestris Overholts,
which has smaller spores and prominent cystidia. See p. 127.

Pholiota ventricosa Earle is P. spectabilis Fries. See p. 146.

Pholiota villosa (Vries) Басс. Syll. Fung. 5: 752. 1887. Speci-
mens so determined by Peck and similar collections from the
Pacific coast seem referable to P. spectabilis.

Pholiota washingtonensis Murrill, Mycologia 4: 259. 1912.
Seems to be unquestionably P. ombrophila. See p. 121.

INDEX TO SPECIES

New species are printed in bold face type; synonyms and less important bi-
nomials in ?/alics; and previously published valid names in ordinary type.

Page Page
Agaricus Hypodendrum
с И eee 1 оғедотетге..,.......,......... 140
о RRTEEREETRTTO TTE 122 Pholiota
SUE Lco ere ya OP ERA т с ЕМ ТТТ СТ 108
ашитпайг.................. 134 o. ПАНА E 154
ا‎ 123 BRUN 6,55 ee e e 139
дасіуұйоіме................... 174 var. strobiloidea............ 140
с. Рег 161 сс с КҮТТІ АНИ
о 2, СІ; 148 aggertcola. .......... 119, 121, 174
Лоғторлогиа................. 175 оз... FEE T EE T ETT ИИ 12
/оһпзотїатив................ 180 albocrenulata................
son Lo ы ыны зэ». Тел ES DE 162 SM, ИЯ 166
татфўїтайиз.................. 182 ПИ: к. 127
DENN МАРИСА 138 аррепйсшаа................ 174
кабы oe 106 Clio. ОР 121
са ETTET 175 аза. ТТС ТУТ 152
йетпорһуИив................. 112 aurivelloides ............... 15
т 535552222. 114 ашичттайв............... 134, 174
ит 134 с АС ЕО 113
Flammula OSDOPAUA. ОА E 1
асе 136 CRORE о hus
BH еее нае ея 148 о «оа cre кен: 169, 174
Hebeloma соштабова..................

1927]

OVERHOLTS—THE GENUS PHOLIOTA IN THE UNITED STATES 177

Page
Pholiota eonim
e К OR NEU Tee 7
үз жумам а тае 38
INNER 161, 174
e а е O 35
p pa Oe EM. 105, 174
var, iy ақын EE HEX д) 1
АР, ТЕР ;
КЕК 25.2.., 25222.
но up. os e acer ES )
BOUND AS Gre EO 3
| ulvo-equamosa gom Ion 137
с VIA uou. SSS a ]
re TA PA РУ воан 169,
оС”, PESE EE T CERES s 7
ТОРКИНИ. у. суу з... rS ER
те” БОГЕТ ТЕРГЕН > 121,
NOlnBOBISBA. recen et e riri E
mr PRU en
pono m ET n
coe MOVE 145,
Tuteofolia Gane e
DR О ВОО NX - |
ИВ TERETEEEEEEER QU 7
eS SPREE у» 137
МетчгрАуі................. 124
ШИШЕ ООРОО Я Е 125
ТООН ЖОЙ, С 175
сыт 1
ВАА" IS 138
г ОНОРЕ ТЕ С 11

о TUNE ТТУ Ў 120

Раде
prede (m
E eA цы Тыр Ser 140
VOTE Eon AU NINE А 175
чалык Ри ЛЕ Жс, Bu ear E 117
DIBOCUE,..): 49 aA о ore 106
тт ls Ree E TE Е 175
с ЕРЕСТА aa 158
с | МАРИ 142
таёов&.'. „4 ЗЕ ое 115
aabulosn. i. Love e ое е 175
Sehraderi; . 2 ЕБ е 159
аребїо8@. o ia e 175
spectabilis.. e
8 eam ЕЕ 109, 111, 175
BqUAITOSS.. ev Soon ce eee 157
equarroscides чи E 155
SUDE. gesso pa a
subsquarrosa......... 151, 156, 176
iemnonhylla.....- 0223: 3
MEN. rrr nara Ea 126
CUT ЕЕ ТЫ ВЕ: 127, 176
ee RS EPEC Ee а 114
ср. О Pee TO 116
о Ж ND QUON > 125
DUDEN аео oe ae 164
be sve eon oh УУ Y 134
ATA ILS. Р Е 146, 176
es Т7 04
SE иЕе АНЫН, pe
washingtonensis...... 119, 121, 176
Stropharia
ДТО Т 110
CECI S dues rame Жон 159

[Vor. 14, 1927]
178 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 8
P. vermiflua. Photo of specimens in Overholts Herb. 3248, Х 2/3. Original.

ANN. Mo. Вот. Ganp., Vor. 14, 1927 PLATE 8

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
180 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 9

. praecox. Photo of specimens in Overholts Herb. 3318. Upper half, X 3/5,
lower half, x 4/5. Original.

ANN. Mo. Bor. Санр., Vor. 14, 1927 PLATE 9

Y

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
182 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 10

P. Acericola. Photo of specimens in Overholts Herb. 3911, Х 3/4. Photo by
E. T. Kirk.

PLATE 10

. Мо. Вот. Санр., Vor. 14, 1927

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
184 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 11

P. erebia. Photo furnished by E. T. Harper from plants collected in Michigan.
About natural size.

PLATE 11

ARD., VoL. 14, 1927

*
x

Mo. Bor. €

ANN.

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
186 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 12

P. aurea. Photo of specimens in Overholts Herb. 8364, collected in British
Columbia by J. Schmidt, October, 1922, and communicated by W. S. Odell. Photo
by Drayton. Reduced.

PLATE 12

27

NN. Mo. Вот. Garp., Vor. 14, 19

A

4

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
188 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 13

P. fulvosquamosa (left half). From Harper, Trans. Wis. Acad. Sei. 17: pl. 60.
1913. About natural size

P. caperata (right hal) Photo of specimens in Overholts Herb. 7967, Х 1.
Small specimen. Original

3

«
€

PLATE 1

27

14, 19

ANN. Mo. Bor. Ganp., Vor.

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
190 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 14
Fig. l. P. marginata. From Harper, Trans. Wis. Acad. Sci. 17: pl. 54, fig. C
1912. About natural size.
igs. 2-3. P. muricata. From Harper, Trans. Wis. Acad. Sci. 17: pl. 52, fig

A, B. 1912. About natural size.

Fig. 4. P. tuberculosa. From Harper, Trans. Wis. Acad. Sci. 17: pl. 41, fig. B.
1912. About natural size.

ANN. Мо. Вот. Garp., Vor. 14, 1927 PLATE 14

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
192 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 15

P. discolor pg half). Photo of specimens collected in South Boulder Сайоп,
Colorado, July 18, 1923, elev. 10,000 ft. About natural size. Origina

P. rugosa die left). Photo of specimens in Overholts Herb. 3623, xL
Original.

P. togularis (upper, right). From Harper, Trans. Wis. Acad. Sci. 17: pl.59. 1913.
About natural size. (Published as P. blattaria.)

ANN. Mo. Вот. Garp., Vor. 14, 1927 PLATE 15

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
194 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 16

P. spectabilis. Photo of specimens in Overholts Herb. 9889. Reduced somewhat.
Photo by L. W. Brownell.

PLATE 16

OVERHOLTS—PHOLIOTA IN THE

UNITED STATES

Vor. 14, 1927]
196 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 17
P. spectabilis. Reduced somewhat. Photo by L. W. Brownell.

PLATE 17

ARD., Vor. 14, 1927

Mo. Bor. G

ANN.

OVERHOLTS

PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
198 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 18

P. aeruginosa. About natural size. Photo by Burtt Leeper.

15

PLATE

ANN. Mo. Bor. GARD., Vor. 14, 1927

25 Wu
47

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
200 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION ОЕ PLATE
PLATE 19

P. albocrenulata — part). From Harper, Trans. Wis. Acad. Sci. 17: pl. 42.
1912. About natural si

P. flammans (lower өлені Right, from Fries, Іс. Hym. pl. 104, fig. 1; left, from
Harper, Trans. Wis. Acad. Sci. 17: pl. 41, fig. C. Both about natural size.

ANN. Мо. Вот. Ganp., Vor. 14, 1927 PLATE 19

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
202 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION ОЕ PLATE
PLATE 20
P. adiposa. About natural size. Photo by L. W. Brownell.

20

PLATE

27

Vor. 14, 19

Вот. GARD.,

ANN. Mo.

OVERHOLTS

PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
204 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 21

P. squarrosa (upper half). About natural size. Photo by L. W. Brownell.
P. adiposa (lower half). About natural size. Photo by L. W. Brownell.

ANN. Мо. Вот. Gan»., Vor. 14, 1927 PLATE 21

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

" [Vor. 14, 1927]
206 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 22

. Squarrosoides - cluster). From Harper, Trans. Wis. Acad. Sci. 17: рі. 36.
1912. Slightly reduced.

P. confragosa Бине. From Harper, Trans. Wis. Acad. Sci. 17: pl. 21, fig. D.
1912.

PLATE 22

XARD.,

ANN. Mo. Bor. €

2
%..

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

[Vor. 14, 1927]
208 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 23
P. angustipes (except lower left НЫ From Harper, Trans. Wis. Acad. Sci.
17: pl. 34. 1912. About natural s

P. confragosa (lower left corner). p^ slightly enlarged. Photo presented
by E. T. Harper.

ANN. Mo. Вот. Ganp., Vor. 14, 1927 PLATE 23

OVERHOLTS—PHOLIOTA IN THE UNITED STATES

° [Vor. 14, 1927]
210 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
_ PLATE 24
P. aurivella, Somewhat reduced. Photo Бу І. №. Brownell.

ANN. Мо. Вот. Garp., Vor. 14, 1927 PLATE 24

LVIS ЧМ AHL NI V.:LOF'IOHd—STL'IOHSIWAO

r
H
GR

К

Annals
of the
Missouri Botanical Garden

Vol. 14 SEPTEMBER, 1927 No. 3

A MONOGRAPH OF THE SECTION OREOCARYA OF
CRYPTANTHA!

EDWIN BLAKE PAYSON?

Professor of Botany, University of Wyom
Formerly Teaching Fellow in the Henry Shaw School of Botany of Washington University

INTRODUCTION

The group of plants treated in the present monograph is re-
markably characteristic of the Upper Sonoran areas of the Great
Basin region, although a few species are native to the eastern
foothills of the Rocky Mountains, one or two have penetrated
to the Canadian plains and two or three occur in northern
Mexico and adjacent Texas. In western Colorado, Utah, Ne-
vada, and southern Wyoming one or more species may be found
on almost any barren hillside. Many seem to prefer soils that
are so strongly impregnated with mineral salts that few other
plants can compete with them. No species seems to be able to
tolerate a moist or undrained situation.

These plants are often transient occupants of any habitat.
They seem particularly at home on shifting or disturbed soil. A
loose hillside or shale outcrop is a favorite locality for the com-
moner kinds. And yet they are never weeds in cultivated
ground—that distinction is reserved for the annual members of
the genus. What the factors are that determine this tendency
to occupy changing habitats is as yet unknown to the author—
one of many ecological problems suggested by the present study.

The members of the section Oreocarya are very similar in
general appearance. They are usually gray with numerous tri-
chomes and in most cases are beset with harsh bristles that render

1 Issued October 8, 1927.

? Since the receipt of the manuscript of this paper, Dr. Payson passed away on
May 15, 1927.

ANN. Mo. Вот. Garp., Vor. 14, 1927 (211)

[Vor. 14
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN

them unpleasant to human hands. As a group, these plants are
inconspicuous in a country where inconspicuous plants are the
rule. They are at home with various species of Atriplex, Arte-
misia, Astragalus, Eriogonum, Abronia, Lesquerella, Physaria,
Mentzelia, Ephedra, Townsendia, and other similar forms. Many
are rather regularly associated with the desert juniper (Juniperus
utahensis) and the pinyon (Pinus edulis).

They are plants of the waste lands of the arid west; lands that
are waste not because man has abandoned them but because he
has never thought them worthy of his attention. From a con-
ventional viewpoint these species are quite unattractive. Even
in the largest-flowered species one can scarcely imagine them
being gathered for ornamental purposes. They are neither suf-
ficiently bizarre to render them noteworthy, sufficiently trouble-
some to make them objects of concern, nor common enough near
the habitations of men to have merited for them the distinction
of a common name.

In spite of the apparent unattractive nature of these plants
they are of remarkable taxonomic interest. As a group they are
not easily distinguishable by superficial peculiarities but possess
technical characters of rather remarkable constancy. The geo-
graphical distribution of the species is decidedly interesting.
Many are very local, and most conform very closely to natural
areas. The present study has shown that this group of plants
exhibits phylogenetic tendencies of importance to students of
evolution in the flowering plants. Doubtless certain features
of their ecological relationships would offer a promising field for
further studies.

PHYLOGENY
I. THE SECTION OREOCARYA

The author finds that the most successful method of attack on
any phylogenetic problem comes from comparing closely re-
lated forms. Some at least of the species that occupy contiguous
areas and differ only by minor characters may be thought to
stand in the father-son relationship to one another. Occasion-
ally, even, we may expect to find series of three species that
stand in the grandfather-father-son relationship. Various con-

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 213

siderations of age, variability, stability of environment, etc. may
be expected to influence the nature of the descendants, but it
does not seem probable that these disturbing factors will so
confuse the phylogenetic pattern as to make it entirely unintel-
ligible. Even if of two, or three, closely related species no one
represents exactly the ancestral form it will often happen that
one remains more primitive than the other, and consequently the
direction of evolutionary change will be quite as evident as though
one species had remained stationary while the other diverged.
If, by studying closely related forms, it can be discovered which
is the ancestor and which the descendant, or which the primitive
member and which the specialized one, evidence may be gained
for use in building a theory of phylogeny. И one bit of such
evidence is consistent with another gained from a consideration
of other characters, then the hypothesis gains in stability. In
this way, and piece by piece, the evolution of the twigs may be
interpreted, and finally by putting all the facts together it may
be possible to make a very probable guess as to the evolutionary
form of the branch as a whole.

Тһе following examples will make clear how this method has
been employed in the present case.

C. Palmeri and C. Jamesii var. multicaulis are very closely re-
lated forms. Тһе differences between them are: (1) C. Palmeri
lacks a circle of crests at the base of the corolla tube, while in
multicaulis the crests are very well developed; (2) C. Palmeri
has а much more conspicuously accrescent calyx than C. multi-
caulis; (3) in C. Palmeri the style is longer than in C. multicaulis
and so also is the corolla tube in proportion to the corolla lobes.
One of the best arguments for the derivative nature of any species
is the absence of parts that are characteristic of the family to
which it belongs. Тһе lack of crests is such a character. Their
presence is widespread in the family, and their loss is held to be a
reduction pointing the direction of evolutionary change. Тһе
presence of accrescent calyx lobes in Palmeri strengthens the
argument that C. multicaulis is a more primitive species than the
other. It is now reasonable to suppose that in correlation with
these characters, the other differences are significant as showing
the course of change.

[Vor. 14
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Another interesting pair of closely related species is C. oblata
and C. Jamesii var. multicaulis. These are not so closely related
as were the other pair and yet their relationship is quite evident.
In the original description of C. oblata, M. E. Jones cited a speci-
men of C. Jamesii var. cinerea as typical of the new species. In
comparing these two we notice the following significant dif-
ferences:

C. oblata C. Jamesüi var. multicaulis
Crests lacking at base of tube Crests present
Corolla tube exceeding the calyx Corolla tube and calyx subequal
Calyx conspicuously accrescent Calyx scarcely accrescent

Flowers heterostyled Flowers uniform

Cymes contracted Cymes elongating

Nutlets somewhat roughened Nutlets smooth dorsally
dorsally

Indument conspicuously setose Setae less conspicuous

Following the lead given in the other case C. multicaulis may
be considered primitive in having crests at the base of the tube
and a scarcely accrescent calyx, while C. oblata is specialized in
these characters. Nor is it unreasonable to think of roughened
nutlets being more specialized than smooth ones and conspicuous
setae developed later than the less setose trichomes. It now
seems probable that the long corolla tube is derived from a
shorter one and that heterostyled flowers have been developed
from uniform flowers within the genus.

Other examples might be given. C. confertiflora and C. flava
are very closely related and have been confused in the past with
C. leucophaea. Тһе relationship of these three species is so close
that any one of them might be considered the ancestral form or
that all three might have come from a very similar common
ancestor. The latter supposition is the more probable. The
phylogenetic conclusions will not be very different in either case.

C. confertiflora C. flava C. leucophaea
Crests usually evi- Crests usually ob- Crests usually evi-
ent solete dent

Conspicuously het- ^ Conspicuously het- ^ Less heterostyled
‚ erostyled erostyled

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 215

Nutlets ovate Nutlets lanceolate Nutlets ovate
4 nutlets mature 1-2 nutlets mature 3-4 nutlets mature
Flowers yellow Flowers yellow Flowers white

On the basis of the absence of crests in the corolla tube it may
be concluded that C. flava is more specialized than are either of
the others. When the data on the number of nutlets that mature
are considered (and these data are very meager in the case of C.
leucophaea) evidence is obtained for a belief in the primitive
nature of C. confertiflora and C. leucophaea as compared with
C. flava. Since the lanceolate nutlets are found in that form where
one or two nutlets quite regularly abort, and the species with
ovate nutlets usually mature all of their fruits, it is logical to
consider the lanceolate nutlets as derived from the ovate ones be-
cause their occurrence in this trio is associated with another char-
acter about which there can be no doubt as to the phylogenetic
interpretation. С. leucophaea seems to be less distinctly di-
morphie in flower structure than is C. confertiflora; also the
corolla tube is not so long, the flowers are white rather than
yellow, and the indument less coarse. These are all charac-
ters that were thought of as primitive in connection with C.
Латезй var. multicaulis when contrasted with C. oblata and С.
Palmeri.

It is of interest to make an especial study of those species that
lack crests at the base of the tube. These species may be con-
sidered as specialized in that regard and since forms that are
highly developed іп one character may be expected to show recent
rather than primitive qualities in other characters, it might be
expected that on the whole the characters of this group of species
would be specialized. Тһе species that quite regularly lack crests
at the base of the tube are: Palmeri, echinoides, fulvocanescens,
flavoculata, Bakeri, oblata, flava, paradoxa, Jonesiana, longiflora,
and tenuis. In addition to these, confertiflora, Osterhoutii,
Wetherillii, and mensana have poorly developed crests or at times
none at all. In the first group, those species that are usually with-
out crests, all but one have corolla tubes that exceed the sepals.
In the second group two have long corollas and two short ones.
Considering both groups together, we have fifteen species of which
twelve have long corollas and only three short corollas. Nine out

[Vor. 14
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of the fifteen have distinctly heterostyled flowers. These three
characters, long corollas, heterostyled flowers, and lack of crests at
the base of the tube, were considered specialized when C. oblata
was compared with C. Jamesii var. multicaulis. Some evidence
has been obtained that confirms the earlier judgment.

If the group of fifteen species listed in the preceding paragraph
is examined as to the relationship of the species involved, one
interesting fact is evident. These forms are not considered as
closely related among themselves. In other words, the group is
almost certainly polyphyletie. С. Palmeri, oblata, confertiflora
and flava belong to the series of which C. Jamesii is the primitive
member. С. Jonesiana, C. echinoides, and C. fulvocanescens are
related to C. breviflora in the short-flowered group. The remain-
ing species are possibly closely related to one another but seem
far removed from the breviflora or the Латезй group. Тһе con-
clusion seems evident that the fifteen species listed above com-
prise a highly specialized artificial group selected from several,
perhaps three, natural groups. If this is the case, then hetero-
styly and the long corolla tube have been evolved more than once
within the section Oreocarya.

Тһе question of the primitive nature of the perennial or annual
habit is of much interest. Morphologists, in general, seem to be
of the opinion that the perennial and even the arborescent habit
is primitive for the Angiosperms. With this generalization the
present author is inclined to agree but he believes he has found
good evidence for the primitive nature of the annual habit in the
Cruciferae. Because of this apparent contradiction the question
in the section Oreocarya has received particular attention. If
C. Sheldonii and C. Bradburiana are compared, it is noticed that
here are two very closely related species, of which one is perennial
and the other biennial. Тһе nutlets in the former are distinctly
smoother, and consequently more primitive, than in the latter.
The geographical location of C. Bradburiana on the periphery of
the generic range makes it seem reasonable also to consider this
species a recent derivative from Sheldonii—so recent а derivative
indeed, that there are difficulties in the way of according it com-
plete specific independence.

Such arguments as these might be continued to include all the

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 217

evidence at hand for the phylogenetic conclusions that have been
reached. This, however, seems of rather doubtful value since
any who may wish to challenge the conclusions will wish to review
the details of specific differences throughout the section. Enough
has been given to show the method of attack, and some evidence
has been adduced for some of the more important conclusions.
Without further justification, the following summary of phylo-
genetic conclusions will be presented. These generalizations are,
of course, only expected to apply to the evolution of the section
Oreocarya of Cryptantha.

1. Smooth nutlets are more primitive than roughened ones.

2. In the course of evolution the nutlets were first roughened
on the dorsal surface and finally on the ventral surfaces.

3. The scar in the primitive nutlets was straight and narrow
and without an elevated margin.

4. The nutlets of the more primitive species are attached to
the gynobase at а lower point than in the specialized species.
(This is in agreement with Dr. Ivan Johnston's theory of the
phylogeny of the genera of this family—see Contr. Gray Herb. 74:
14-19. 1925).

5. Тһе ovate nutlets are more primitive than the elongated or
lanceolate type.

6. The primitive corollas in this group were uniform and did
not exceed the sepals.

7. Heterostyly has apparently been developed а number of
times within the section and seems to be regularly associated with
long corolla tubes.

8. Crests at the base of the corolla tube were present in the
primitive species and have become obsolete in the more specialized
ones.

9. Тһе accrescent sepals denote specialized species, and those
forms with sepals that enlarge but slightly in fruit are to be con-
sidered primitive in this regard.

10. The primitive inflorescence was confined to the upper part
of the stem and was composed of conspicuously elongating scor-
poid eymes. Asdevelopment proceeded the cymes became shorter
and the inflorescence tended to extend over more and more of the
stem.

ГҮ ош. 14
218 ANNALS OF THE MISSOURI BOTANICAL GARDEN

11. The rather slender unbranched tall stem is considered more
primitive than the very stout stem or the short, branched one.

12. An extremely setose indument is thought to be a mark of
specialization, and in like manner a very sericeous and uniform
covering is a specialization in another direction.

13. The primitive species were perennials and the less enduring
forms are specialized.

14. Тһе linear-oblanceolate leaf is more primitive than the
broader ovate or spatulate types.

П. THE GENUS CRYPTANTHA AS А WHOLE

A detailed study of each section of Cryptantha in which the
evolutionary tendencies were traced from species to species and
from group to group would prove of very great phylogenetic
interest. When that study is made, then, and only then, will it
be possible to discuss the phylogeny of the genus in a compre-
hensive way. However, even with only a superficial knowledge
of the genus as a whole, certain conclusions seem certain enough
to warrant some discussion of them.

Two of the sections are characterized by the absence of cleis-
togamous flowers. This is so evidently a primitive character
that these two sections, Oreocarya and Krynitzkia, are considered
the primitive sections of the genus. Of these two sections one,
Oreocarya, is prevailingly perennial, the other prevailingly annual.
On this basis, then, Oreocarya is to be considered the most prim-
itive of the four. Oreocarya differs from Krynitzkia also in other
ways that are significant. In the former the flowers are mostly
large and the cymes are usually bracteate, while in the latter the
flowers are usually minute and the cymes usually ebracteate.
In Krynitzkia the nutlets are often heteromorphic, while in Oreo-
carya the nutlets may not all develop but there is no tendency to
heteromorphism. АП these differences go to show that Oreocarya
is, аз а whole, more primitive than is Krynitzkia. However, the
section Krynitzkia may not be considered as made up merely of
specialized descendants of recent species of Oreocarya. For one
thing, the elongated cymes, a primitive character, that are so
characteristic of Krynitzkia must be traced back to a primitive
rather than to a specialized Oreocarya species or Oreocarya an-
cestor.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 219

The other two sections of Cryptantha, Geocarya and Eucryp-
tantha, are, asit was stated before, characterized by the presence of
cleistogamous flowers. Іп the former these are highly specialized
and curious structures from near the surface of the ground, in the
latter they occur in the inflorescence or in the axils of the cauline
leaves and are not so highly transformed. It seems evident that
of the two, Geocarya is to be considered the most specialized
section of the genus. Both Geocarya and Eucryptantha have
perennial primitive representatives as well as annual specialized
ones.

Тһе author interprets these facts concerning the characteristics
of the sections as follows:

The perennial species we now place in the Section Oreocarya
are the most primitive we know in the genus. Тһе most primi-
tive of these is C. Jamesii var. multicaulis. This variety has so
many primitive characters that it may be used to visualize the
ancestral form from which the others developed without doing
violence to the known facts of phylogeny. It would seem reason-
able to suppose that the primitive Oreocarya species evolved in
four rather distinct directions. The species of Oreocarya re-
mained perennial and showed & marked tendency to develop
larger flowers and to reduce the length of the cymes. This group
is probably monophyletic in the sense that a single primitive
species gave rise to several species that in their turn produced
others. Development in another direction produced annuals
very early in the history of the genus. "These annuals showed a
marked tendency to reduce the size of the flowers, differentiate
the nutlets in a single flower, and to elongate the cymes. This
group we now know as the section Krynitzkia. It is probably
truly polyphyletic, and species grouped here have arisen indepen-
dently from several perennial ancestors.

Тһе development of cleistogamous flowers of the two kinds
undoubtedly began before the perennial habit had been lost since
both the sections Geocarya and Eucryptantha have primitive
perennial members. These sections followed lines of development
very similar to those of Krynitzkia and so the species of these
three sections are very similar in appearance. It seems rather
doubtful if Geocarya is polyphyletic because of the very peculiar

[Уоь. 14
220 ANNALS OF THE MISSOURI BOTANICAL GARDEN

cleistogamous flowers common to the species. Hucryptantha,
however, may have arisen a number of times from Krynitzkia—
whenever cleistogamous flowers were developed.

III. THE MECHANISM OF EVOLUTION

It is the author's contention that a taxonomic and phylogenetic
study furnishes one of the best backgrounds for an evaluation of
various theories of the mechanism of evolution. To make the
most of this opportunity the taxonomic-phylogenetic study should
be followed by an investigation into the variability, heredity,
ecology, etc., of the group of plants under consideration. In the
present case this has not been done, and it is realized that any
conclusions the author may have reached must be offered merely
as suggestions. Тһе following speculations are presented for
what they may be worth.

Natural selection.— The species of the section Oreocarya are
much too beautifully adapted to their surroundings to permit the
theory of natural selection to be discarded in any consideration
of the evolution of the group. However, when the differences
between species are taken into account, it seems evident that
natural selection has had little to do with the evolution of species.
For example, it is hard to imagine that the difference between the
muricate and rugose nutlets would have any survival value.
Even the wings on such nutlets as C. confertiflora or С. setosissima
scarcely seem to be large or efficient enough to be of real value to
the species. Тһе distribution of pustulate hairs on the surfaces
of the leaves can hardly be supposed to determine whether a
species will survive or not. Even though specific differentiation
in Oreocarya may be non-adaptive yet it is quite possible that
natural selection has been constantly operative in eliminating
many evolving forms that were not nicely adjusted to their sur-
roundings.

Geographical isolation.—The author is inclined to believe that
geographical isolation has been one of the chief factors in speci-
ation. The way in which species conform to natural areas in
their distribution, the consistency with which closely related and
incipient species occupy adjoining areas, and the trivial characters
that distinguish species and varieties, all seem to argue in favor

1927] :
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 221

of isolation, and in the present case mainly geographical isolation,
as being one of the most important evolutionary factors.

'The subject of Age and Area has been much discussed recently.
The author can see no indication that there is any correlation
between these two factors in the present group of species. The
species that is considered most primitive, C. Jamesii, occupies the
largest area of any species in the section. However, C. flavocu-
lata, which is certainly one of the most specialized of all, occupies
an area that is much larger than that occupied by most species.
Although the comparative ages of the species have not been
established, it might be expected that there is a close positive
correlation between age and degree of specialization.

Orthogenesis.—The frequency with which orthogenetic ex-
planations lend themselves to phylogenetic description seems to
be impressing itself more and more upon those who are interested
in evolutionary theories. Orthogenesis is a phenomenon of
evolution and no explanation. There is, of necessity, nothing
teleological about this mode of development. In the present
study one very interesting tendency that may be considered or-
thogenetic seems to be in operation. This is the tendency for
long-lived ancestral species to produce short-lived descendant
species. In the section Oreocarya this tendency has certainly
occurred several times independently although the annual habit
has never been attained. The sections Eucryptantha and Geo-
carya exhibit the same tendency to reduce the length of life.
Therefore, in looking over the genus as a whole it can be said
that again and again long-lived species have given rise to shorter-
lived ones. Is this an inherently orthogenetic tendency or is it
to be interpreted merely as a general response to wide-spread
conditions of a common habitat? The only approach to an
answer that the author can give to this question at the present
time is to call attention to the phylogeny of Lesquerella. Here
is a genus of comparable size to the section Oreocarya that occupies
an area, in North and South America, where very similar habitats
prevail. In Lesquerella, the tendency that appears again and
again as the species evolve is to increase the duration of life. In
similar, and often identical, habitats two unrelated genera do
opposite things: Cryptantha produces annuals or biennials again

[Vor. 14
222 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and again, while Lesquerella produces perennials time after time.
And yet the tendency is not absolute. At least one species in
Lesquerella, L. Cusickii, has certainly been produced from peren-
nial рагепіз,! and it seems quite probable that in South America
Cryptantha argentea and C. amplexicaulis have developed from
annual parents.

GEOGRAPHICAL DISTRIBUTION

A careful study of geographical distribution is at once one of the
most interesting and necessary parts of any taxonomic or phylo-
genetic monograph. It is of great interest because species must
evolve in some relation to their distribution and theories of iso-
lation are becoming of greater and greater importance. Geo-
graphical distribution is of great practical importance in deter-
mination of species, since closely related forms rarely exist in the
same area.

In a genus previously studied, i.e., Lesquerella of the Cruciferae,
there was a most interesting direct relationship between geo-
graphical distribution and phylogeny. In this case the primitive
species are located rather near the center of distribution and the
farther any species occurs from this central region the more likely
is it to be specialized. In the present group of species, this re-
lationship seems not to hold—or at least not to be conspicuous.
The only explanation that suggests itself at present is that the
section Oreocarya is too old to show such a relationship. In this
connection it would be most interesting to study the distribution
of the more specialized sections of the genus in order to discover
if other relationships exist there.

A study of a map on which are plotted all the specific ranges
reveals the fact that the center of distribution is in western Colo-
rado and eastern Utah. A conspicuous area of concentration
extends from northeast to southwest, from southern Wyoming to
southern Nevada.

If the distribution of the genus Cryptantha as a whole is ex-
amined it is seen that the primitive section, Oreocarya, is confined
(if we neglect one or two possible South American species) to

‘See Payson, E. B. A monograph of the genus Lesquerella. Ann. Mo. Bot.
Gard. 8: 104. 1921.

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 223

North America, and the most primitive species is in the southern
part of the sectional range. Тһе other sections have their most
primitive species in South America. It would seem reasonable
to suppose that the genus originated in the southwestern part of
North America and in its perennial primitive form spread to the
desert regions of Chile and Argentina. In that region it became
differentiated into three main groups. Тһе most primitive of
these invaded North America as Cryptantha in the limited sense
of recent American floras. Тһе other somewhat more specialized
sections have not spread beyond South America. Before any
confidence could be placed in such a theory the other sections of
the genus would have to be studied very carefully.

THE SPECIFIC CONCEPT

The attempt has been made in the present study to adopt a
specific concept that is intermediate between extremes of segre-
gation and of aggregation although it is recognized that the
species are not of uniform value. In general it has been the
policy to accept as a species any unit that could be distinguished
from related groups by two or more constant differences of sig-
nificant value, provided that intermediates were lacking between
the groups, or that these intermediates were so few as to be con-
sidered relatively unimportant. If the differences between dif-
ferent groups are of very small size, or if numerous intermediates
exist, then the units concerned are best treated as varieties.

A conservative attitude toward proposed species has been
maintained in case the evidence at hand is rather meager. This
has led to the retention of certain units as species when possibly
they will eventually be accorded varietal rank only.

The author is firmly convinced that all the units that are here
accorded taxonomic recognition are real genetic units of consider-
able size and distinctness. Further study may shift the status
of the various groups but this will be only of minor importance.

The device given below has been employed to illustrate the
specific concept and, at the same time, to indicate something of
the relative value of the different units. It is to be understood
that names associated in certain positions in this outline are not
indications of new nomenclatorial combinations.

224

[Vor. 14

ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

Species and subordinate
groups that author
would recognize if a broad-

P ew and subordinate
Oups recognized in
o present study

Species and блесна
groups аз they might
recognized if Pig specific

er specific concept were ой were smaller and
employed the peer of intermedi-
ates igno
Jamesit Jamesit
multicaulis multicaulis multicaulis
laxa laxa
cinerea cinerea
disticha disticha
/атевй
pica typica
abortiva abortiva abortiva
pustulosa pustulosa pustulosa
Palme almeri Palmeri
ta oblata oblata
leucophaea
confertiflora confertiflora confertiflora
flava flava
typica leucophaea leucophaea
salmonensis salmonensis salmonensis
icta stricta stricta
? nubigena nubige nubigena
Clem Clemensae Clemensae
setosissima setosissima setosissima
virgata virgata virgata
humilis
insolita insolita insolita
virginensis virginensis virginensis
tumulosa tumulosa tumulosa
modesta esta odest
typica humilis humilis
? nubigena
caespitosa caespitosa pitosa
thyrsiflora thyrsiflora thyrsiflora
? elata elata elata
sericea sericea
typica typica
perennis perennis
ta aperta aperta
Bradburiana
elata
? aperta
rugulosa rugulosa rugulosa
interrupta interrupta interrupta
spiculifera spiculifera spieulifera
celosioides celosioides
Sheldonii Sheldonii тус е
acountt Macounii
sobolifera sobolifera sobolifera
са Bradburiana Bradburiana
nana
commixta comm commixta
Shanizii Shantzii Shantzii
ovina ovina
typica typica nana
cana cana cana
propria propria propria

1927]

PAYSON—SECTION OREOCARYA OF CRYPTANTHA

225

Species and subordinate
groups that the author
would recognize if а broad-
er specific concept were
employed

рое апа чта aes
oups as recognized i
the present study

E

Species and qa ng
groups as they might

cept we
the pressnos eot intermedi-
ates ignore

ее»

reviflora
typica
echinoides
Jonesiana

ongiflora
Wetherillii
typica

Osterhoutii

breviflora
fulvocanescens
echinoides
Wetherillii
те flora

Otorhoutét
ратадота

mensana
flavoculata

breviflora

J onesiana
Wetherillit
longiflora
tenuis
Osterhoutii
akeri

mensana
Ж

? Shockleyi

TAXONOMIC CHARACTERS OF SPECIFIC VALUE IN THE SECTION
OREOCAR

Nutlets—Nutlet differences are remarkably constant between

individuals of any species in this group. With rather few ex-
ceptions, it is possible to determine any species by means of а
mature nutlet. Differences in the character of the surface are
most important, although size, form of the scar, and outline are
usually characteristic. Were the nutlet peculiarities the only
differences that exist, the species would be doubtfully tenable as
species. However, when the plants are well known certain other
characters are evident that are correlated with nutlet characters
as well as with geographical distribution.

Corolla.—Certain differences in the corolla are of great im-
portance. The relative length of the corolla tube and calyx lobes
is very constant for any given species. "Тһе form of the fornices,
the presence or absence of crests at the base of the tube are valu-
able diagnostic characters. Two species have yellow corollas
while all others have white ones. In a few the corolla tube is
In most species the fornices are distinctly yellow.
The
amount of elongation of the lobes after anthesis is comparatively

[Vor. 14

226 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

)
/ г
f 2
К Е
\ Й f
2
І В ш Е

Fig. 1. Comparison of floral and fruit details of four species of Cryptantha: I. C.
virginensis, section Oreocarya; II. C. capituliflora, section Eucryptantha; ТП. C. Kingii,
section Geocarya; IV. C. ambigua, section Krynitzkia.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 227

small in а few primitive species. Certain species have sepals
that are nearly linear and in others they are fairly broad.

Style.—The length of the style, as well as the position of the
stamens in the corolla tube, is of little specific value in the long-
flowered species, since most of them are distinctly dimorphic in
these characters. In the short-flowered species, however, the
distance that the style exceeds the mature nutlets is of some
value.

Indument.—Some species are characteristically setose and
others conspicuously sericeous but most of the different forms are
so similar in general hairiness that the differences are unnoticed
by any one unfamiliar with the group. A few species are nicely
characterized by the absence of pustulate hairs on the upper leaf
surface, and one is peculiar in being nearly glabrous.

Inflorescence.—Differences in inflorescence are difficult of de-
scription. Most of the species are similar with respect to the
manner іп which the flowers are borne. А few have an unusual
appearance because of the regular and extreme elongation of the
cymes. One, C. virgata, is unique in the possession of elongated
foliar bracts in an otherwise narrow inflorescence. Most of the
species with a narrow inflorescence have a tendency to elongate
the branches in age and so come to resemble those species with
characteristically broad inflorescence.

Persistence.—A few of the species are evidently biennial and
some are especially long lived. In general, differences in duration
are not easy of definition and interpretation, and so these char-
acters are only of exceptional value.

Leaves.—Leaf size and shape vary considerably between in-
dividuals and are ordinarily of little value in identification.

Stem.—Most of the species are similar with respect to the
characters of the stem. A few are unusually tall and some are
very low and caespitose.

Gynobase.—This structure is difficult of examination and in
only a few species is it of diagnostic value. А series of careful
drawings of the gynobase was made and these are available in
the Rocky Mountain Herbarium.

[Vor. 14
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN

THE GENERIC CONCEPT

One of the most important results of the present study has
developed from a suggestion made to the author early in 1924 by
Dr. Ivan Johnston of the Gray Herbarium. This was to the
effect that Oreocarya could only with difficulty be maintained as
a genus separate from Cryptantha. Since that time Dr. Johnston
has continued his studies of the genera of this family and has
become convinced that his earlier conjecture was correct. He
has studied and collected plants of this relationship in South
America as well as in western North America and has accumulated
much evidence bearing on this point. Through the kindness of
Dr. Johnston and of Dr. B. L. Robinson of the Gray Herbarium
the author has been permitted to study a considerable series of
South American specimens. This study, although much more
superficial than that given to it by Dr. Johnston, has served to
confirm his conclusions and to convince the author that as a
genus Oreocarya can not be separated from the older Cryptantha.

In this enlarged sense, Cryptantha consists of about 150 species
of the arid regions of western North and South America. These
species are annual, biennial, perennial or even fruticose herbs
with linear, oblanceolate, spatulate or nearly obovate leaves.
Most of the species are clothed with a conspicuous indument of
unbranched trichomes and all possess some enlarged epidermal
pustules. The inflorescence consists of variously elongating,
2-ranked, scorpioid cymes. The sepals are persistent and
usually somewhat accrescent in fruit. The corolla is salverform
and white, white with yellow fornices, or entirely yellow in color.
The fornices are always present at the throat of the corolla tube;
a circle of crests at the base of the tube is present or absent.
Nutlets are homomorphous or heteromorphous, one to four
maturing, straight or nearly so, attached laterally to a subulate
gynobase, smooth or variously roughened.

The considerations that have led the author to submerge
Oreocarya into Cryptantha may be summarized as follows:

FUNDAMENTAL PREMISES

1. Genera can not be maintained that are only provincially
distinct. In other words, genera should not be recognized in

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 229

one part of the world if they are not satisfactorily distinct in
another.

2. It is admitted that genera are partly limited by a consider-
ation of matters of convenience but fundamentally there must be
some morphological character or characters on which to base
genera.

3. If genera are based entirely on considerations of convenience,
then taxonomic groupings will become so artificial and distorted
as to furnish no sound basis for geographic or phylogenetic gener-
alizations and taxonomy will become no more than a system of
cataloguing.

4. Habital differences only may be of great value in specific or
varietal distinctions but are not usually of generic value. Fur-
thermore it is easy to say that this group differs from that in
aspect but if this difference in aspect is not capable of morpho-
logical definition it can be accorded very little weight.

IMMEDIATE CONSIDERATIONS

5. Тһе species of Cryptantha in South America, maintained by
Johnston as valid, are about 40 in number. "These include forms
with considerable diversity of structure. "There are forms with
cleistogamous flowers of various kinds, others with only chas-
mogamic ones. There are fruticose, persistent, and annual species.
However, the intermediates are so evident between the various
categories that there seems no possibility of separating them
except specifically. In South America, then, we have one genus.

6. It seems quite impossible to separate the annual North
American Cryptanthas from the annual South American species
except specifically. Some forms from the two continents are so
similar that they could even be confused as belonging to the same
species.

7. 'There are in South America a few species which in North
America would unhesitatingly have been referred to Oreocarya
because of floral and fruit characters as well as characteristic
habit and aspect. These species are not aberrant Cryptanthas
in South America. |

8. The author has found it impossible to locate а single mor-
phological character that will definitely separate Oreocarya and

[Vor. 14
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Cryptantha when the South American species are taken into
account. Even in North America the only character that has
been found that will separate the two groups is the biennial or
perennial root in Oreocarya and the annual one in Cryptantha.

9. From a standpoint of geographical distribution it is not re-
markable to find a single genus common to western North America
and western South America in Chile and Argentina. Many
examples might be given: Lesquerella mendocina of northern
Argentina, adjacent Chile and Bolivia is so similar to certain
Rocky Mountain species as to make determination difficult if the
labels are concealed; Coldenia Nuttallii is common to both conti-
nents with an enormous gap between (see Johnston, Contr. Gray
Herb. 75: 43-44. 1925); Alamisquaea emarginata is common to
Lower California and to Chile.

If only the North American species were concerned, the author
would keep the genera Oreocarya and Cryptantha as distinct on
the basis of annual as opposed to perennial or biennial herbs. In
addition, the inflorescence of Cryptantha is usually elongated and
bractless while that of Oreocarya is usually contracted and brac-
teate. There аге, however, many exceptions to these distinctions.
The nutlets in the North American Cryptanthas show a strong
tendency to definite heteromorphism, while in Oreocarya no
such tendency has been noted. There are North American
Cryptanthas with homomorphous nutlets. Іп some species of
Cryptantha the calyx tends to fall as a unit and enclose the per-
sistent nutlets. This has been given as a character distinguishing
Отеосатуа from Cryptantha but is not satisfactory, since many
Cryptanthas do not shed their calyces and in one Oreocarya, at
least, the pedicels have a tendency to separate from the stem
and so drop the calyx with the persistent nutlets. This is C. nana
var. commixta. Taking the South American species into con-
sideration then, the one definite character separating Cryptantha
from Отеосатуа disappears entirely, and the union of the two
groups becomes a necessity if the premises given above are
worthy of acceptance.

The unfortunate result of uniting the Oreocarya group with the
Cryptanthas comes because of the necessary shift in names.
'This seems unavoidable and is in the direction of simplification

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 231

rather than of complication. There has been much demand of
late, on the part of taxonomists as well as non-taxonomists, for
larger generic concepts and a consequent reduction in generic
names. And it therefore happens that the change necessary in
the present case is in line with tendencies that cannot be ignored.

Dr. Johnston (Contr. Gray Herb. 74: 1-114. 1925) has re-
viewed the historical development of the genus Cryptantha (ex-
clusive of Oreocarya) and has discussed the question of segregation
and available generic names. This material need not be ге-
peated here. Suffice it to say that the author accepts this treat-
ment of the genus in North America as regards generic concept
but would now add to it Oreocarya.

Taxonomic HISTORY OF THE SECTION OREOCARYA

The first species belonging to this group to be described was
C. Bradburiana. This was collected by Bradbury in ‘“ Upper
Louisiana" in 1810 and described by Pursh as a species of
Cynoglossum in 1814. In 1818 it was transferred to Myosotis by
Nuttall. In 1828, a second species was described (C. Jamesii) and
assigned to Myosotis by Torrey. Douglas described the third
species (C. leucophaea) in 1830 and also placed it in Myosotis.

De Candolle іп the ‘Prodromus’ transferred Bradburiana and
leucophaea to the genus Eritrichium in 1846 and so started а
treatment that persisted until 1885. During this period four
species and two varieties were described in the group. Тһе most
important publication affecting the genus was by Gray in Proc.
Am. Acad. 10: 61. 1875. Аза part of a revision of Eritrichium
Gray grouped the known species and varieties together under the
section Pseudo-M yosotis which had been proposed by A. De
Candolle in the ‘Prodromus’ for C. leucophaea. Four species
and three varieties were recognized as valid in this revision.

The next important treatment of this genus was by Gray in
Proc. Am. Acad. 20: 276-280. 1885. Тһе species are here
transferred to the section Pseudokrynitzkia of Krynitzkia with the
exception of C. setosissima which was placed in the section
Pterygium. The group was still held essentially as a unit and
eight species were recognized that are now placed in the section
Oreocarya.

[Vor. 14
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Тһе genus Oreocarya was proposed by Greene in 1887 (Pittonia
1: 57-58) and made to contain nine species—eight of which are
now held in this section. It is evident that this paper of Greene's
is concerned with the question of the generic relationship of
certain Boraginaceous plants rather than with specific deter-
minations, since the species are those recognized by Gray as of
this relationship in 1885. In 1896 Greene again published on
this genus, and this time it was to describe eight new species and
to redescribe several old ones. With this paper, Greene's most
important contributions to the study of this genus end. In 1899
he described two new species and in 1901 two more.

The period from 1896-1915 is chiefly characterized by the ap-
pearance in literature of some thirty-five species proposed as new.
These were proposed by a number of workers on the flora of the
Rocky Mountains and the Great Basin, and except for those
described by M. E. Jones were all assigned to the genus Oreocarya.
The most active in this work of characterizing new species were
Alice Eastwood, Aven Nelson, P. A. Rydberg, and M. E. Jones.
Іп 1906 Rydberg published his ‘Flora of Colorado’ in which he
recognized nineteen species as occurring in Colorado alone, and
in 1909 Aven Nelson in the Coulter-Nelson ‘Manual of the Flora
of the Central Rocky Mountains,’ maintained the same number of
valid species for the whole region covered by the ‘Manual.’

The first comprehensive revision of the group as included in
the genus Oreocarya was published by J. F. Macbride in Contr.
Gray Herb. 48: 20-38. 1916. In this very excellent study forty-
five valid species are maintained and differentiated by means of
a dichotomous key. Specimens are cited under each specific
name and comments are made relative to the different species.
In 1918 Rydberg published a treatment of the species of the Rocky
Mountain region in his ‘Flora of the Rocky Mountains and Ad-
jacent Plains.’ In this he follows Macbride very largely but
recognizes a few species that Macbride had not accorded specific
rank. There are a few extra-limital species that Rydberg does
not consider. However, he recognizes thirty-nine species in the
range of his flora.

The history of Oreocarya subsequent to the important work
done by Macbride in 1916 has been comparatively unimportant.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 233

New species have been described from time to time by Osterhout,
Brand, and Payson. The generic acceptance of Oreocarya has
been almost universal since the genus was proposed until the
present time. М. Е. Jones and К. Brandegee seem to have been
the only recent taxonomists who have refused to accept the
generic separation of the perennial American forms (Oreocarya)
from the annual species (Cryptantha or Krynitzkia). Their re-
fusal to accept this segregation, it is of interest to note, was not
based upon the most convincing evidence, i.e. the South American
material, but upon the similarity of the North American species.

What the author believes to be the beginning of a new concept
in the taxonomic treatment of the plants under consideration
was made by Dr. Johnston in Contr. Gray Herb. 70: 46. 1924,
when he questioned the generic value of the Oreocarya group.

In summary, the history of this group may be said to reflect
the history of taxonomy in America. First we notice the great
increase in the number of described species. From two in 1828
the group has grown to eight in 1887, sixteen in 1896, sixty-two
in 1916, and seventy-five in 1926. This increase shows two great
factors at work. First is the great activity in exploration and
description of species from the desert regions of western North
America. Second is a tendency to reduce the value of a species.
The generic shifts in the position of the species under consider-
ation reflect another phase of taxonomic history. In the earlier
treatments the species were placed under older genera to which
they were not closely related. These associations proved un-
satisfactory and so the species shifted until in Krynitzkia Dr.
Gray thought he had associated them with their relatives—and
so we think to-day. The author thinks there is no doubt but
that Dr. Gray would have united Krynitzkia with Cryptantha if
he had known the South American species as we now know them.
This earlier period of taxonomic activity—ending with the work
of Gray and Watson—was characterized by synthetic studies.
Genera were revised as a whole and a great effort was made to
arrange the groups so they might stand in the proper relationship
to one another. Following this early period came a time of great
expansion of population in the ited States, and this was fol-
lowed by a flood of floristic works on the plants of the regions so

[Уог. 14
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN

newly occupied and colonized. Floras and manuals are notori-
ously provincial in their generic and specific concepts and so a
great impetus was given to segregation of genera and species.
Units were maintained in one region that could not be separated
in another and there resulted a great confusion in the taxonomy
of the plants of western America. This period of regional floras
rather than of monographic work on whole genera or families
seems now to be giving way to a period of synthetic and even
experimental taxonomic work. With the great activity in the
botanical exploration of South America, botanists must come
to recognize that a local view of a genus is not sufficient and that
before species or genera can be successfully characterized they
must be seen in relation to all of their relatives. The union of
Oreocarya and Cryptantha is a result of a larger systematic view-
point and seems inevitable when such an enlarged view is ob-
tained.

ILLUSTRATIONS

Since the nutlets present characters of the greatest taxonomic
value, and since they are difficult of adequate description, it was
thought worth while to illustrate all of those species in which the
nutlets are significantly different. These drawings were made in
pencil from the nutlets as seen through a binocular dissecting
microscope giving a magnification of thirty diameters. Each
nutlet is shown in three positions, and all were drawn twenty
times the actual size. In reproduction this enlargement was
reduced one-half. All the illustrations were prepared by the
author.

ACKNOWLEDGEMENTS

The author is under great obligations to many people for as-
sistance in the present work. Any taxonomic treatment borrows
much from earlier workers. Sources of this kind are indicated
in a bibliography. In the present case the revision by Mr. J. F.
Macbride has been of the greatest value. That was the pioneer
work that assembled the materials and made the present study
much less difficult than it would otherwise have been. To Dr.
Ivan Johnston and Dr. A. Brand the author is much indebted for

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 235

suggestions concerning the plants under consideration. Both of
these workers are at present busied with studies in the Boragin-
aceae. Dr. Aven Nelson, Curator of the Rocky Mountain
Herbarium, has done much to encourage and assist the author in
many ways.

Herbarium material has been borrowed from a number of
herbaria. Some collections have been visited by the author in
person. То the curators of these various herbaria the author
wishes to express his gratitude.

To Dr. George T. Moore, Director of the Missouri Botanical
Garden, and to Dr. J. M. Greenman, Curator of the Herbarium
at that institution, the author is under especial obligations for
making possible the publication of this paper.

Specimens have been examined from the following herbaria,
the abbreviations in parentheses being those used in the citation
of specimens:

1. Colorado State Museum (Colo.).

2. Field Museum (Field).

3. Gray Herbarium (Gray).

4. Missouri Botanical Garden (Mo.).

5. Montana State College (Mont.).

6. New York Botanical Garden (N. Y.).

7. George E. Osterhout Herbarium (Osterh.).

8. Philadelphia Academy of Natural Sciences (Phila.).
9. Pomona College (Pomona).
10. Rocky Mountain Herbarium (R.Mt.).

11. United States National Herbarium (U.S.).
12. University of California (Calif.).

13. Victoria Memorial Museum, Ottawa (Canada).
14. Washington State College (Wash.).

TAXONOMY

Dr. Ivan Johnston would divide the genus Cryptantha into
three sections based upon the character of the flowers. To these
three, the present author would add a fourth by dividing one of
those maintained by Dr. Johnston. This classification may be
indicated as follows:

[Vor. 14
236 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Plant n only chasmogamic flowers
nial or biennial herbs, native to North America....Section I. Oreocarya.
inen (or rarely perennial) herbs, native to North sod South America
О irae Жаы о СТРУС eei Section II. Krynitzkia.
Plant producing some cleistogamic flowers.
Cleistogamic flowers not highly specialized, differing from the chasmogamic
ones merely in the closed corolla; borne in the axils of не cauline leaves

and frequently i in the spikes above.............. Section. ПТ. Eueryptantha.
СІ ly specialized, becoming iau ықы borne
at the base of the stem ИЕ ee ТОРЕ Section IV. Geocarya.

The author proposed to retain Oreocarya as a section of Cryp-
tantha for the following reasons:

1. The group of species that has been treated as comprising

the genus Oreocarya is a fairly homogeneous unit rather
easily separable from the other sections of the genus.
That it is not entirely and consistently separable from the
other sections is an argument for its reduction from generic
rank but is not a reason for failing to recognize it as a section.
As a matter of convenience, it is desirable to retain a name
that has been so completely incorporated into literature as
has Oreocarya. Since it can not be maintained as a genus,
it is of some value to retain it in subgeneric rank.

In the present treatment the section Oreocarya is limited to the
North American species. It is recognized that this is a more or
less arbitrary division, and it might be argued that the separation
would be better made to include all the perennial species without
cleistogamic flowers. This would result in including at least
one South American species, С. gnaphalioides, in the section
Oreocarya. Two others might also be included, namely C. argen-
tea and C. amplexicaulis. These two last ones are apparently
more nearly related to some of the annual rather than to the
perennial species and so would introduce an anomalous element
into the section. The author believes that because the sectional
separation must be made more or less arbitrarily, since no distinct
line of cleavage exists, it is well to make it conform to distinct
geographical lines. This seems particularly desirable since it is
probable that even С. gnaphalioides, which species has the best
right to be included in Oreocarya of all the South American forms,
is probably more closely related to the annual Krynitzkias than
to the perennial North American Oreocaryas. If it were certain

ho

оз?

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 237

that a perfectly phylogenetie classification would result from
dividing the sections strictly according to morphological char-
acters, then there would be no question as to the preferable
procedure. Аз it is, the sections are convenient, mainly mono-
phyletie, groups, and the question of expediency has a right to
consideration.

CRYPTANTHA section OREOCARYA (Greene) Payson new comb.

Eritrichium, section Pseudomyosotis A.DC. Prodr. 10: 129.
1846; Gray, Proc. Am. Acad. 10: 61. 1875; Benth. & Hook.
Gen. Pl. 2:850. 1876; Gray, Syn. Fl. №. Am. 2::196-197. 1878.

Krynitzkia, section Pterygium Gray, Proc. Ат. Acad. 20: 276.
1885, in part.

Krynitzkia, section Pseudokrynitzkia Gray, Proc. Am. Acad.
20: 276-280. 1885, in large part; Coulter, Manual Rocky Mt.
Region, 164. 1885.

Oreocarya Greene, Pittonia 1:57-58. 1887; Ibid. 3: 109-115.
1896; Gürke in Engl. & Prantl, Nat. Pflanzenfam. IV. Abt. 3a &
3b. 109. 1897; A. Nelson, Erythea 7: 65-68. 1899; Howell,
Fl. Nw. Am. 486. 1900; Eastwood, Bull. Torr. Bot. Club 30:
238—246. 1903; Britton, Manual, ed. 2, 770. 1905; Piper, Contr.
U. S. Nat. Herb. 11: 481-482. 1906; Rydb. Fl. Colo. 287-288.
1906; A. Nelson in Coulter & Nelson, Man. Cent. Rocky Mts.
416-419. 1909; Britton & Brown, Ill. FL, ed. 2, 3: 80. 1913;
Wooton & Standley, Contr. U. S. Nat. Herb. 19: 544-546. 1915;
Macbride, Contr. Gray Herb. 48: 20-38. 1916; Rydb. Fl. Rocky
Mts. 719-725. 1917; Tidestrom, Contr. U. S. Nat. Herb. 25:
457-460. 1925; Payson, Univ. Wyo. Publ. Bot. 1: 164-171.
1926. |

Perennial or biennial herbs with a conspicuous setose, hirsute
or sericeous indument (except in C. pustulosa). Leaves entire,
oblanceolate, spatulate or linear. Stems solitary from the root
or caespitose, commonly unbranched below the inflorescence,
0.5-9 dm. high. Inflorescence a continuous or glomerate cluster
of elongating or reduced, simple or branched, bracteate or nearly |
ebracteate, two-ranked, unilateral, scorpioid cymes. Sepals
distinct, usually conspicuously accrescent. Corollas white or
yellow, salverform; limb 4-12 mm. broad; tube equalling or ex-

[Vor. 14
238 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ceeding the sepals. Stamens included in the tube, anthers sessile
or nearly so. Style equalling or much exceeding the mature
nutlets; stigma entire. Nutlets from nearly circular in outline to
narrowly lanceolate, margined or winged, smooth or variously
roughened on the different surfaces, attached to the gynobase at
a point !/,-*/, of the distance from the base to the apex of the
nutlet. Scar of nutlets various, open or closed and margin
elevated or plane. Generic type: C. Bradburiana Payson.

ARTIFICIAL КЕҮ TO THE SPECIES OF CRYPTANTHA,
SECTION OREOCARYA

a. Tube of the corolla distinctly longer than the calyx lobes in anthesis.
b. Nutlets smooth on the dorsal and ventral surfaces; leaves linear-oblan-
ceolate, usually acute, densely appressed-strigose, petioles ciliate.

с. Corollas yellow, individuals distinctly dimorphic as regards stamen
insertion; species Tanging from central Wyoming south and west to
southern California
d. Nutlets broadly ovate in outline, distinctly wing-margined, all

four usually maturing; inflorescence distinctly glomerate; native
from Utah to southern California.................. 5. C. confertiflora
. Nutlets lanceolate in outline, scarcely wing-margined, some usu-
ally aborting; inflorescence scarcely glomerate; native from central
Wyoming through western Colorado to northern New Mexico wn
eastern Utah AIRES A Е С. Лаға
сс. Flowers wl lividuals 1 idently dimorphi regards 44.
insertion; native to Washington and (?) Oregon УЕ 7. C. leucophaea
bb. Nutlets more or less roughened or wrinkled on dorsal surfaces; leaves
various.
с. Nutlets uniformly muricate or рарШове, not evidently tuberculate,
rugose or wrin
d. Leaves silky-strigose, pustulate hairs small or lacking; fornices
elongated
e. Muriculations usually acute and sometimes minutely setose;
eastern Utah, c Colorado to northern New € and
northeastern Агізопа......................... 36. C. fulvocanescens
ee. Muriculations ақы setose, sometimes almost arbor-
escent; southern and southwestern Utah and adjacent Arizona.
37. C. echinoides
dd. Leaves appressed-setose and coarsely strigose, conspicuously
pustulate on both leaf surfaces; fornices low and rounded
viu оо о у о г 38. C. J'onesiana
cc. Nutlets more or less rugose or tuberculate, sometimes conspicuously
muricate also.
d. Inner or ventral surfaces of the nutlets smooth or nearly so; margin
evidently but rather narrowly wing-margined.............. 4. C. oblata

с.
Qa

ee ee ee ee ee ee ee ee a oe ee ae oe er ee % е 69%%%%8%%%

1927]
PAYSON—SECTION OREOCARYA ОЕ СКУРТАМТНА 239

dd. Inner surfaces of the nutlets distinctly nd
e. Nutlets conspicuously muricate, some of these confluent into
rugae; scar narrow, straight, open, no bebes of an elevated
ОА РР РИ. 38. C. Jonesiana
ee. Жайка rugose ог tubereulate, these more conspicuous than the
murications; scar various, margin distinctly elevated in some

species.

f. Margins of the nutlets not in contact, fruit oblate-ovoid;
densely caespitose perennials; pustulate hairs lacking on upper
ау... ПИ СТРОИТЕ: 43. С. ратадоха

ff. Margins of the nutlets in contact.

g. бсаг of the nutlets i-a open and surrounded by а
definite elevated гоагріп.................... 46. C. flavoculata
gg. Scar of the nutlets died or slightly open but then not sur-
rounded by a definitely limited and conspicuous elevated
margin.
h. Leaves sparsely if at all pustulate ventrally, even in age.
i. Scar of the nutlets surrounded by an elevated margin
but tightly closed, leaves oblanceolate or broader. 44. C. Bakeri
ii. Scar neither tightly closed nor surrounded by an evident
elevated margin.
j Leaves narrowly linear-spatulate, nutlets sharply
and deeply тир Leer rr 41. C. tenuis
jj. Leaves obovate or broadly oblanceolate; nutlets with
rounded rugae and tubercles; fornices он е”
К. ТЕ . C. Майей я
hh. Leaves conspicuously pustulate ventrally, т їп
rolla tube 1 ста. or more long; Colorado. .40. С. longiflora
aa. Tube of the corolla equalling or shorter than the calyx lobes in anthesis.
b. Nutlets smooth on the dorsal surfaces, not rugose, muricate or tuber-
ulate

с у
c. Fruit depressed, globular, nutlets not in contact by their margins.
d. Leaves conspicuously pubescent on both surfaces.
e. Crests at base E corolla tube conspicuous; calyx not conspicu-
1. C. Ja

ously accrescent....... оне ева ео nehm mesit

ee. > өлен T base ۴ corolla tube obsolete; calyx MM der Y. ас-
раз она а ds rm o . C. Palmeri
dd. Шы B above, sparingly hairy below........... Ж с. pustulosa

се. Fruit conical or ovoid, nutlets in contact by their margins.
d. Stout, peny erect plants with many elongated and conspicuous
n the inflorescence; eastern Colorado and southeastern
io) eesosa O o. 00D. 13. C. virgata
dd. Smaller, somewhat caespitose plants with few or inconspicuous
racts in the inflorescence; central Idaho............ е. а
bb. Nutlets more ог less roughened (muricate, wrinkled, йан, at
least on the dorsal surfaces.
c. Nutlets densely and uniformly muricate, not rugose or tuberculate.
d. Pubescence of leaves silky-strigose or strigillose but not sub-
tomentose.

240

[Vor. 14
ANNALS OF THE MISSOURI BOTANICAL GARDEN

e. Leaves nearly or quite uniformly silky-strigose.
f. Plants scarcely or only moderately caespitose; leaves broadly
oblanceolate or spatulate; native to northeastern Utah and ad-
jJücent Colorado (T)... cc eran о acea Шы $5. C. breviflora
ff. Plants densely caespitose, caudex multicipital; leaves linear-
oblanceolate; native to eastern Wyoming and adjacent Ne-
кетир and Colorado

КҮТУІ PUO FM Ие: 33. C. cana
with two distinctly different kinds of trichomes....32. C. nana
dd. 5. устай subtomentose аз well as appressedackdas .82. С. nana
ec. Ми е{з not exclusively muricate.
d. Inner surfaces of the mature nutlets smooth or nearly so, not con-
spicuously rugose, tuberculate or muricate.

e. Nutlets ovate, papery with a broad, thin wing-margin; stout
erect plants 2.5-8 dm. tall; native to Utah and Arizona
кезде т OTR TERT eT Ae re esi ELE EE RM . C. setosissima

ee. Nutlets ovate to lanceolate, not with a broad, thin wing-margin
if the nutlets are ovate in outline
f. Stout, strictly erect plants with many elongated and con-
spicuous foliar bracts in the spiciform inflorescence....13. C. virgata
ff. Lower, more slender plants; inflorescence not spiciform nor

with numerous elongated foliar bracts that greatly exceed the

flowers

g. Inflorescence very broad and rounded in outline; nutlets not
wing-margined; native to the eastern slope of the conti-
nental divide from Wyoming to central New ее”
И АЛ А бөзі ҚҰЛАҚ va тата ES . С. thyrsiflora

ЕЕ. Inflorescence narrower; nutlets usually with an bris
wing-margin (except in C. rugulosa); native west of the con-
tinental divide.
h. Nutlets with evident transverse rugae, pri more con-
жен than the tubercles that тау be
Ee scarcely or not at all muricate а the

№ үа егесі, conspicuously веїозе — of
d

о’ a ТТТ ТТТ” . C. stricta
jj. Caespitose, | i егеппі f high
elevation i in Idaho, Nevada, д екінің ала —

ИАА . С. nubigena
. Nutlets distinetly muricate between the rugae ЖР near
the margins of the nutlets.

j. Erect perennials from western Utah........ 24. C. rugulosa

jj. Caespitose and more or less soboliferous perenni
from Montan

ials
UICE TNT RETURN ELE . C. sobolifera
hh. Nutlets with distant tubercles - no инон rugae;

native of the southern Sierra Nevada........ lemensae
dd. Inner surfaces of the be кыйа ан. and definitely 700086,
tuberculate ог muricate.
e. Scar of the nutlets evidently open some distance above the base.
f. Scar somewhat constricted some distance below the middle of
the open portion.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 241

g. Elevated margin of the scar definitely limited; pustules
present on both Іеаѓ-виѓасез................... . C. mensana

gg. Elevated margin indefinitely limited; pustules кедені, only
on the lower surface of the leaves........... . С. Osterhoutii

ff. Scar triangular and not constricted below the midi.

. Some tendency to elevated margin evident around the scar.

h. Cymules elongating and so the inflorescence broad; bien-

nial or short-lived perennials; nutlets with an evident

dorsal ridge.

i. Surfaces of the leaves conspicuously setose with spread-
ing bristles; inflorescence crowded. ......... 15. C. virginensis

ii. Surfaces of the leaves with inconspicuous appressed
bristles; inflorescence ореп................... 14. C. insolita

hh. Cymules shorter and ры narrow; long-lived

‚ perennials; nutlets with only a slight dorsal ridge, if any.

i. Nutlets indefinitely tuberculate апа rugose; California

16

Е NE UU вене вех оа . C. tumulosa
ii Nutlets definitely tuberculate or rugose; Utah and
eastern Nevada. С 9n адь 17. C. modesta

gg. No tendency to an elevated margin around the scar.
h. Style exceeding the mature nutlets by 1 mm. or more.
i. Leaves finely strigose and appressed-setulose; densely
eaespitose perennials of eastern Oregon and adjacent
Idaho... dR ema Ee 34. C. propria
ii. Leaves setose and tomentose, less densely caespitose.
т тоске elas pe T 18. С. humilis
hh. DENM not exceeding the mature nutlets by more than 0. 5
mm.; densely caespitose perennials of southern Wyoming.
TT se e МА E к Е 19. C. caespitosa
ee. Scar at the nutlets closed or nearly во, no conspicuous triangular,
open area near the base
f. Upper surface of the lens uniformly appressed-strigose and
without pustules.
g. Nutlets ria ғ and tuberculate; scar surrounded by
au elevated пасе чье. . C. Bakeri
gg. Nutlets not so над rugose ог tuberculate; scar not sur-
rounded by an elevated margin
h. Densely caespitose from а multicipital caudex; native to
eastern Oregon and western Idaho............. 34. C. propria
hh. Less evidently or not че all caespitose; native р Р
Colorado, and Wyo ET Ca . С. sericea
ff. Upper surface of the leaves kon two distinct kinds 3 AE
pustulate.
g. Densely caespitose perennials; style not over 0.5 mm.
longer than mature nutlets; stems not over 1.5 ies high.
E E AuGcesaA 5 реки . C. caespitosa
gg. Кесә ды or solitary plants; stems usually more ri 1.5
m. high.

ү енен A mostly native in the Rocky Mountains or
west of them.

[Vor. 14
242 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

i. Mature sepals exceeding the nutlets by 2-3 mm.; in-
florescence broad-topped; western Colorado...... 21. C. elata
. Mature sepals exceeding the nutlets by 4-8 m
mn Nutlets tuberculate, scarcely if at all но
к. Ven en aiios of the nutlets тес 5.
MOS. oL rx bero rr азан . C. sobolifera
kk. = нан peris of the nutlets E sende
ened.

1 Stems 1-2 dm. high; inflorescence пее
western Colorado..................... . C. aperta
ll. Stems 2-9 dm. high; inflorescence d nar-
rower; Nevada to
m. Nutlets broadly lanceolate; murications
lacking or indefinite; Oregon, тее
and Мопќёапа.................. ‚С. Sheldonià
mm. Nutlets narrowly lanceolate; 5.
vory QUEM. ОР 25. C. interrupta
11. Nutlets more iM rugose.
k. Sear of the nutlets somewhat open at е Баве
clie and California................. ‚ С. humilis
kk. Sear closed or very nearly so; northerly in penta
1. Nutlets 4-5 mm. long; along the Columbia River
and tributary streams in Oregon and pc
ЛКК ГЕТ КЕТО ERES . С. celosioides
ll. Nutlets less than 4 mm. lon
Leaves енды iain ie strongly hir-

sute-cilia
n. Stems ix I 3-4 mm. long; Idaho
and Washington.............. 26. C. spiculifera

nn. Stems lor nutlets 2-3 mm. 5;
Canada, Washington, Wyoming. .29. С. Macounii
mm. Leaves broadly oblanceolate or spatulate.
n. Ventral surfaces of the ме. nearly

smooth; Montana.............. . C. sobolifera
nn. Ventral surfaces of the nutlets Fee
rough; Oregon to Montana. ..... 28. C. Sheldontt

hh. Biennials; native to the plains east of the Rocky Moun-

tains and to the foothill region east of the continental
QUE. АИ SA REV EE» 31. C. Bradburiana
Series 1. JAMESIANAE. Nutlets smooth on all surfaces, ог
rugose or tuberculate but not at all muricate on the dorsal surface,
ventral surfaces quite or nearly smooth. Nutlets often wing-
margined. Scar narrow, straight and closed or nearly so.

Margins not elevated. Species 1-11.

1. C. Jamesii (Torr.) new comb.
Myosotis suffruticosa Torr. Ann. Lyc. N. Y. 2: 225. 1827, not
Cryptantha suffruticosa Piper, Proc. Biol. бос. Wash. 32:42. 1919.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 243

Perennial; leaves linear-oblanceolate, obtuse or acute, 3-15
ст. long, indument usually appressed, scarcely hispid (in some
forms conspicuously setose), pustules small, sometimes confined
to the dorsal surface, always more numerous dorsally than ven-
trally; inflorescence usually open, cymes elongating, setose-
hirsute and subtomentose; sepals lanceolate, acute, in anthesis
3-4 mm. long, in fruit 5-6.5 mm. long, exceeding the nutlets by
3-4 mm.; corolla white, tube 2.5-3 mm. long, usually distinctly
shorter ihin the sepals, crests at the base of the tube conspicuous
and well developed, fornices probably yellow, elongated, usually
emarginate, about 1 mm. long, slightly papillose, limb 6-8 mm.
broad, tube and limb subequal, lobes united for 7/; to !/, their
length; fruit strongly oblate-ovoid, 1-4 nutlets maturing, style
exceeding the nutlets by 1-2 mm. ; nutlets rather narrowly ovate-
lanceolate in dorsal outline, 2-2.5 mm. long, acute, with margins
widely separated, acute, surfaces of nutlets smooth, glossy (with
30 diameters magnification the surfaces are evidently puberulent),
scar tightly closed, extending from the base to near the apex, no
elevated margin.

Distribution: Upper Sonoran and Transition Zones of the
southern Rocky Mountain region from central Wyoming to
Chihuahua and from western South Dakota and Texas to
Nevada and southern California.

The varieties of C. Jamesii are contrasted as follows:

Stems branched from the base, simple above, erect, 2-4.5 dm. tall; xi
tufted at the base; from southern Colorado and Utah to northern Mexic

гы EM Eee. c cerne re re RED и а. уаг. sa WM
Stems branched from near the base and upwards, erect, 2.5-4 dm. tall; leaves

apparently not tufted at the base, linear or nearly so; cymes usually much

elongated; northern Mexico and western Техав.................... 1b. var. (ата
Stems branched from the base, simple or sparingly branched above, erect,
1.2-2.5 dm. high; leaves somewhat tufted at the base; southern Colorado

and Utah to northern New Mexico and Arizona................. 1с. var. cinerea
Stems rather sparingly branched from the base and upwards, erect, 2.5-3.5

dm. high; leaves green, sparsely strigose on both surfaces, conspicuously

pustulate below, sparsely so above; southeastern Utah and northeastern

ATIZOhA..... ome Mec eie ose ces еж EEUU eee 1d. var. disticha
Stems branched upwards as well as from the base, more or less decumbent,

1-3 dm. long; leaves usually not tufted at the base; western Dakota and

Stems branched from the base, usually simple upwards, prostrate or nearly so,
m. long; leaves somewhat tufted at the base; eastern deserts of
southern California and adjacent Nevada..................... 1f. var. abortiva

[Vor. 14
244 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

1а. Var. multicaulis (Torr.) new comb. Plate 25, figs. 1-4.

Eritrichium multicaule Torr. in Marcy, Expl. Red River, 262.
1854.

Oreocarya multicaulis (Тогг.) Greene, Pittonia 3: 114. 1896.

Krynitzkia multicaulis var. setosa Jones, Contr. West. Bot. 13:
4. 1910. (Туре: “near Ft. Cove, Utah, June 27, 1901, growing
under junipers." М. Е. Jones.)

Oreocarya suffruticosa var. multicaulis (Torr.) Payson, Univ.
Wyo. Publ. Bot. 1: 171. 1926.

Perennial from а woody root; stems branched from the base,
simple above, erect, rather slender, 20-45 cm. tall, strigose and
pilose-hirsute; leaves tufted at the base, 5-15 cm. long, dorsal
surface strigose, appressed-setulose (in occasional forms setose-
hirsute), pustulate, ventral surface uniformly strigose, rarely
sparsely pustulate, petioles long-hairy toward the base, finely
ciliate; inflorescence becoming open by the elongation of the
cymes, usually confined to the upper 14, or less, of the stem, foliar
bracts inconspicuous; 2-4 nutlets commonly maturing.

Distribution: Upper and Lower Sonoran Zones, northwestern
Oklahoma, western Texas, southwestern Colorado, New Mexico,
southern Utah, eastern Nevada, Arizona and northern Mexico.
Type: from near Santa Fe, New Mexico, Fendler 636.

Specimens examined:

Oklahoma: near Knowles, Beaver Co., May 5, 1913, Stevens 336
(Мо., Gray); near Shattuck, Ellis Co., May 10, 1914, Clifton 3039
(Mo.); near Shattuck, May 16 and 17, 1914, Stevens 3039, 3024
(Gray). |

Texas: sandhills оп Canadian, Hemphill Co., June 4, 1901,
Eggert (Mo.); Big Springs, May 15, 1902, Tracy 7835 (Minn.,
U. S); Dalhart, June 24, 1920, Jones 339 (Gray); Ft. Davis,
June, 1881, Havard (U. S.); Clarendon, 1888, Nealley (U.S.).

Colorado: Durango, July 10, 1896, Tweedy 576 (U.S.) ; Arboles.
June, 1899, Baker 563 (Pomona, К. Mt., Mo., U.S., Gray).

New Mexico: Tunitcha Mts., Aug. 8, 1911, Standley 7830
(U.S.); near Cedar Hill, San Juan Co., Aug. 16, 1911, Standley
7967 (U.S.); near Farmington, San Juan Co., July 20, 1911,
Standley 7125 (U.S.); near Santa Fe, 1847, Fendler 636 (N.Y.,
U.S., Gray); near Las Vegas, June, 1920, Anect 164 (U.S.);

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 245

Albuquerque, Sept. 21, 1884, Jones 6687 (Pomona); near Ресов,
San Miguel Co., Aug. 15, 1908, Standley 4899 (U.S., Mo.);
Florita Mts., Sept. 7, 1903, Jones (Pomona); Las Palomas,
Sandia Mts., April-May, 1914, Ellis 366 (Mo., U.S.); Balsam
Park, Sandia Mts., Aug. 4, 1914, Ellis 463 (U.S.) ; Datil, Socorro
Co., бері. 29-Oct. 1, 1919, Eggleston 16184 (Field) ; near Roswell,
June, 1914, Wooton (U.S.); Dofia Ana, April, 1851, T'hurber 293
(Gray); Jornada del Muerto, April, 1851, T'hurber 278 (Gray);
Organ Mts., Aug. 28, 1897, Wooton 401 (Pomona, R. Mt., Calif.,
Minn., Mo., U.S.); Organ Mts., July 15, 1897, Wooton 595
(Minn., Mo., U.S.); Animas Valley, Oct. 2, 1893, Mearns 2522
(U.S.); Carrizallito Mts., April 17, 1892, Mearns 111 (U.S.);
Silver City, April 28, 1919, Eastwood 8366 (U.S., Gray); Middle
Fork of the Gila, Mogollon Mts., Aug. 5, 1900, Wooton (U.S.);
north of Carrizozo, Aug. 28, 1904, Wooton 2818 (U.S.); Mangas
Springs, n.w. of Silver City, May 17, 1903, Metcalfe 70 (Pomona,
В. Mt., Calif., Minn., U.S., Gray); Mogollon Mts., Aug. 9, 1903,
Metcalfe 431 (Pomona, Minn., U.S., Calif., Gray); Gila Hot
Springs, Mogollon Mts., Aug. 27, 1903, Metcalfe 863 (В. Mt.,
Mo.); Mimbres River, Grant Co., July 1, 1904, Metcalfe 1061
(U.S.); Santa Rita del Cobre, 1877, Greene 32 (Gray).

Utah: Pahria Canyon, May 26, 1894, Jones 5297q, 52986
(U.S.); canyon above Tropic Мау, 28, 1894, Jones 5300 (Pomona,
U.S.).

Arizona: Fort Valley, Coconino Nat’l. Forest, July 14, 1909,
Pearson 210 (U.S.) ; Metcalfe, Oct. 1, 1900, Davidson 608 (Gray);
Ft. Apache, June 21-30, 1890, Palmer 591 (U.S., in рагі); Skull
Valley, April 28, 1903, Jones (Pomona); Flagstaff, Aug. 6, 1884,
Jones 4007 (Pomona, В. Mt., U.S.); Flagstaff, June 4, 1898,
MacDougal 49 (R.Mt., Calif., U.S., Gray); near Flagstaff, June
17, 1901, Leiberg 5545 (U.S.); Flagstaff, Aug. 25, 1883, Rusby
749 (Calif., U.S.); northeast of Flagstaff, June 8, 1922, Hanson
4120 (Mo.); San Francisco Mts., Aug. 3, 1923, Jaeger (Pomona);
San Francisco Mts., April, 1881, Rusby 283 (Minn., U.S.); San
Francisco Mts., Oct. 1881, Lemmon & wife (R. Mt.); Williams,
Aug. 6-15, 1903, Griffiths 4912 (U.S.); Bowie, Sept. 18, 1884,
Jones 6685 (Pomona); Cosnino, Aug. 9, 1884, Jones 4047 (Po-
mona, U.S.); Santa Rita Forest Reserve, March 31-Аргі 23,

[Vor. 14
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1903, Griffiths 4268 (U.S.) ; Alpine, July 31, 1912, Goodding 1264
(В. Mt., Calif., U.S.); Outlaw Canyon, Chiricahua Mts., July 30,
1907, Goodding 2349 (R.Mt., Gray).

Nevada: Ely, June 26, 1907, Jones (Pomona).

Chibuahua: Mexican boundary near White Water, June 18,
1892, Mearns 359, 360 (U.S.); base of San Luis Mts., Sept. 5,
1893, Mearns 2091 (U.S.); Casas Grandes, May 13, 1899, Goldman
407 (U.S., Gray); plains of Guerrero, Sept. 8, 1887, Pringle
(Calif.).

1b. Var. laxa (Macbr.) new comb.

Oreocarya multicaulis var. lara Macbr. Contr. Gray Herb.
48: 35. 1916.

Perennial; stems branched from near the base and upwards,
erect, rather stout, 2.5-4 dm. high, strigose and hirsute with
ascending hairs; leaves numerous, linear or linear-oblanceolate,
acute, 6-12 ст. long, rather coarsely and sparsely appressed-
strigose below, finely strigose above, pustulate on both surfaces
but much more abundantly so on lower; inflorescence broad and
open, the cymes becoming much elongated in age (10 cm.), con-
fined to the upper 14-14 of the stem, foliar bracts not conspicu-
ous, linear.

Distribution: Lower Sonoran Zone in northern Chihuahua,
Mexico, and western Texas. Type: sand hills near Paso del
Norte, Chihuahua, C. G. Pringle 776.

Specimens examined:

Texas: Marfa, Sept. 1883, Havard (U.8.).

Chihuahua: sand hills near Paso del Norte, Sept. 20, 1886,
Pringle 776 (Field, Mo., Phila., U.S., Gray, TYPE); between Casas
Grandes and Sabinal, Sept. 4-5, 1889, Nelson 6350 (U.S., Gray).

1c. Var. cinerea (Greene) new comb.

Oreocarya cinerea Greene, Pittonia 3: 113. 1896.

O. Lemmoni Eastw. Bull. Torr. Bot. Club 30: 239. 1903.
(Type: Arizona, without definite locality, 1884, Lemmon.)

O. multicaulis var. cinerea (Greene) Macbr. Proc. Am. Acad.
51:54. 1916

O. suffruticosa var. cinerea (Greene) Payson, Univ. Wyo. Publ.
Bot. 1: 171. 1926.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 247

Perennial; stems branched from the base, simple or sparingly
branched above, erect, rather slender, 12-25 cm. high,strigose and
setose-hirsute; leaves somewhat tufted at the base, 5-15 cm. long,
dorsal surface strigose and appressed-setulose (rarely setose-hir-
sute), pustulate, ventral surface densely and uniformly strigose,
with few or no pustules, petioles long-hairy at the base, finely cili-
ate; inflorescence becoming open by the elongation of the cymes,
usually confined to the upper 15 or 24 of the stem, bracts of the
lower part of the inflorescence rather large and leaf-like; 2-4
nutlets commonly maturing.

Distribution: Upper Sonoran and Transition Zones of south
central Colorado, northern New Mexico, southern Utah, northern
Arizona and eastern Nevada. Туре: ‘‘Confined, as far as I
know, to the Arkansas Valley, in southern Colorado, where it oc-
cupies low subsaline clayey soils," Greene.

Specimens examined:

Colorado: Canyon City, June 28, 1917, Payson 1019 (R.Mt.);
Canyon City, June 27, 1895, Osterhout 628 (R.Mt., Minn.,
Osterh.); Rocky Ford, June 18, 1900, Osterhout 2087 (R.Mt.,
Osterh.); Florence, June 14, 1892, Cowen (Mo.); Wet Mountain
Valley, June 20, 1873, Brandegee (Mo.); plains, Pueblo, 1873,
Greene (Gray); Pueblo, June, 1890, Mr. & Mrs. G. H. Hicks 178
(Minn.); mesas near Pueblo, May 14, 1900, Rydberg & Vreeland
5702 (R.Mt.); Buena Vista, July 6, 1892, Sheldon 527 (0.8.);
Gunnison, July 29, 1925, Smith 7680 (R.Mt.) ; Gunnison, July 17,
1901, Baker 455 (Pomona, R.Mt., Minn., Mo., U.S., Gray).

New Mexico: 15 miles w. of Sante Fe, May 22, 1897, A. A. &
E. G. Heller 3577 (Pomona, Minn., Mo., U.S., Gray).

Utah: along Bullion Creek, above Marysvale, July 21, 1905,
Rydberg & Carlton 7041 (R.Mt., U.S.); Marysvale, May 31, 1894,
Jones 5328 (Pomona, Calif., Mo., U.S.); near Fort Cove, June 27,
1901, Jones (Pomona).

Arizona: Moki Indian Reservation, Aug. 1-бері. 5, 1897,
Hough 8 (U.S.); Grand Canyon, June, 1915, Macbride & Payson
950 (R.Mt., U.S., Gray); vicinity of Flagstaff, July 2, 1898, Мас-
Dougal 204 (R.Mt., Calif., Phila., U.S., Gray); Cosnino, Aug. 9,
1884, Jones 4042 (Pomona, R.Mt., U.S.); near Flagstaff, May-
Oct., 1901, Purpus 8195 (Calif., Mo.); Cosnino, June 9, 1890,

[Vor. 14
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Jones (Pomona); San Francisco Mts., June 9, 1893, Jones 6688
(Pomona); San Francisco Mts., May-Oct., 1900, Purpus 8048a,
7067 (Calif.); Ash Fork, June 18, 1901, Barber 112 (U.S.); Pres-
cott, 1876, Palmer 375 (Gray); Peach Springs, May 26, 1884,
Jones 6684, (Pomona, R.Mt., Calif.).

Nevada: Ely, Aug. 11, 1913, Hitchcock 1198 (U.S.); Mt.
Magruder, May-Oct., 1898, Purpus 6068 (Pomona, U. 5.).

1d. Var. disticha (Eastw.) new comb.

Oreocarya disticha Eastw. Bull. Torr. Bot. Club 30: 238. 1903.

Perennial; stems branched from the base and more or less
branched upwards, strigose and sparingly setose with slender as-
cending trichomes, 2.5-3.5 dm. high, erect; leaves narrowly ob-
lanceolate, obtuse or acute, 5-8 ст. long, green, sparsely strigose
on both surfaces, densely pustulate beneath, sparingly so above;
inflorescence open, cymes elongating, lax, foliar bracts rather
conspicuous; 1-4 nutlets maturing.

Distribution: Upper Sonoran and Transition Zones in south-
eastern Utah and northeastern Arizona. Type: “оп the mesa
above the San Juan River, Utah, on what was known as Barton’s
Range, July 13, 1895," Eastwood.

Specimens examined:

Utah: Rabbit Valley, Aug. 12, 1875, Ward 557 (U.S., Mo.);
Thousand Lake Mt., July 14, 1875, Ward 393 (0.5.); Barton
Range, s.e. Utah, July 13, 1895, Eastwood (Calif., U.S., Mo.,
Gray).

Arizona: Laguna Canyon, July 10-11, 1920, Clute 37 (U.S.,
R.Mt., Gray).

le. Var. typica n. var.

Муозойз suffruticosa Torr. Ann. Гус. №. Y. 2: 225. 1827.

Eritrichium Jamesii Torr. in Marcy, Expl. Red River, 262.
1854.

Krynitzkia Jamesii (Torr.) Gray, Proc. Am. Acad. 20: 278.
1885, in part.

Oreocarya suffruticosa (Torr.) Greene, Pittonia 1: 57. 1887.

Perennial, but probably shorter-lived than in the varieties
cinerea and multicaulis; stems branched upwards as well as from

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 249

the base, more or less decumbent, 10—30 em. long, rather slender,
strigose and setose-hirsute; leaves usually not tufted at the base,
linear-oblanceolate, 3-8 ст. long, dorsal surface densely strigose
and appressed-setulose, pustulate, ventral surface uniformly
strigose, only rarely appressed-setulose or pustulate, petioles
somewhat ciliate near the base; inflorescence open, mostly con-
fined to the upper 14-24 of the stem, cymes elongating, bracts of
the lower part of the inflorescence elongated and leaf-like; 2-4
nutlets commonly maturing.

Distribution: Upper Sonoran Zone in the western part of South
Dakota, Nebraska, and Kansas, northwestern Oklahoma and
Texas, southeastern Wyoming, and eastern Colorado. Туре:
“barren desert along the Platte," James.

Specimens examined:

South Dakota: Indian Creek, Aug. 26, 1891, Williams (Wash.).

Nebraska: Alliance, June 29, 1913, Bates 5768 (Minn.); АІ-
liance, May 16, 1914, Bates 5897 (Minn.); near Plummer Ford,
Thomas Co., July 8, 1893, Rydberg 1514 (N.Y., U.S., Gray);
Wild Cat Mts., Banner Co., July 16, 1891, Rydberg 254 (U.S.);
Sidney, May 23, 1922, Nelson (R.Mt.).

Kansas: near Hay Springs, June 6-7, 1901, MacDougal 79
(№. Y., Mont.); Hamilton Co., Aug. 3, 1895, Hitchcock 347 (В.
Mt., U.S., Gray); Syracuse, Hamilton Co., July 11, 1893, Thomp-
son 97 (N.Y., U.S.); Comanche Co., June 17, 1891, Carlton 246
(U.8.); Arkalon, Seward Co., June 27, 1888, Kellerman 6 (U.S.).

Texas: Canadian, July 8, 1912, Condit (Calif.).

Wyoming: Casper, July 6, 1901, Goodding 208 (R.Mt., Pomona,
Mo., U.S., Gray); Platte River at Ferris, July 19, 1898, E. Nelson
4906 (R.Mt., Mo., U.S., Gray); Douglas, July 20, 1915, Hess 106
(R.Mt.); Uva, July 31, 1896, Nelson 2567 (R.Mt., Minn.); Platte
River Canyon (Wheatland), July 14, 1894, Nelson 477 (R.Mt.,
Wash., Minn., Mo., U.S., Gray); Powder River, June 28, 1910,
Nelson 9379 (R.Mt., Minn., Gray); Ft. Laramie, June 29, 1901,
Nelson 8305 (R.Mt.); Pine Bluffs, June 28, 1889, Bodin (Minn.);
Pine Bluffs, May 15, 1897, Nelson 2882 (R.Mt.).

Colorado: Wray, July 14, 1909, Osterhout 3992 (R.Mt., Osterh.) ;
Wray, July 1-4, 1919, Eggleston 15168 (Pomona) ; Sterling, Logan
Co., June 12, 1896, Osterhout (Minn.); Ft. Lupton, July 8, 1916,

[Vor. 14
250 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Johnston 845 (Mo., U.S.); White Rocks, Boulder Co., July 19,
1918, Cockerell (U.S.); near Boulder, July, 1902, Tweedy 5219
(R.Mt.); barren deserts high up on the Platte, James (N.Y.,
TYPE); Denver, July 10, 1919, Payson & Bethel 1608 (R.Mt.);
North Denver, Aug. 12, 1910, Eastwood (Gray); Golden, June 3,
1870, Greene 300 (Gray) ; Castle Rock, June 30, 1917, Payson 1027
(R.Mt.); Table Rock, Е Paso Co., July 15, 1891, Crandall
(Calif.); Colorado Springs, June 26, 1903, Shantz 571 (U.S.);
Colorado Springs, July 3, 1895, Osterhout 627 (R.Mt., Osterh.);
Colorado Springs, July 22, 1920, Johnston 2809 (Gray) ; Colorado
Springs, June 19-30, 1915, Eggleston 11194a (U.S.); Colorado
Springs, May 28, 1879, Jones (Pomona); Colorado Springs, Мау
15, 1878, Jones 66 (U.S.); Salida, Aug. 2, 1925, Smith 3912 (R.
Mt.); Blanca, Costilla Co., June 27, 1921, Bethel, Nielley &
Clokey 4258 (R.Mt., Phila.).

1f. Var. abortiva (Greene) new comb.

Oreocarya abortiva Greene, Pittonia 3: 114. 1896.

Krynitzkia multicaulis var. abortiva (Greene) Jones, Contr.
West. Bot. 13: 5. 1910.

Oreocarya suffruticosa var. abortiva (Greene) Macbr. Proc. Am.
Acad. 51:547. 1916.

Perennial from a woody root; stems branched from the base,
usually simple upwards, rather slender, prostrate or nearly so,
6-15 em. long, strigose and hirsute; leaves somewhat tufted at
the base, 5-10 em. long, dorsal surface densely strigose, appressed-
setose and pustulate, petioles long-hairy at the base, finely ciliate;
inflorescence not raised above the leaves, extending over 34 of
the stem, bracts conspicuous and elongated; 2-3 nutlets fre-
quently aborted.

Distribution: Upper Sonoran Zone in the eastern deserts of
southern California and in southern Nevada. Type: “Bear Val-
ley, San Bernardino Mountains, California," S. B. Parish.

Specimens examined:

Nevada: Lee Canyon, Charleston Mts., Clark Co., July 28,
1913, Heller 11016 (Calif., Phila., Field, U.S., Gray).

California: north side of Bear Valley, June 12, 1922, Munz 5721
(Pomona, Calif., Gray); east of Big Meadows, San Bernardino

1927]
PAYSON—SECTION OREOCARYA ОЕ СКУРТАМТНА 251

Mts., June 21, 1922, Pierson 3103 (Pomona); Bear Valley, San
Bernardino Co., July 19, 1900, Jones 5316} (Pomona); Bear
Valley, June 16-20, 1895, Parish 3694 (СаШ., Gray); Bear
Valley, May, 1882, Parish 1480 (Calif, Mo., U.S., Gray);
Bear Valley, 6500 ft., June 15, 1894, Parish 3238 (0.8.).

C. Jamesii, in the aggregate sense, is the most widely distrib-
uted and most heterogeneous species in the genus. Ав а species
it is very easily recognized by the smooth nutlets that are not in
contact by their edges. It is likely to be confused with only two
other units that are here accorded specific rank, C. pustulosa and
C. Palmeri. 'The glabrous stems and ventral leaf surfaces dis-
tinguish the former, and the accrescent calyces, longer corolla
tubes, and lack of crests at the bases of the tubes identify the
latter.

The varieties of C. Jamesii are quite difficult of determination,
and intermediates between the various groups are of fairly com-
mon occurrence. These intermediates are usually in inter-
mediate ranges but occasionally occur far inside the range of a
different variety. It is possible, of course, that some of these
anomalies are produced by unusual habitats. This is the more
probable since the differences between the varieties are entirely
those of habit. Since the different forms are fairly well segregated
geographically it is to be expected that local floras will continue
to treat the variants as separate species. This is greatly to be
deplored, since it will overemphasize the distinctions and serve
to make determinations difficult.

On the other hand, the general consistency of the varietal ranges
makes it certain that the varieties are really incipient species and
so worthy of named recognition in any critical work. "The author
has no doubt but that some specimens cited are incorrectly placed.
Sometimes in examining two duplicates of the same collection one
is tempted to place one in one variety and the other in a different
one. Botanists working in an intermediate range, such as at
Flagstaff, Arizona, will probably find the varieties, as in this case,
cinerea and multicaulis, in hopeless confusion.

'The setose-hispid form of the variety multicaulis is, according
to Macbride, to be considered as typical multicaulis. He would
make the matter of spreading or appressed pubescence the cri-

[Vor. 14
252 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

terion of distinction between cinerea and multicaulis. It seems
to the present author that such a classification cuts across the
lines of real relationship. Cinerea is more than a form of multi-
caulis with appressed setae. There are more specimens of multi-
caulis with appressed than with spreading setae. Cinerea is a
growth form intermediate between multicaulis and typica and as
such is perhaps the least definite of the varieties. It might be
better to submerge the variety cinerea in multicaulis. It is on
the southern edge of the range of cinerea that it is most difficult
of delimitation. The var. setosa Jones is an exact synonym of
Macbride’s idea of multicaulis.

The variety (ата seems certainly more than the ecological
variety that Macbride suggested. Other collections of it have
confirmed its peculiarities.

The variety disticha needs much more study, but the similar
collections from the type region seem to assure the general preva-
lence of an unusually green form of the species in southern and
southeastern Utah and adjacent Arizona. The character of the
single nutlet maturing is certainly of no value. On part of the
type preserved in the herbarium of the University of California
some fruits were found with three nutlets. In another collection
of similar general appearance four mature nutlets were found.

Other varieties may need to be characterized from time to
time. А study of the different varieties in intermediate ranges
should prove of great interest to local botanists.

2. C. pustulosa (Rydb.) new comb.

Oreocarya pustulosa Rydb. Bull. Torr. Bot. Club 40: 480. 1913.

Perennial; stems slender, branched at the base, 3-5 dm. high,
glabrous or nearly so throughout; leaves linear-oblanceolate, ap-
parently not clustered at the base, numerous on the stems, 3-10
ст. long, acute or obtuse, glabrous and without pustules above,
conspicuously and densely pustulate and short-hairy below; in-
florescence confined to the upper М or less of the stem, cymes
elongating and probably lax in age, foliar bracts inconspicuous;
sepals lanceolate, acute, about 4 mm. long in anthesis, probably
not conspicuously elongated in fruit, strigose and rather sparsely
setose with short, appressed bristles; corolla white, tube 2.5-3

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 253

mm. long, somewhat shorter than the sepals in anthesis, crests at
the base of the tube evident but not large, fornices about 1 mm.
long, slightly papillose, probably yellow, limb 5-6 mm. broad,
tube and limb subequal, lobes united for 14 their length; mature
fruit not seen, presumably very like C. James.

Distribution: Upper Sonoran Zone, southeastern Utah. Туре:
“Hammond Canyon, Elk Mountains, July 31, 1911, Rydberg &
Garrett 9320.”

Specimens examined:

Utah: Hammond Canyon, Elk Mts., Aug. 10, 1911, Rydberg &
Garrett 9569 (R.Mt., U.S.).

This species is quite unlike any other species of the genus in
general appearance, due chiefly to the absence of any conspicuous
setae and the quite glabrous stems and upper surfaces of the
leaves. Тһе original description characterizes the leaves as
being ‘‘glabrous beneath, sparingly hairy above." In the speci-
men at hand it is certainly the upper leaf-surface that is glabrous.
Тһе specimen available of this species has more the appearance
of a Heliotropium or of а Plagiobothrys than а Cryptantha but
there seems no doubt that it is correctly placed in the latter genus.
Тһе variety disticha of Jamesii is probably something of а con-
necting link between Jamesii var. multicaulis and the present
species.

3. C. Palmeri (Gray) new comb.

Krynitzkia Palmeri Gray, Proc. Am. Acad. 20: 278. 1885.

Oreocarya Palmeri (Gray) Greene, Pittonia 1: 57. 1887.

Caespitose, long-lived perennial; stems erect, rather stout,
15-30 сто. tall, densely setose with long, rather slender, divaricate
hairs; radical leaves linear to linear-oblanceolate, acute, 4-7 cm.
long, 2-7 mm. broad, tomentulose and subappressed-setose,
abundantly pustulate on the dorsal surface, pustules smaller and
fewer on the ventral surface; cauline leaves similar but smaller;
inflorescence conspicuously, but rather softly, setose, mainly
limited to the upper l4 of the stem, cymes elongating, foliar
bracts inconspicuous; calyx setose, sepals in anthesis linear-
lanceolate, acute, about 5 mm. long, in fruit 7-10 mm. long, ex-
ceeding the nutlets by 6-8 mm.; corolla probably white, tube 4-5

[Vor. 14
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mm. long, equalling or shorter than the calyx lobes in anthesis,
crests lacking at the base of the tube, fornices elongated, papillose,
0.5-1 mm. long, limb about 8 mm. broad, lobes united for 14-16
their length, shorter than the tube; fruit strongly oblate-ovoid,
all 4 nutlets maturing, style exceeding the nutlets by 3-4 mm.;
nutlets very similar to those of C. Jamesii, about З mm. long, apex
acute, margins not in contact, acute, all surfaces smooth, glossy,
scar straight, closed or nearly so, extending from the base to
beyond the middle, margin not elevated.

Distribution: Lower Sonoran Zone in western Texas and ad-
jacent Mexico. Type: “Coahuila, Mexico, forty miles south of
Saltillo, Dr. Palmer, March, 1880, no. 895 of the distribution."

Specimens examined:

Texas: rocky hills near Big Spring, Howard Co., June 11, 1900,
Eggert (Mo., Gray); sandy hills and plains, Upper Concho, April,
Reverchon 2120 (Minn., Field, N. Y., Mo., U.S., Gray); Toyah,
Reeves Co., May 2, 1902, Tracy & Earle 424 (U.S.) ; from western
Texas to El Paso, May-Oct. 1849, Wright 489 (U.S., Gray).

Coahuila: 40 miles south of Saltillo, March, 1880, Palmer 895
(Gray, TYPE).

This species has long remained obscure because of the im-
maturity of the type. Тһе author had designated the plant col-
lected by Reverchon (no. 2120) as the type of a new species
before he had examined the type of C. Palmeri. Dr. Gray had
described Palmeri as having subrugose nutlets and this is not
true of the Reverchon plant. With the type of Palmeri at hand
it is evident that the nutlets are subrugose only because they are
immature. Тһе other floral characters of Palmeri agree very
well with the Texan specimen, and it seems very certain that at
last the identity of Palmeri has been established.

This species is very close to Jamesii var. multicaulis and is
certainly to be considered as a recent derivative from it. Its
distinguishing characters are: (1) the accrescent sepals, (2) the
corolla tube with no crests at the base, (3) the longer style.

4. C. oblata (Jones) new comb. Plate 25, figs. 5-7.

Krynitzkia oblata Jones, Contr. West. Bot. 13: 4. 1910.

Oreocarya hispidissima Wooton & Standley, Contr. U. S. Nat.
Herb. 19: 545. 1915, not О. hispidissima (Torr.) Rydb.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 255

О. oblata (Jones) Macbr. Proc. Am. Acad. 51: 548. 1916.

O. Paysonii Macbr. Contr. Gray Herb. 48: 36. 1916 (Type:
limestone hills, Berendo Creek, Sierra Co., New Mexico, May 12,
1905, Metcalfe 1576).

Caespitose perennial; stems several to many from the base,
rather slender, 1.5-2.5 dm. high, hirsute and becoming more con-
spicuously setose upwards; leaves tufted at the base, linear-ob-
lanceolate, usually acute, 5-10 cm. long, dorsal surface rather
coarsely strigose and appressed setose, pustules rather numerous,
ventral surface more finely strigose, weakly and sparsely ap-
pressed-setose, sparingly pustulate; petioles more or less ciliate
toward the base; inflorescence confined to the upper 1% of the
stem, cymes somewhat elongated in age and so the thyrsus not
very narrow, densely setose with rather slender bristles, foliar
bracts inconspicuous; calyx abundantly setose with rather weak
bristles, sepals in anthesis nearly linear, 6-7 mm. long, acute, in
fruit 10-12 mm. long, exceeding the nutlets 7-8 mm.; corolla
white, tube 8-10 mm. long, exceeding the sepals by 2-3 mm.,
crests lacking at the base of the tube, fornices probably yellow,
rounded, broad and low (0.5-1 mm.), minutely papillose, limb
8-10 mm. broad, lobes united for about 14 their lengths, scarcely
half as long as the tube; fruit oblate-ovoid, all four nutlets usually
maturing, style exceeding the mature nutlets by 3-5 mm. (species
evidently moderately heterostyled) ; nutlets 2.5-3 mm. long, ovate
or nearly circular in outline, obtuse at the apex, margins acute,
not quite in contact, surfaces of nutlets glossy, the dorsal rather
sparsely tuberculate and more or less rugose, these elevations low
and rounded, ventral surfaces smooth, scar closed, extending from
the base to about the middle of the nutlet, no elevated margin
present.

Distribution: Lower Sonoran Zone of southern New Mexico
and adjacent Texas. Probably occurs also on the Mexican side
of the Rio Grande near Е Paso. Туре: “по. 3759 [М. Е.
Jones], El Paso, Texas, April 23, 1884.”

Specimens examined:

Texas: Marfa, March 26, 1919, Hanson 399 (U. S., Gray); near
J. Davis’ В. C., West Texas, Sept. 1883, Назата (U.S.); Fort
Bliss, April 22, 1915, Carlson (Gray); El Paso, April, 1881, Vasey

[Vor. 14
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(Mo.); near El Paso, March, 1851, T'hurber 14? (Gray); El Paso,
April 23, 1884, Jones 3759 (Pomona, түре, R.Mt., 0.5.);
northern base of Chenate Mts., Sept. 1892, Nealley 167 (Field,
N.Y., 0.5.).

New Mexico: 1851-52, Wright 1566 (Field); mesa west of
Organ Mts., May 6, 1906, Standley (U.S.); Organ Mts., April 4,
1903, Wooton (R.Mt., Pomona); Tortugas Mt., Aug. 29, 1902,
Wooton (U.S.); Tortugas Mt., April 22, 1894, Wooton (U.S.);
Organ Mts., March 30, 1905, Wooton (U.S.); Lake Valley, south-
ern Sierra Co., 1915, Beals (U.S.); Berendo Creek, Sierra Co.,
Мау 12, 1905, Metcalfe 1576 (Pomona, Field, Mo., U.S., Gray).

This species is readily distinguished from all other species by
the combination of an exserted corolla tube and nutlets that are
smooth on the inner surfaces and distinctly roughened on the
outer. In general appearance it is not very unlike C. Jamesii
var. multicaulis or var. cinerea. In the original description of
C. oblata, Mr. Jones also cited а specimen from Peach Springs,
Arizona. This proves on examination to be C. James var.
cinerea. |

C. oblata is most closely related to C. Palmeri and to С. Jamesii
var. multicaulis. It differs from both in having exserted corolla
tubes and somewhat roughened nutlets that are quite distinetly
wing-margined.

5. C. confertiflora (Greene) new comb. Plate 25, figs. 8-10.

Krynitzkia leucophaea var. alata Jones, Proc. Calif. Acad. Sci.
II. 5: 710. 1895. (Туре: “Хо. 5289 May 23, 1894, Johnson,
Utah, 5,000 ft. alt., on sandstone cliffs," M. E. Jones).

Огеосатуа confertiflora Greene, Pittonia 3: 112. 1896.

O. leucophaea var. confertiflora (Greene) Parish, Erythea 7: 95.
1899.

O. lutea Greene, Muhlenbergia 2: 240. 1906; Brand, Fedde,
Rep. Sp. Nov. 19: 73. 1923. (Туре: “по. 8211, collected May
9, on rocky slopes in Silver Canyon in the White Mountains, op-
posite Laws, Inyo County,” California, A. A. Heller).

O. alata (Jones) A. Nels. Coulter & Nelson, Man. Cent. Rocky
Mts., 417. 1909; Rydb. Fl. Rocky Mts. 725. 1917.

рава peremit from à woody root; stems few to many

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 257

from a branching caespitose caudex, slender, simple, 15-50 cm.
tall, densely white-hairy at base, appressed-strigose and sparingly
setose upwards, bristles mostly appressed; leaves somewhat
clustered near the base, linear-oblanceolate, acute, 3-10 cm.
long, dorsal surface densely strigose and with rather numerous
small appressed bristles with pustulate bases, ventral surface
uniformly strigose, in age minutely pustulate, petioles ciliate;
inflorescence mainly restricted to the upper 4% of the stem or
less, usually distinctly glomerate, cymules short, foliar bracts
inconspicuous, bristles yellowish in age, divaricate; calyx setose
with rather short, weak bristles, sepals in anthesis linear-lance-
olate, acute, 7-10 mm. long, in fruit 10-12 mm. long, exceeding
the nutlets by about 6 mm.; corolla yellow, tube 9-13 mm.
long, 2-3 mm. longer than the sepals, crests at base of the
tube usually evident, confluent into a ring (sometimes obsolete),
fornices broad, emarginate, about 1 mm. high, limb about 10 mm.
broad, lobes united for about 14 their length, about 15 as long
as the tube; fruit broadly ovoid (nearly square in cross-section
due to the flat dorsal surfaces of the nutlets), all four nutlets com-
monly maturing, style exceeding the nutlets 3-6 mm. (species
conspicuously heterostyled) ; nutlets broadly ovate in dorsal view,
obtuse or subacute, about 3 mm. long, nearly triangular in cross-
section, somewhat papery margins in contact, acute, almost wing-
margined, surfaces of nutlets glossy, smooth, scar straight, nearly
closed, extending from near the base to above the middle of the
nutlet, margin not elevated.

Distribution: Upper Sonoran Zone, western Utah, northern
Arizona, southern Nevada, and the eastern desert region of
southern California. Type: “at Cushenberry Springs on the
north side of the San Bernardino Mountains, southern California,"
S. B. Parish.

Specimens examined:

Utah: Dutch Mt., Tooele Co., June 8, 1900, Jones (Pomona);
Gold Hill, near Clifton, June 13, 1891, Jones (Pomona) ; Dugway,
w. Utah, May 27, 1891, Jones (Pomona); Fish Springs, June 4,
1891, Jones (Pomona, R. Mt., Calif., Mo., U.S.) ; Elsinore, Sevier
Co., June 13, 1899, Jones (Pomona); red sand, Springdale, May
17, 1894, Jones 5261) (Pomona, U.S.); Wa Wa, 30 miles west of

[Vor. 14
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Frisco, June 25, 1906, Jones (Pomona); Vermilion, June 4, 1901,
Jones (Pomona); canyon south of Glenwood, Мау 25, 1875,
Ward 108 (Mo., U.S., Gray); Johnson, May 23, 1894, Jones 5289t
(Pomona); Johnson, June 20, 1890, Jones (Pomona); Silver Reef,
Мау 3, 4, 1894, Jones 5144 (Pomona, R.Mt., Calif., Mo., 0.8.);
Diamond Valley, May 16, 1902, Goodding 814 (Pomona, R.Mt.,
Mo., U.S., Gray); Beaverdam Mts., May, 1874, Parry 166 (Mo.,
Gray).

Arizona: 3 miles below Tanner’s Crossing, Little Colorado,
May 27, 1901, Ward (№. Y., U.S.); Red Canyon trail, Grand
Canyon, June 10, 1901, Ward (0.5.).

Nevada: The Muddy Range, April 10, 1905, Goodding 2221
(R.Mt., Minn., Mo., Gray); Mormon Mts., Lincoln Co., July,
1906, Kennedy & Goodding 110 (Calif.); Meadow Valley Wash,
mile 16, April 28, 1904, Jones (Pomona); Good Springs, May 1,
1905, Jones (Pomona); Good Springs, May, 1915, К. Brandegee
(Calif.) ; Goldfield, May 14, 1909, Heller 9619 (Phila.) ; Esmeralda
Co., Shockley (Calif.).

California: Andrew's Camp, Bishop Creek, Inyo Co., July, 1911,
Davidson 2722 (Gray); Andrew's Camp above Bishop Creek,
July, 1913, К. Brandegee (Pomona, Calif.); Bishop Creek, May
31, 1906, Hall & Chandler 7234 (Calif.); northern slope of San
Bernardino Mts., May, 1882, Parish 1819 (СаШ.); near summit of
Wild Rose Canyon, Inyo Co., Parish 19167 (Calif.) ; Cactus Flat,
desert slope of San Bernardino Mts., June 2, 1901, Parish 4887
(Calif., U.S.); San Bernardino Mts., June 17, 1894, Parish 3240
(Mo., U.S.); Water Canyon, San Bernardino Mts., May, 1882,
Parish 1316 (Mo., Gray); near Cushenberry Springs, Mojave
Desert, May, 1882, Parish & Parish 1316 (Calif., Mo.); Erskine
Creek, May, 1897 (7982), Purpus 5323 (СаШ., Mo., U.S., Gray);
White Mts., 1898, Purpus 5802 (Calif., U.S.); Silver Canyon
east of Laws, May 9, 1906, Heller 8211 (Cahf., Mo., Phila., U.S.,
Gray); south of Mono Lake, July 8, 1863, Brewer 1822 (0.8.);
Pleasant Canyon, Panamint Mts., May 6, 1897, Jones (Pomona,
U.S.); Kern River, Austin (Calif.); Olancha Mt., Tulare Co.,
June 25-30, 1904, Hall & Babcock 5270 (Cahf.).

C. confertiflora and its close relatives, C. flava and C. leucophaea,
are readily distinguishable from all other species of the section

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 259

by their long corolla tubes and smooth nutlets. They are not so
easily separable from one another, however, and a broader specific
concept would unite them as varieties of one species. "The three
units are definitely segregated geographically, and the mor-
phological differences, when once noted, seem to be constant.
The principal characteristics may be summarized as follows:
C. confertiflora and C. flava have yellow flowers, while C. leuco-
phaea has white ones. Тһе nutlets in the first and last are defi-
nitely ovate in outline while in the second they are much nar-
rower. 'The inflorescence in confertiflora is distinctly glomerate
while in the related species it is continuous or nearly so.

Dr. Brand legally published the nomen nudem Oreocarya lutea
Greene. He did this in the belief that lutea Greene was specific-
ally distinct from confertiflora. He says ‘‘Macbride stellt in
seiner im Jahre 1916 erschienen Monographie der Gattung
Oreocarya den Namen O. lutea Greene als Synonym zu O. con-
fertiflora. Aber abgesehen von der goldgelben Blütenfarbe sind
bei O. lutea die Staubblütter an der Spitze des Kronentubus dicht
unter den Hohlschuppen eingefügt, wahrend sie bei О. confertiflora
(O. flava) in der mitte der Kronenrohre sitzen und von den Hohl-
schuppen durch einen deutlichen Zwischenraum getrennt sind."
These differences in position of the stamens in the tube are ob-
viously of no taxonomie value in the present case, since the
present species is very definitely dimorphic as regards stamen in-
sertion and length of style and both forms are commonly found
growing in close proximity.

6. C. flava (А. Nels.) new comb. Plate 25, figs. 11-13.
Oreocarya flava А. Nels. Bull. Torr. Bot. Club 25: 202. 1898.
О. lutescens Greene, Pittonia 4: 93. 1899. (Туре: “соттоп
on hills about Aztec, New Mexico, 25 April, 1899," C. F. Baker.)
Very similar in habit of growth to C. confertiflora; stems 15-85
em. tall, bristles somewhat соатзег and more widely spreading
than in confertiflora; leaves linear or linear-oblanceolate, acute,
3-9 em. long, dorsal surface densely appressed-strigose and with
numerous small appressed bristles with pustulate bases, ventral
surface uniformly strigose in age, with rather numerous pustulate
hairs, petioles ciliate; inflorescence usually restricted to the upper

[Vor. 14
260 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

6-34 of the stem, scarcely glomerate, indument very similar to
that in confertiflora, possibly more strongly setose; calyx setose
with rather strong divaricate bristles, sepals in anthesis linear-
lanceolate or quite linear, acute, about 10 mm. long, in fruit
scarcely enlarged, exceeding the nutlets by about 6 mm.; corolla
yellow, tube 9-11 mm. long, 2-3 mm. longer than the sepals,
crests at the base of the tube usually obsolete, rarely vestigial,
fornices very similar to those in confertiflora, limb 8-9 mm. broad,
lobes united for about 14 their length, less than 1% as long as the
tube; fruit lance-ovoid, usually less than 4 nutlets maturing (often
only 1 or 2), style exceeding the nutlets 2-7 mm. (species con-
spicuously heterostyled) ; nutlets lanceolate in dorsal view, acute,
3—4 mm. long, firmer in texture than in confertiflora, margins in
contact, acute, not wing-margined, surfaces of nutlets glossy,
smooth, dorsal somewhat more rounded crosswise than in con-
fertiflora, ventral not so distinctly keeled, scar similar.

Distribution: Upper Sonoran Zone in southern Wyoming,
western Colorado, northwestern New Mexico, eastern Utah, and
northeastern Arizona. Type: Point of Rocks, Sweetwater Co.,
Wyoming, June 1, 1897, A. Nelson 3074.

Specimens examined:

Wyoming: Alcova, Natrona Co., July 1, 1901, Goodding 164,
(R.Mt., Pomona, Field, Mo., U.S., Gray); hills of the Platte in
the mountains, Nuttall (Phila.); Cooper Creek, June 18, 1892,
Nelson 22 (Gray); Medicine Bow, July 9, 1898, Е. Nelson 4397
(R.Mt., Field); T. B. Ranch, Carbon Co., June 20, 1901, Goodding
53 (R.Mt.); Ft. Steele, May 25-June 10, 1901, Tweedy 4262
(U.S.); 3 miles north of Saratoga, July 3, 1922, Payson & Payson
2533 (R.Mt., Pomona, Colo., Mo., Gray); Ft. Steele, June 16,
1907, Nelson (R.Mt.); northeast corner of Sweetwater Co., July 6,
1926, Nelson 10693 (R.Mt.); Bitter Creek, June 16, 1898, Nelson
4771 (R.Mt.); Bitter Creek, June 2, 1897, Nelson 3098 (R.Mt.);
Point of Rocks, June 1, 1897, Nelson 3074 (R.Mt., TYPE);
Steamboat Mt., Sweetwater Co., June 9, 1900, Nelson 7067
(R.Mt., Pomona, Minn., Mo., U.S., Gray); 20 miles east of Point
of Rocks, July 4, 1922, Payson & Payson 2557 (R.Mt.); near
Leucite Hills, June 17, 1901, Merrill & Wilcox 497 (R.Mt., U.S.,
Gray); same locality and date, 486 (U.S.); near Washington's

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 261

Ranch, June 30, 1901, Merrill & Wilcox 726 (R.Mt., U.S., Gray);
Rock Springs, July 7, 1925, Payson & Payson 4307 (R.Mt.);
Green River, June 23, 1896, Jones 6725 (Pomona).

Colorado: 8-10 miles west of Craig, June 17, 1925 Osterhout
6389 (R.Mt.); Grand Junction, May 17, 1892, Eastwood (Calif.,
Mo., U.S., Gray); Grand Junction, May 28, 1894, Crandall
(Calif.); Grand Junction, June 20, 1898, Bethel (Colo.); Grand
Junction, June 19, 1915, Macbride & Payson 705 (R.Mt., Gray);
Dolores River near Mesa Creek, June 11, 1914, Payson 405 (R.
Mt., Gray).

New Mexico: Aztec, April, 1899, Baker 562 (Pomona, R.Mt.,
Calif., Mo., Gray); Fort Defiance, 1869, Palmer (U.S.).

Utah: Uinta Mts., 1902, Langille 117 (U.S.); Myton, May 20,
1908, Jones (Pomona) ; Theodore to Myton, May 19, 1908, Jones
(Pomona); between the K. Ranch and Jensen, June 18, 1925,
Osterhout 6406 (R.Mt.); Price, June 10, 1900, Stokes (Calif.);
Green River, June 12, 1901, Stokes (Minn., U.S.); Green River,
May 8, 1909, Tidestrom 2031 (U.S.) ; Thompson's Springs, May 3,
1891, Jones (Pomona); Thompson's Springs, May 7, 1891, Jones
(R.Mt.); Moab and vicinity, July 1-2, 1911, Rydberg & Garrett
8431 (N.Y.); Cisco, May 2, 1890, Jones (Pomona, R.Mt., Calif.);
San Rafael Swell, May 15, 1914, Jones (Pomona); in clay, near
Orangeville, June 18, 1894, Jones 5464 (Pomona, Calif., Mo.,
0.5.); 2 miles north of Ferron, June 18, 1894, Jones 5455c (U.S.,
Pomona).

Arizona: 5 miles east of Tuba, May 30, 1901, Ward (N.Y.,
0.$.); plants of the Hopis, Voth 103, 15 (Field); Adamana,
Petrified Forest, June 27, 1913, Hitchcock 7 (U.S.).

The interpretation here given to flava is somewhat more in-
clusive than that in Macbride’s revision inasmuch as Macbride
referred all plants from western Colorado to confertiflora while
they are now placed in flava. The characters ascribed to the
two species by Macbride are those accepted here. The discrep-
ancy in range is probably due to the comparative immaturity and
paucity of the specimens cited by Macbride from Colorado. For
a discussion of the distinction between flava and related species
see under C. confertiflora.

[Vor. 14
262 ` ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

7. C. leucophaea (Dougl.) new comb. Plate 25, figs. 14-16.

Myosotis leucophaea Dougl. in Lehm. Pug. 2:22. 1830.

Eritrichium leucophaeum (Dougl.) A.DC. Prodr. 10: 129. 1846.

Krynitzkia leucophaea (Dougl.) Gray, Ргос. Am. Acad. 20: 280.
1885.

Oreocarya leucophaea (Dougl.) Greene, Pittonia 1: 58. 1887.

Very similar in habit of growth to C. confertiflora; stems 15-40
ст. tall, white-hairy at the base, densely strigose but scarcely
getose upwards except in the inflorescence; leaves linear to linear-
oblanceolate, acute, 3-9 ст. long, dorsal surface densely strigose
and with numerous weak bristles with pustulate bases, ventral
surface uniformly strigose, pustules few or none, leaf-bases and
petioles ciliate; inflorescence mainly restricted to upper Уз of the
stem, scarcely glomerate, indument rather softly setose, bristles
divaricate, white, not turning yellowish in age; calyx setose with
rather weak bristles, sepals in anthesis linear or nearly so, acute,
about 10 mm. long, in fruit 13-14 mm. long, exceeding the nutlets
by 7-8 mm.; corolla white, tube 8-11 mm. long, 1-2 mm. longer
than the sepals, crests at the base of the tube evident, fornices
linear-oblong, emarginate, 1 mm. long, probably yellow, limb
8-10 mm. broad, lobes united about l4 their length, less than
half as long as the tube; fruit ovoid, usually less than 4 nutlets
maturing, style exceeding the mature nutlets 1.5-7 mm. (species
heterostyled); nutlets ovate, acute, 3-4 mm. long, rather thin,
margins in contact, acute, almost wing-margined (less so than in
C. confertiflora, more than in C. flava), surfaces of nutlets glossy,
smooth, scar as in C. confertiflora.

Distribution: Upper Sonoran Zone in south-central Washing-
ton—reported from southern British Columbia (Macoun, Cat.
Canad. Pl. 2: 338. 1884, and Henry, Fl. Southern British
Columbia, 254. 1915), and probably to be found in northern
Oregon. Туре: “arid barrens of the Columbia, and of its north-
ern and southern tributaries." Collected by Douglas.

Specimens examined:

Washington: “ех dupl. Hook." Douglas (Gray); Columbia
River from lat. 46° to 49? N., 1860, Lyall (Gray); near Morgan's
Ferry, Yakima River, June 8, 1884, Suksdorf 407 (Gray); near
Egbert Spring, Douglas Co., July 4, 1893, Sandberg & Leiberg 373

1927]
PAYSON—SECTION OREOCARYA OF СКУРТАМТНА 263

(Wash., Pomona, R.Mt., Calif., Minn., Field, Mo., Phila., U.S.,
Gray); Pasco, May 26, 1899, Piper 2987 (Wash.); Pasco, July 11,
1897, Piper (R.Mt.); Pasco, May 20, 1899, Piper 2987 (Gray);
Pasco, May 25, 1896, Hindshaw 2 (Wash.); Pasco, July, 1898,
Elmer 1056 (Wash., Minn., U.S.); Scott, Klickitat Co., May 16,
1898, Leckenby (Wash.); 2 miles s.w. of Attalia, May 6, 1911,
Beattie 3921 (Wash.); Burbank, Walla Walla Co., May 29, 1922,
Lechiner 3 (Wash.); Walla Walla region, June, 1883, Brandegee 997
(Phila., Calif., Gray); Wallula, May 23, 1903, Cotton 1027 (Wash.,
U.S., Gray).

For a discussion of the differences between this species and the
closely related ones see under C. confertiflora. C. leucophaea is
of especial interest because it is so widely separated geographically
from its very near allies.

8. C. salmonensis (Nels. & Macbr.) new comb.
Plate 25, figs. 17-19.
Oreocarya salmonensis Nels. & Macbr. Bot. Gaz. 61: 43. 1916.
Perennial and caespitose; stems rather stout, 1.5-3 dm. high,
setose with rather weak, white hairs, as well as strigose-tomentu-
lose; leaves rather narrowly oblanceolate to spatulate, clustered
at the base, numerous upwards, acute or obtuse, more or less
tomentulose on both surfaces, setose with rather weak hairs,
pustulate dorsally and ventrally; inflorescence on upper 1% of the
stem, rather narrow or sometimes broader by the elongation of
the cymules, foliar bracts not conspicuous; calyx abundantly
setose with white, rather slender hairs; sepals in anthesis linear-
lanceolate, acute, about 3 mm. long, in fruit 5-7 mm. long, ex-
ceeding the nutlets by about 3 mm.; corolla white, tube 3 mm.
long, equalling or slightly shorter than the sepals, slightly shorter
than the limb, crests at the base of the tube well developed, for-
nices distinctly papillose, probably yellow, 0.5-1 mm. long,
emarginate, limb 8-10 mm. broad, lobes united for about 4
their length; fruit lanceolate-ovoid, 3—4 nutlets usually maturing,
style 1.5 mm. longer than the mature nutlets; nutlets lanceolate,
obtuse, 3-4 mm. long, with margins in contact, rather indefinitely
wing-margined, all surfaces smooth, glossy, scar straight, closed,
extending the full length of the nutlets, no elevated margin evi-
dent.

[Уоь. 14
264 ANNALS ОҒ THE MISSOURI BOTANICAL GARDDN

Distribution: Upper Sonoran areas in central Idaho. Туре:
Charles L. Kirtley, Salmon, Idaho.

Specimens examined:

Idaho: Salmon, June, 1896, Kirtley (R.Mt., TYPE); Salmon,
July 3, 1920 Payson & Payson 1880 (R.Mt., Mo., Gray); Challis,
July 21, 1916, Macbride & Payson 3348 (R.Mt., U.S., Mo., Gray).

There can be no doubt as to the right of this plant to specific
recognition. It is perhaps most closely related to C. leucophaea
but is at once separable from that by the short corollas, the more
hispid indument, and the broader leaves.

9. C. stricta (Osterhout) new comb. Plate 26, figs. 20-22.

Oreocarya stricta Osterh. Bull. Torr. Bot. Club 50: 217. 1923.

Perennial; stems solitary or 2 or 3 from а tap-root, rather
slender, strictly erect, 1.5-3 dm. high, conspicuously setose with
strong, divaricate bristles and sparsely retrorse-strigose; leaves
clustered at the base of the stem, oblanceolate, usually acute,
2—5 cm. long, conspicuously veined longitudinally, strongly setose
with spreading bristles and strigose pustulate hairs about equally
numerous on both leaf surfaces, cauline leaves similar to the
basal, reduced upwards; inflorescence apparently rather narrow
and crowded, extending over the upper 14 or 14 of the stem,
densely setose, foliar bracts quite small and inconspicuous; calyx
strigose and setose with spreading bristles, sepals narrowly
lanceolate, acute, 4-5 mm. long in anthesis, 6-8 mm. long in
fruit, exceeding the nutlets by about 3 mm.; corolla quite yellow
in dried material but Mr. Osterhout believes it to have been
white with yellow fornices, tube 3.5 mm. long, equalling or shorter
than the sepals, crests well developed at the base of the tube,
fornices low, rounded, limb 8-10 mm. broad, tube and limb sub-
equal, lobes united for 14 to 15 their length; fruit lanceolate-ovoid,
all four nutlets usually maturing, style exceeding the mature
nutlets 1-1.5 mm.; nutlets lanceolate or elliptical, obtuse or
subacute, with margins in contact, narrowly winged, acute,
surfaces of nutlets glossy, the dorsal definitely rugose and
sometimes tuberculate, not muricate, ventral surfaces smooth or
nearly so, scar straight, extending from the base to near the apex,
nearly closed, no elevated margin present.

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 265

Distribution: Upper Sonoran Zone, northwestern Colorado.
Type: collected ‘‘some distance south of the Yampa or Bear
River in Moffat PURI (Colorado) along the Victory Highway."

Specimens examine

Colorado: some distina south of the Yampa River along the
Victory Highway, June 21, 1922, Osterhout 6195 (R.Mt., Osterh.,
TYPE, Gray).

C. stricta is a very distinct species and although only known at
present from a single collection, must take its place as one of the
major units in any study of the genus. It is difficult to say to
what species it is most closely allied. Тһе nutlet characters
would place it nearest to C. nubigena but it is more than probable
that these similarities represent merely a convergent development
rather than an actual immediate relationship. С. stricta may
represent a separate line of evolution from C. James var.
multicaulis or an offshoot from the confertiflora-flava branch.

10. C. nubigena (Greene) new comb. Plate 26, figs. 23-25.

Oreocarya nubigena Greene, Pittonia 3: 112. 1896

Perennial and caespitose from a woody root; stems several,
slender or rather stout, 16—20 cm. high, setose and hirsute, bristles
divaricate; leaves oblanceolate to spatulate, obtuse, tapering
gradually to а somewhat ciliate petiole, 2-4 cm. long, setose with
curved bristles, hirsute and more or less tomentulose, pustulate
bases to the bristles present on both surfaces, more abundant and
earlier evident on the dorsal surface; inflorescence confined to the
upper L4 or less of the stem, narrow, somewhat glomerate or
compact, foliar bracts inconspicuous, densely setose; calyx setose
with divaricate or somewhat reflexed bristles, hirsute, sepals in
anthesis linear-lanceolate, somewhat obtuse at the very tip,
3-4 mm. long, in fruit 7-9 mm. long, exceeding the nutlets by
2-3 mm.; corolla white, tube 3-4 mm. long, equalling the calyx
lobes, erento at base of the tube evident, fornices prominent,
broad, рарШове, yellow, about 0.5 mm. high, limb 4-7 mm. broad,
lobes slightly shorter than the tube (measuring from the fornices),
united for about L4 their length; fruit elongated and narrowly
ovoid, all 4 nutlets commonly maturing, style equalling the
mature nutlets or exceeding them 0.5-1.5 mm.; nutlets narrowly

[Vor. 14
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lanceolate, acute or obtuse, 4-5 mm. long, somewhat glossy,
margins in contact, acute, apices spreading, dorsal surface of nut-
lets nearly smooth or (usually) indefinitely rugose and tubercu-
late, somewhat margined, ventral surface smooth or indistinctly
wrinkled, scar narrow, straight, extending from near the base
nearly to the apex.

Distribution: in the high mountains, south-central Idaho,
northwestern Nevada, Oregon and northern and eastern Cali-
fornia. Туре: “Оп Cloud's Rest, Mariposa Co., California, 10
July, 1889," Messrs. Chestnut & Drew.

Specimens examined:

Idaho: Smoky Mts., Blaine Co., Aug. 13, 1916, Macbride &
Payson 3771 (R.Mt., Mo., U.S., Gray).

Nevada: Santa Rosa Mts., July 11, 1898, Cusick 2028 (R.Mt.,
Calif., Minn., Mo., U.S., Gray).

Oregon: Pine Creek, Baker Co., Sept. 1879, Cusick (Gray);
dry banks above the John Day River, Prairie City, Grant Co.,
July 1, 1919, Ferris & Duthie 734 (R.Mt.); Warner Mts., July
1898, Austin & Bruce 2270 (Pomona); Crater Lake, Sept. 14,
1902, Coville 1514 (U.S.).

California: Scott’s Mt., north Calif., Aug. 30, 1880, G. Engel-
mann (Mo.); Mt. Eddy, Siskiyou Co., July 9, 1920, Heller 18435
(Field, N. Y., Mo., U.S.); head North Fork, Parker Creek,
Warner Mts., Modoc Co., July 13, 1910, Taylor & Bryant (Calif.) ;
mountain near Sonora Pass, July 16, 1863, Brewer 1887 (Calif.) ;
summit of Cloud's Rest, July 13, 1889, Chestnut & Drew (Calif.).

It is not felt that the treatment here accorded nubigena is
particularly satisfactory. Тһе specimens at hand are com-
paratively few and from widely separated localities. They are
not particularly homogeneous among themselves. It is possible
that the species may be separable into several geographie varieties
when it is better known. Perhaps it is simply an especially
polymorphic unit and any one locality will give great extremes of
variation. Even its relationship to other species is not clear.

Because of its range and habitat, nubigena is most likely to be
confused with humilis. 'That it is really closely related to it is
not so certain. Тһе outstanding peculiarities of nubigena are its
greatly elongated nutlets that are nearly smooth on the inner sur-

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 267

faces. А satisfactory concept of this species must await further
study and exploration.

11. C. Clemensae Payson! Plate 26, figs. 26-28.

Perennial, probably short-lived; stems numerous from the
rather slender root, the underground portions densely clothed
with the leaves of previous years, very slender, 6-12 cm. high,
setose; leaves narrowly oblanceolate or spatulate, rather thin,
2-3 em. long, obtuse or subacute, rather sparsely hairy with
spreading, hirsute trichomes and small, pustulate setae, both
surfaces pustulate, the upper much less densely so than the lower;
inflorescence subcapitate, rarely over 3 cm. long, foliar bracts in-
conspicuous; calyx densely setose with rather slender bristles,
rather sparsely strigose, sepals in anthesis 3-4 mm. long, linear-
lanceolate, in fruit about 7 mm. long, exceeding the nutlets by
3-4 mm.; corolla white, tube 2.5 mm. long, shorter than the
sepals in anthesis, crests at the base of the tube evident, fornices
well developed, probably white, 0.5-1 mm. long, scarcely papil-
lose, limb about 4 mm. broad, tube distinctly longer than the
limb, lobes united for about 4 their length; fruit elongated and
narrowly ovoid, all four nutlets commonly maturing, style ex-
ceeding the mature nutlets by 0.5-1 mm., margins in contact or
nearly so, acute; nutlets somewhat papery, linear-lanceolate,
acute, 3 mm. long, very definitely but narrowly wing-margined,
surfaces slightly glossy, the dorsal sparsely tuberculate, the
tubercles low and sometimes elongated but apparently not often
forming rugae, ventral surfaces nearly smooth except for the
veining of the pericarp, scar straight, extending from near the
base nearly to the apex, open but narrow, no elevated margin
present.

1 Cryptantha Clemensae sp. nov., perennis; caulibus multis gracilibus setosis
6-12 cm. altis; foliis anguste oblanceolatis aut spathulatis tenuibus obtusis aut
subacutis hirsutis et setosis supra et subter pustulosis; асын subcapitatis;
sepalis setosis linearo-lanceolatis 3-4 mm. longis, fructiferis ca. 7 mm. longis, quam
nuculae 3-4 mm. 27 corolla alba, tubo 2.5 mm. КЕ quam sepalis brevi-
oribus, limbo ca. 4 mm. ; stylo nuculas 0.5-1 mm. superante; nuculis linearo-
lanceolatis acutis 3 mm. n anguste alatis, facie exteriore sparse tuberculosis,
faciebus ventralibus fere laevibus, sulco recto angusto, margine non edito.—Col-
lected by Mrs. Joseph Clemens, Glenn's Pass, California, July 22, 1910, (R. Mt. Herb.,
TYPE).

[Vor. 14
268 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

Distribution: in the southern Sierra Nevada of California.
Type: Glenn's Pass, California, July 22, 1910, Mrs. Joseph
Clemens.

Specimens examined:

California: Glenn's Pass, July 22, 1910, Clemens (R.Mt.,
TYPE, Pomona); Mt. Whitney, July 11, 1910, Clemens (Pomona);
near Mt. Whitney, Sept. 1, 1891, Bailey, Coville & Funston
"DRY Mt. Whitney, alt. 13,000 ft., Aug. 17, 1899, Copeland 52

Presumably this species is most closely related to C. nubigena,
and its geographical location and habitat make it reasonable to
suppose that it is a derivative from that plant. From nubigena,
С. Clemensae is very different in general appearance. Тһе thin,
green and rather flaccid leaves give it an appearance that is quite
unusual in the genus. Тһе small, rather turgid but narrowly
winged nutlets are also quite characteristic. It seems as distinct
а unit as has been described in the genus; indeed the author's
chief hesitation in describing the plant has been because it seemed
so aberrant in the section Oreocarya that it might be best placed
in some other group. А study of the flowers and fruit seems to
make it certain that it must be placed in Cryptantha, and the
perennial root and compact inflorescence definitely place this
plant in the section Oreocarya.

The species is named in honor of the indefatigable collector,
Mrs. Joseph Clemens.

Series 2. VriRGATAE. Stout, erect biennials or perennials
with unbranched stems. Nutlets smooth, or nearly so, on the
ventral surfaces, variously roughened on the dorsal — or in some
forms of C. virgata — quite smooth. Species 12-13.

12. C. setosissima (Gray) new comb. Plate 26, figs. 29-31.
Eritrichium setosissimum Gray, Proc. Am. Acad. 12:80. 1877.
Krynitzkia setosissima Gray, Proc. Am. Acad. 20: 276. 1885.
Oreocarya setosissima (Gray) Greene, Pittonia 1: 58. 1887.
Short-lived perennial or possibly a biennial from a stout tap-
root; stem usually solitary, unbranched, stout, 2.5-8 dm. tall,
setose and hirsute; leaves clustered at the base, linear-oblance-
olate, obtuse, the lower 6-12 сш. long, somewhat reduced up-

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 269

wards, setose and hirsute, bristles not so coarse nor so widely
spreading as in C. virgata, pustulate hairs numerous on both sur-
faces but more numerous dorsally ; inflorescence rarely extending
over more than 1% of the stem, foliar bracts scarcely exceeding
the branches of the inflorescence, this elongating and spicate
rather than glomerate; calyx densely hirsute and setose; sepals
in anthesis lanceolate, acute or obtuse, about 6 mm. long, in
fruit linear-lanceolate, acute or obtuse, about 10 mm. long,
exceeding the nutlets by about 5 mm.; corolla white, tube about
4 mm. long, equalled by the calyx lobes, constricted above the
ovary with a thickened ring of crests, fornices distinctly emargin-
ate, 0.5 mm. high, probably yellow, limb about 8 mm. broad,
lobes united about '/, their length; fruit ovoid, all four nutlets
commonly maturing, style exceeding the nutlets by 1.5 mm.;
nutlets in contact, ovate, obtuse, 4-5 mm. long, papery rather
than bony, surrounded by a conspicuous wing-margin nearly 1
mm. wide, surfaces somewhat glossy, dorsal uniformly and openly
muriculate with a few larger tubercles or short rugae, ventral
surface smooth or nearly so, scar straight, narrow, open, extending
from very near the base to near the apex.

Distribution: Transition Zone from south-central Utah to
southern Arizona. Туре: “shores of Fish Lake, Utah, at 8,700
feet, L. F. Ward,’’ in Powell's Expedition, 1875.

Specimens examined:

Utah: Bromide Pass, Henry Mts., July 27, 1894, Jones 5692al
(U.S.); Panguitch Lake, Sept. 7, 1894, Jones 6015al (Pomona,
U.8.); Fish Lake, Aug. 9, 1894, Jones 5790m (Pomona, U.S.);
Fish Lake, Aug. 10, 1894, Jones 5812 (Pomona, R.Mt., Calif.,
Mo., U.S.); Fish Lake, Aug. 25, 1875, Ward 646 (Phila., Mo.,
U.S., Gray, TYPE); St. George, 1877, Palmer 857 (Mo., Gray).

Arizona: Grand Canyon of the Colorado, June 26, 1898, Mac-
Dougal 165 (R.Mt., Calif., Phila., U.S., Gray); Grand Canyon,
July 9, 1892, Тоитеу 224 (U.S.); rim of Grand Canyon, July 12,
1892, Wooton (U.S.); San Francisco Mts., Sept. 1884, Lemmon &
wife (Calif.); San Francisco Peaks, July 26, 1901, Leiberg 5748
(U.S.); Flagstaff, Aug. 7, 1884, Jones 6686 (Pomona); Flagstaff,
May-Oet. 1900, Purpus 8048 (Pomona, Calif., Mo., 0.8.);
Flagstaff, Aug. 1883, Rusby 748 (Calif., Phila., 0.5.); Mt. Hum-

[Vor. 14
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN

phrey, July 26, 1897, Kunze (N.Y.); Bill Williams Mt., July 5,
1889, Greene (Calif., U.S.) ; Nagle’s Ranch (northwestern Arizona),
Sept. 17, 1894, Jones 6054s (U.S.); Ft. Apache, June 21-30, 1890,
Palmer 591 (Calif., Gray, U.S., in part); 66 miles south of Hol-
brook, July 4, 1901, Hough 88 (U.S.); Thompson’s Ranch, Black
River, White Mts., July 14, 1910, Goodding 589 (R.Mt., U.S.,
Gray); White Mts., Aug. 6-15, 1903, Griffiths 5300 (U.S.); near
Santa Catalina Mts., Aug. 1881, Lemmon & wife (R.Mt.).

This is certainly one of the most distinct units in the section
Oreocarya. It is remarkably uniform in character and is to be
confused with no other species. In general appearance it is
somewhat similar to C. virgata, and for that reason is associated
with it in the series. This resemblance may be no indication of
real relationship. The nutlets of certain annual species of
Cryptantha, notably C. utahensis, C. holoptera, and C. pterocarya,
are remarkably similar to those of С. setosissima and suggest the
possibility that some of the annual forms of the genus may have
developed from perennial forms in North America. It was doubt-
less this similarity of nutlets that led Greene (Pittonia 1: 58.
1887) to transfer one of the annual species to Oreocarya. This
was О. holoptera (Gray) Greene; now treated as Cryptantha
holoptera (Gray) Macbride.

13. C. virgata (Porter) new comb. Plate 26, figs. 32-34.

Eritrichium virgatum Porter, Hayden Rept. 479. 1870.

E. glomeratum var. virgatum Porter in Porter & Coulter, Syn.
Fl. Colo. 102. 1874.

Krynitzkia virgata (Porter) Gray, Proc. Am. Acad. 20: 279.
1885.

Oreocarya virgata (Porter) Greene, Роща, 1: 58. 1887.

О. spicata Rydb. Bull. Torr. Bot. Club 36: 678. 1909. (Туре:
Artist's Glen, Pikes Peak, Colorado, Aug. 1, 1901, Clements 102.)

O. virgata forma spicata (Rydb.) Macbr. Proc. Am. Acad. 51:
546. 1916.

Biennial from a stout taproot; stem usually solitary, un-
branched, stout (when the terminal bud is injured a number of
rather slender stems may be developed), 2.5-7 dm. tall, densely
setose with long, stout, divaricate bristles, hirsute; leaves narrowly

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 271

oblanceolate, obtuse, 3-12 ст. long, hirsute and setose, appressed
pubescence lacking, pustulate hairs numerous on both surfaces
but somewhat more numerous dorsally; inflorescence usually
extending along at least 34 of the stem, uniformly cylindrical in
outline, conspicuously bracteate with linear-oblanceolate foliar
bracts that much exceed the cymules in length; calyx setose and
hirsute, indument not essentially different from that of the leaves,
sepals in anthesis lanceolate, acute, 4 mm. long, in fruit linear-
lanceolate, acute or slightly obtuse, 11 mm. long, exceeding the
nutlets by about 6 mm.; corolla white, tube about 3.5 mm. long,
slightly exceeded by the calyx lobes, with a conspicuous thickened
ring of crests just above the ovary, fornices papillose, yellow,
distinctly emarginate, 0.5 mm. high, limb 8-10 mm. broad, lobes
united about 15 their length; fruit broadly ovoid, all four nutlets
commonly maturing, style exceeding the nutlets 1.5-2 mm.;
nutlets in contact, ovate, obtuse, 2.5-3.5 mm. long, with an in-
distinct thinner margin, surfaces somewhat glossy, dorsally
sparingly tuberculate and usually more or less rugose with low,
rounded rugae (smooth in forma spicata), ventrally nearly
smooth, often with a few indistinct tubercles or rugae, scar
straight, nearly or quite closed, extending from about 15 the
distance from the base to about the same distance from the apex.

Distribution: Transition Zone, eastern foothills of the Rocky
Mountains, southeastern Wyoming to south-central Colorado;
also in North Park, Colorado. Туре: “пеаг Denver, Colorado
Territory, 1869," B. H. Smith.

Specimens examined:

Wyoming: Sheep Mt., Albany Co., June 24, 1925, Payson &
Payson 4248 (R.Mt.); Chug Creek, Albany Co., June 29, 1900,
Nelson 7338 (R.Mt., Pomona, Minn., Mo., U.S., Gray); Tele-
phone Canyon, Albany Co., June 15, 1894, Nelson 281 (R.Mt.,
Minn., Mo., U.S., Gray); Laramie Hills, June 13, 1896, Nelson
1937 (R.Mt., Minn., Mo.); Centennial Valley, June 8, 1895,
Nelson 1267 (R.Mt., Wash.).

Colorado: Walden, June 25, 1925, Payson & Payson 4253
(R.Mt.); King’s Canyon, Jackson Co., June 25, 1925, Payson &
Payson 4291 (R.Mt.); Horsetooth Gulch, June 30, 1893, Baker
(Pomona); foothills, Larimer Co., May 25, 1895, Osterhout

[Уог. 14
272 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(R.Mt.); Front Range, July 2, 1896, Crandall (R.Mt., Wash.,
СаШ., Mont.); Estes Park, July 12, 1904, Cooper 117 (R.Mt.);
St. Vrain Creek, June 9, 1906, Dodds 1832 (R.Mt.); Estes Park,
Aug. 1895, Osterhout (Minn.); Boulder, June 24, 1901, Osterhout
2463 (Pomona, Calif., Gray, Osterh.); between Tolland and Rol-
linsville, July 8, 1913, Overholts 10155 (Calif.); near Boulder,
June 10, 1905, Ramaley 1078 (R.Mt.) ; near Boulder, July 9, 1900,
Ramaley A.107 (R.Mt.); near Boulder, July, 1902, Tweedy 5222,
5222 (R.Mt.); near Boulder, June, 1903, Tweedy 5676 (R.Mt.);
Georgetown, July and Aug. 1885, Patterson 110 (Calif., Phila.,
Mo., U.S., Gray); near Golden, June 24, 1878, Jones 296 (Po-
mona); Green Mt. Falls, Ute Pass, Aug. 2, 1892, Sheldon 216
(U.S.); Colorado Springs, June 2, 1879, Jones (R.Mt., Pomona,
U.S.); Pikes Peak region, July 27, 1920, Johnston 2818 (Gray);
Artist's Glen, Pikes Peak, Aug. 1, 1901, Clements & Clements 102
(R.Mt., Minn. Mo., U.S., Gray); mountains n.e. of Canyon
City, 1874, Brandegee 898 (Mo.).

C. virgata is one of the most conspicuous herbaceous plants on
the eastern foothills of the Rocky Mountains in eastern Colorado.
The strictly erect, rod-like stems with the closely set white flowers
are unique. Тре numerous elongated leaf-like bracts of the in-
florescence add to the peculiar appearance and serve to separate
the species from its relatives. Its relationship to C. setosissima
is decidedly problematical. Although it was for a time held to
be a variety of Bradburiana there is no reason to believe it is
really related to that plant.

Additional evidence is at hand to support Macbride's conten-
tion (Proc. Am. Acad. 51: 546. 1916) that О. spicata Rydb. is
not even deserving of varietal rank. It is simply to be regarded
as а form of virgata. ‘Thanks to the help of Mr. W. T. Penfound,
of the Alpine Laboratory on Pikes Peak, specimens have been
examined that prove the existence of typical virgata from that
region as well as the smooth-fruited form.

Series 3. HuMiLAE. Nutlets distinctly and variously rough-
ened on all surfaces. Scar open and triangular, margins some-
times elevated. Corolla tubes never longer than the calyces.
Plants of Utah, Nevada, southwestern Idaho, and California.
Species 14-19.

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 273

14. C. insolita (Macbr.) new comb. Plate 26, figs. 35-37.

Oreocarya insolita Macbr. Contr. Gray Herb. 48: 28. 1916.

Biennial (or short lived perennial?) from a rather slender root ;
stems one to several, rather stout, erect, 1.5-4 dm. high, abun-
dantly setose and rather coarsely strigose; leaves clustered at the
base, spatulate, obtuse, 3—5 ст. long, dorsal surface subtomentose
and rather sparsely appressed-setose and pustulate, ventral sur-
face similar but setae smaller and fewer, pustules rather few and
small, petioles abundantly long-hairy at the base, otherwise
scarcely ciliate; inflorescence mostly confined to the upper 14 or
15 of the stem, cymes rather few (for the genus) and conspicu-
ously elongating, inflorescence abundantly but rather weakly
setose, bracts inconspicuous; calyx densely hirsute, conspicuously
setose with rather short weak bristles; sepals in anthesis linear-
lanceolate, acute, about 4 mm. long, in fruit about 8 mm. long,
exceeding the nutlets by about 4 mm.; corolla white, tube about
3 mm. long, equalling or somewhat shorter than the sepals,
crests at the base of the tube well developed, fornices 0.5-1 mm.
long, probably yellow, minutely рарШове, slightly emarginate,
limb about 7 mm. broad, lobes and tube subequal, lobes united for
about 14 their length; fruit broadly ovoid, 1-4 nutlets maturing,
style exceeding the nutlets by 1-1.5 mm.; nutlets ovate, obtuse or
acute, 4 mm. long, margins in contact or nearly so, acute, surfaces
of nutlets dull or slightly glossy, the dorsal more or less carinate,
tuberculate, granulo-muriculate and sometimes slightly rugose,
ventral surface tuberculate and somewhat rugose, scar narrow but
slightly open, the margin showing some tendency to be elevated.

Distribution: lower part of the Upper Sonoran Zone, southern
Nevada. Туре: Las Vegas, Nevada, May 4, 1905, Г. М.
Goodding 2286.

Specimens examined:

Nevada: Las Vegas, Мау 4, 1905, Goodding 2286 (R.Mt.,
Minn., Mo., Gray, түре); Las Vegas, К. Brandegee (Calif.);
Las Vegas, April 29, 1905, Jones (Pomona).

C. insolita is probably most elosely related to C. virginensis and
C. tumulosa. Тһе differences between them, which it will be
noticed are mostly those of habit rather than of nutlet peculi-
arities, may be summarized in the following way:

[Vor. 14

274 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN
C. tumulosa C. insolita C. virginensis
Perennial. Biennial. Biennial.
Stems 1-2.5 dm. Stems 1.5-4 dm. Stems 1.5-4 dm.
high. high. high

Leaves setose below, Leaves inconspicu- Leaves conspicuous-
with subappressed ously setose below ly setose below

bristles. with appressed with ^ spreading
bristles. bristles.
Inflorescence narrow, Inflorescence broad, Inflorescence broad,
cymes short. cymes elongating, cymes elongating,
very numerous, few, and flowers
flowers crowded. rather distant.
Nutlets slightly Nutlets distinctly Nutlets ^ distinctly
carinate. carinate. carinate.

Dorsal surface with Dorsal surface with Dorsal surface with
low indefinite tu- short definite tu- sharp definite tu-
bercles. bercles. bercles.

In general appearance this species is very similar to C. elata
of Grand Junction, Colorado. However, it is doubtful if the two
species are closely related.

15. C. virginensis (Jones) new comb. Plate 27, figs. 38-40.

Krynitzkia glomerata var. virginensis Jones, Contr. West. Bot.
13: 5. 1910.

Oreocarya virginensis (Jones) Macbr. Proc. Am. Acad. 51: 547.

16.

Biennial from a rather stout taproot; stems 1-many from the
base, stout, 15-40 cm. high, conspicuously setose with divaricate
bristles; leaves oblanceolate or spatulate, obtuse, 5-12 ст. long,
rather sparsely setose, slightly hirsute and distinctly subtomentose
on the dorsal surface, abundantly pustulate, ventral surface
rather sparsely pustulate, weakly appressed-setose and subto-
mentose, at times subsericeous; inflorescence extending over 34
or more of the stem, very floriferous, not interrupted, becoming a
broad thyrsus in which the individual cymes are much elongated,
densely setose, in age usually fulvous, lower foliar bracts often
conspicuous and somewhat reflexed; calyx densely setose, sepals
in anthesis lanceolate, about 4 mm. long, acute, in fruit linear,

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 275

10-12 mm. long, exceeding the nutlets by about 6 mm.; corolla
white, tube 3-4 mm. long, equalling or slightly shorter than the
sepals, crests at base of tube well developed, fornices conspicuous,
probably yellow, nearly 1 mm. long, oblong, emarginate, conspicu-
ously papillose, limb 6-8 mm. broad, lobes and tube subequal,
lobes united for about 15 their length; fruit ovoid, 1-2 nutlets
usually maturing, style exceeding nutlets by about 1 mm.;
nutlets lance-ovate, obtuse, about 4.5 mm. long, margins in
contact, acute, surfaces of nutlets somewhat glossy, the dorsal
usually distinctly carinate, sharply tuberculate and somewhat
rugose, surface between the tubercles uneven, ventral surface very
uneven with indeterminate rugae and tubercles, scar narrowly
triangular, some tendency evident to an elevated margin.

Distribution: lower part of Upper Sonoran Zone, southwestern
Utah, southern Nevada, southeastern California. Туре: Га
Verken, Utah, May 8, 1894, M. Е. Jones 5195а.

Specimens examined:

Utah: Valley of the Virgin near St. George, 1874, Parry 173
(Field, N.Y., Gray); Diamond Valley, Мау 16, 1902, Goodding
830 (R.Mt., Pomona, Мо., U.S., Gray); Diamond Valley, April
28, 1894, Jones 5125 (R.Mt., Pomona, СаШ., Mo., U.S.); La
Verken, May 8, 1894, Jones 5195a (Pomona, түре, R.Mt., Calif.,
Mo., 0.8.).

Nevada: Muddy Valley, Lincoln Co., May 16, 1906, Kennedy
& Goodding 78 (Calif., Mo., U.S.); Vegas Valley, Lincoln Co.,
May 1, 1891, Bailey, Coville & Funston 1888 (U.S.); Las Vegas,
К. Brandegee (Pomona); Gold Mountain, 1898, Purpus (СаШ.);
Indian Spring, Charleston Mts., May 7, 1906, Jones (Pomona);
Good Springs, April 30, 1905, Jones (Pomona); Tonopah, April
24, 1907, Jones (Pomona); Amargosa Desert, April 27, 1907,
Jones (Pomona).

California: Panamint Canyon, May 4, 1897, Jones 6728
(Pomona) ; above Barstow, April 18, 1921, Jaeger 1113 (Pomona).

С. virginensis is in general appearance more like C. thyrsiflora
than any other species of the genus. It is not likely to be con-
fused with that species because of the great distance between
their ranges. Тһе nutlets of the two species are entirely dif-
ferent, and it seems quite doubtful if they are in reality closely

[Vor. 14
276 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

related. Its nearest relatives are probably C. tumulosa, C. mo-
desta, and C. insolita. Тһе distinguishing characteristics of these
species are contrasted under C. insolita.

16. C. tumulosa (Payson) new comb. Plate 27, figs. 41-43.

Oreocarya tumulosa Payson, Univ. Wyo. Publ. Bot. 1:164. 1926.

Long-lived caespitose perennial from a woody root; stems few
to many from a branching caudex, rather stout, 10-25 em. high,
hirsute and densely setose with divaricate bristles; leaves numer-
ous near the base, oblanceolate, obtuse, 3-5 cm. long, blade
gradually narrowed into a long, slender, scarcely hirsute petiole,
lower leaves conspicuously tomentulose, rather sparsely setose
with weak, appressed bristles, dorsal surface slightly more pus-
tulate than the ventral; inflorescence extending over l4 to 34
of the stem, floriferous, rather narrow, uninterrupted, densely
setose with yellowish bristles (at least in age), foliar bracts incon-
spicuous, reflexed; calyx densely divaricate or retrorse-setose,
hirsute, sepals in anthesis linear-lanceolate, acute, about 4 mm.
long, in fruit 8-10 mm. long, exceeding the nutlets by 4—6 mm.;
corolla white, tube 3.5-4 mm. long, equalling or slightly shorter
than the sepals, crests at the base of the tube evident but not
conspicuous, fornices conspicuous, probably yellow, about 1 mm.
long, limb 7 mm. broad, limb and tube subequal, lobes united
for about 1$ their length; fruit asymmetrical, 1-2 nutlets only
maturing, style exceeding the nutlets by about 0.5 mm.; nutlet
ovate-lanceolate in outline, obtuse, 4 mm. long, very pale in color,
dull or slightly glossy, margins in contact, acute, dorsal surface
of nutlets with a low but evident medial ridge, indefinitely tuber-
culate and sometimes indistinctly rugose, ventral surface rough-
ened with indefinite tubercles and rugae, scar triangular, open,
short, margin only slightly elevated.

Distribution: Upper Sonoran Zone on or near the Providence
Mountains, San Bernardino Co., California. Type: Providence
Mts., May, 1902, T. S. Brandegee.

Specimens examined:

California: Ivanpah Mts., San Bernardino Co., June 4, 1915,
Parish 10243 (Calif.); Providence Mts., May, 1902, Т. S. Bran-
degee (Calif., түре); Providence Mts., May, 1892, T. S. Brandegee

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 277

(Phila.) ; Barnwell, May 14, 1911, К. Brandegee (Pomona, Calif.) ;
vicinity of Bonanza King Mine, east slope of Providence Mts.,
May 21-24, 1920, Munz, Johnston & Harwood 4209 (R.Mt.,
Pomona, Calif.).

This species has been confused with С. humilis and C. nubigena.
Тһе nutlets are quite different from either of those species. Its
nearest relatives are probably C. virginensis, C. insolita and С.
modesta. It is contrasted with the first two in the discussion of
C. insolita. From C. modesta it differs in the lack of an elevated
margin around the scar and by the indefinite roughening on the
dorsal surface.

17. C. modesta new name. Plate 27, figs. 44—46.

Krynitzkia depressa Jones, Contr. West. Bot. 13:5. 1910, not
Cryptantha depressa Nelson, Bot. Gaz. 34: 29. 1902.

Oreocarya depressa (Jones) Macbr. Contr. Gray Herb. 48: 32.
1916.

Densely caespitose, long-lived perennial from a woody root;
stems few to many from the branching caudex which is densely
clothed with the leaf-bases of former years, 8-15 cm. high, rather
slender, setose with slender, weak bristles; leaves numerous on
the crowns of the caudices, spatulate, obtuse, 1.5-4 ст. long,
closely tomentose and setose with weak appressed bristles, pus-
tulate on both surfaces, more abundantly on dorsal, petioles about
as long as the blades, conspicuously ciliate near the base; inflores-
сепсе extending over 1% to 34 of the stem, narrow, not inter-
rupted, cymules not elongating, rather softly setose-hirsute,
foliar bracts inconspicuous; sepals setose-hirsute, linear-lanceolate,
acute, in anthesis 3—4 mm. long, in fruit 7-8 mm. long, exceeding
the nutlets by 2-3 mm.; corolla white, tube 3-4 mm. long, equal-
ling the sepals, crests at the base conspicuous, fornices probably
yellow, rounded, nearly closing the throat, minutely papillose,
0.5-1 mm. long, limb 7-8 mm. broad, lobes equalling or exceeding
the tube, united for about 14 their lengths; fruit ovoid, 2-3 nut-
lets usually maturing; nutlets broadly ovate, obtuse or acute,
with slight tendency to be carinate, 3-4 mm. long, margins in
contact, acute, surfaces of nutlets glossy, the dorsal tuberculate
and often more or less rugose, tubercles and rugae (if present)

[Vor. 14
278 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

rounded and rather low, ventral surface deeply rugose and tuber-
culate, scar triangular, open, margin somewhat inclined to be
elevated.

Distribution: Upper Sonoran Zone in southwestern Utah and
eastern Nevada. Туре: Aurum, Nevada, alt. 7,300 ft., June 20,
1893, M. E. Jones 6692.

Specimens examined:

Utah: hills east of Grass Valley, May 17, 1875, Ward 49 (U.S.,
Gray); Modena, June 2, 1902, Goodding 996 (R.Mt., Pomona,
Mo., U.S., Gray).

Nevada: Aurum, June 20, 1893, Jones 6692 (Pomona, TYPE,
Mo., U.S.); Muncy, July 6, 1891, Jones 6713 (Pomona).

This species is perhaps most closely related to tumulosa, vir-
ginensis, and insolita. In general appearance it is not unlike
tumulosa or some of the varieties of C. nana.

18. C. humilis (Greene) new comb. Plate 27, figs. 47-49.

Eritrichium glomeratum var. (?) fulvocanescens Wats. Bot.
King’s Exp. 243. 1871.

Е. glomeratum var. humile Gray, Proc. Am. Acad. 10:61. 1875,
in part.

Oreocarya humilis Greene, Pittonia 3: 112. 1896.

O. hispida Nelson & Kennedy, Proc. Biol. Soc. Wash. 19: 156.
1906. (Type: “Collected in Carson Valley, Ormsby County,
Nevada, April 24, 1904," No. 865, G. H. True.)

O. echinoides Macbride, Contr. Gray Herb. 48: 31. 1916, not
Krynitzkia echinoides Jones.

О. Macbridii Brand, Fedde, Rep. Sp. Хоу. 19: 73. 1923.
(Type: Mt. Jarbidge, Nevada, July 6, 1912, Nelson & Macbride
1960.)

Caespitose perennial; stems 1-зеуега] from the woody caudex,
rather stout or slender, 10-30 cm. high, setose with rather weak
bristles; leaves obovate-spatulate to oblanceolate, 3-8 em. long,
usually obtuse, rather weakly setose with appressed or slightly
spreading bristles, the older leaves distinctly tomentose, petioles
conspicuously ciliate near the base with long, white, weak hairs,
pustules about equally numerous on both leaf-surfaces; inflores-
cence rather narrow, mostly continuous, extending over upper %

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 279

or 24 of the stem, foliar bracts not conspicuous; calyx densely
setose with rather slender bristles, sepals in anthesis linear or
linear-lanceolate, acute, 4-5 mm. long, in fruit 8-13 mm. long,
exceeding the nutlets by 5-8 mm.; corolla white, tube 4-5 mm.
long, tube and calyx lobes subequal, crests at the base of the tube
small but evident, fornices low (0.5 mm.), rounded, рарШове,
probably pale yellow, limb 8-10 mm. broad, lobes and tube
subequal, lobes united for about 14 their length; fruit lance-ovoid,
all four nutlets commonly maturing, style exceeding the mature
nutlets 1.5-2.5 mm.; nutlets ovate-lanceolate to lanceolate,
acute or obtuse, 3-4.5 mm. long, margins in contact, acute or
obtuse, surfaces of nutlets somewhat glossy, the dorsal often
densely muricate, more or less tuberculate and usually rugose,
ventral surface rather indistinctly muricate or tuberculate, scar
open at the base or nearly closed, no elevated margin.

Distribution: mostly in mountainous parts of southwestern
Idaho, Nevada, and northeastern California. Туре not desig-
nated: “Frequent in the mountains of Nevada and adjacent
eastern California: the Californian plant, as collected by Mr.
Sonne, having nutlets nearly twice as large as in the more typical
form of eastern Nevada, yet otherwise quite the same."

Specimens examined:

Idaho: Hot Hole, e. fork of Bruneau, July 3, 1912, Nelson &
Macbride (R.Mt.).

Nevada: near Holborn, July 16, 1896, Greene (R.Mt., frag-
ment, TYPE in Notre Dame Herb.); Palisade, June 12, 1903,
Stokes (Calif.) ; Palisade, June 14, 1882, Jones (Pomona) ; Jarbidge,
July 6, 1912, Nelson & Macbride 1960 (R.Mt., Minn., Mo., U.S.,
Gray); Jarbidge, July 8, 1912, Nelson & Macbride 1980 (R.Mt.,
Minn., Mo., U.S., Gray); Wadsworth, June 16, 1897, Jones (Ро-
mona); Monitor Valley, July, 1868, Watson 863, in part (U.S.,
Gray); Coleman Valley, n.w. Nevada, July 29, 1896, Coville & Lei-
berg 91 (U.S.); Carson City, May, 1882, Jones 6732 (Pomona);
Carson Valley, April 24, 1904, True 865 (R.Mt.); Hunter Creek,
Washoe Co., Мау 17, 1907, Kennedy 1664 (Calif., Mo., U.S.).

California: sandy banks of Truckee River at Verdi, May and
June, 1887, Sonne (Calif., Mo.); on Truckee River, Placer Co.,
July, 1886, Sonne (Calif.); Squaw Valley on Truckee River, June-

[Vor. 14
280 ANNALS OF THE MISSOURI BOTANICAL GARDEN

July, 1885-6, Sonne (Calif.); Mt. Stanford, Nevada Co., July,
1892, Sonne (Calif, Mo.); Susanville, June 28, 1897, Jones
(Pomona); Castle Peak near the highest point, Nevada Co.,
Aug. 3, 1903, Heller (Phila., Mo., U.S., Gray); Sierra Co., 1874,
Lemmon 165 (Mo.); Castle Peak, 9,000 ft., Aug. 7, 1900, Leiberg
5994 (U.8.); summit of Sierra Nevada, July 10, 1879, Kellogg
(Calif.).

Thanks to Dr. J. A. Nieuwland, of Notre Dame University,
I am in receipt of fragments from specimens labeled in Dr.
Greene's handwriting as Oreocarya humilis Greene. Мо one of
these is designated as the type but doubtless they serve to confirm
our notion of Greene's concept of this species. Greene used a
name proposed by Gray as a variety of glomerata and later trans-
ferred by him as the major part of sericea. С. humilis, as we
now know it, was a small part of Gray's concept of the var.
humilis of glomerata. However, it seems only reasonable to
adopt the name for the species in the sense in which Greene used
it. The specimens designated as humilis by Greene are as follows:

1. Near Holborn, Nevada, July 16, 1896, Edw. L. Greene.

2. Banks of ТгисКее River, near Verdi, California, April and
May, 1889, C. F. Sonne.

3. Mt. near Truckee River, 8000 ft., July 18, 1886, C. F.
Sonne 77 (?).

It is always desirable to select a type specimen for every species
whenever possible. In the present case the choice definitely falls
upon the specimen collected by Dr. Greene near Holborn,
Nevada. There are three reasons for this choice. First, Greene
describes the nutlets as “rather densely tuberculate but not
rugose." This is more characteristic of the specimen from
eastern Nevada than of those from California. Second, Greene
refers to the form from eastern Nevada as ‘‘the more typical
form." ‘Third, an author might be expected to select a plant
of his own collecting as the center around which his concept is
formed rather than one collected by another collector.

In eastern Nevada the boundary between this species and
nana var. commixta seems rather weak. Тһе rugae tend to be
reduced to scarcely continuous tubercles or indistinct wrinkles.
It was one of these forms that Macbride classified as nana var.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 281

commixta. Brand, examining it more critically, noticed the rugae
and thinking that it could not be placed in commixta, proposed а
new species for it—O. Macbridii. An examination of а nutlet
from Greene’s plant from Holborn, Nevada, makes it evident
that the elevations are two sizes. The smaller ones we call
murications and the larger, tubercles. In neither are the rugae
entirely absent. The specific limits between humilis and nana
var. commizta are rather uncertain in the eastern Nevada region.
It is unfortunate that this intermediate form should have been
selected as the type rather than the more different forms from
the Sierra Nevada. Oreocarya Macbridii is evidently an exact
synonym of C. humilis.

19. С. caespitosa (A. Nels.) new comb. Plate 27, figs. 50-52.

Oreocarya caespitosa A. Nels. Erythea 7: 65. 1899.

Densely caespitose, long-lived perennial; stems many from the
ends of the numerous caudices, rather slender, 0.5-1 dm. high,
weakly setose or hirsute; leaves mainly clustered on the caudices,
spatulate to oblanceolate, obtuse or acute, 2-5 cm. long, densely
strigose and appressed-setose with little-differentiated hairs,
these so densely matted as to give an appearance similar to
tomentum, pustules abundant on the lower surface, somewhat
fewer above but quite evident in the older leaves; inflorescence
narrow, confined to upper 44-34 of the stem, foliar bracts incon-
spicuous; calyx subtomentose and shortly setose or hirsute,
sepals linear to linear-lanceolate, acute, 3-4 mm. long in an-
thesis, in fruit 6-7 mm. long, exceeding the nutlets by about 3
mm.; corolla white, tube З mm. long, equalling or slightly shorter
than the sepals, crests well developed at the base of the tube,
fornices low (0.5 mm.), rounded, pale yellow, limb 5-6 mm.
broad, tube and limb subequal or the tube slightly longer, corolla
lobes united for 14-15 their length; fruit lance-ovoid, less than
four nutlets usually maturing (usually 2?), style scarcely, if at all,
exceeding the nutlets; nutlets lanceolate, acute, 3-3.5 mm. long,
margins in contact, obtuse or acute, surfaces of nutlets white, dull,
the dorsal definitely rugose with low, rounded rugae, somewhat
tuberculate, these larger elevations surrounded by numerous low,
rounded muriculations, the ventral surfaces minutely and rather

[Vor. 14
282 ANNALS OF THE MISSOURI BOTANICAL GARDEN

indefinitely muriculate, scar extending from the base nearly to
the apex, open and narrowly triangular, no elevated margin
present.

Distribution: Upper Sonoran and lower part of the Transition
Zones in southern Wyoming. Туре (as designated in R.Mt.
Herb.): Point of Rocks, A. Nelson 4749.

Specimens examined:

Wyoming: Rocky Mts., east side, Nuttall (Phila.); Cooper
Creek, June 5, 1898, E. Nelson 5448 (R.Mt.); Freezeout Hills,
July 10, 1898, E. Nelson 4497 (R.Mt.); T. B. Ranch, Carbon Co.,
June 20, 1901, Goodding 60 (R.Mt.); Ft. Steele, May 25-June 10,
1901, T'weedy 4260 (U.S.); Ft. Steele, June 16, 1900, Nelson 7255
(R.Mt., Gray); Point of Rocks, June 15, 1898, Nelson 4749
(R.Mt., түре, U.S.); Point of Rocks, June 19, 1901, Merrill &
Wilcox 617 (R.Mt., U.S., Gray); Red Desert, June 3, 1897, Nelson
3120 (R.Mt.); Bush Ranch, Sweetwater Co., June 10, 1900,
Nelson 7078 (R.Mt., Pomona, Minn., Мо., U.S., Gray); n.e.
corner of Sweetwater Co., July 6, 1926, Nelson 10696 (R.Mt.);
Bitter Creek, June 16, 1898, Nelson 4772 (R.Mt., Mo., Gray);
Green River, June 1, 1897, Nelson 3072 (R.Mt.); Green River,
June 23, 1896, Jones 6726 (Pomona); red marl hills оп Gros
Ventre Fork, June 10, 1860, Hayden (Мо.); between Eden & Big
Piney, July 6, 1922, Payson & Payson 2578 (R.Mt., Pomona,
Mo., Gray); between Opal and Kemmerer, June 19, 1923, Payson
& Armstrong 3223 (R.Mt., Mo., Gray); Fossil, June 12, 1898,
Nelson 4671 (R.Mt.); east of Evanston, June 28, 1922, Osterhout
6248 (R.Mt.).

When Dr. Nelson first described this species he included in his
specific concept specimens of the plant he later described as
Oreocarya cana. The two are in reality very different in char-
acters of pubescence and range. С. caespitosa is perhaps most
closely related to C. modesta although it is perhaps not very far
removed from C. humilis. Whatever its exact relationship
caespitosa is a very definite unit with an isolated range and it is
doubtful if any treatment, however large the specific concept,
would fail to accord it full rank as a species.

Series 6. BnADBURIANAE. Nutlets always roughened dorsally
and distinctly rugose or tuberculate, or both, and often muricate

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 283

also. Ventral surfaces smooth or variously roughened. Scar
straight, closed or nearly so, margins not elevated. Corolla tubes
never longer than the calyx lobes. Species 20-31.

20. C. thyrsiflora (Greene) new comb. Plate 27, figs. 53-55.

Eritrichium glomeratum var. hispidissimum Torr. Bot. Mex.
Bound. Surv. 140. 1859, at least in part.

Oreocarya thyrsiflora Greene, Pittonia 3: 111. 1896.

O. hispidissima (Torr.) Rydb. Bull. Torr. Bot. Club 33: 150.
1906; Fl. Colo. 288. 1906.

O. urticacea Wooton & Standley, Contr. U. S. Nat. Herb. 16:
166. 1913. (Type: Canyoncito, Santa Fe Co., New Mexico,
June 18, 1897, A. A. & E. G. Heller 3731).

O. dura Nels. & Macbr. Bot. Gaz. 62: 144. 1916. (Type: E.
L. Johnston 418, 1907, Central Colorado).

O. monosperma Osterhout, Bull. Torr. Bot. Club 46: 55. 1919.
(Type: Trinidad, Las Animas Co., Colorado, July 20, 1918,
Osterhout 5754).

Short-lived perennial (probably sometimes biennial); stems
l-several from the base, rather stout, 2-4 dm. high, densely
setose; leaves oblanceolate, obtuse, 4-10 cm. long, densely and
rather coarsely strigose, conspicuously pustulate on both leaf-
surfaces, sometimes less densely so above, petioles conspicuously
and coarsely setose-ciliate; inflorescence broad and round-topped,
dense, abundantly setose, cymes much elongating, foliar bracts
large but not conspicuous because of the width of the inflorescence;
calyx densely setose, sepals in anthesis linear to lanceolate-linear,
3—4 mm. long, acute, in fruit 6-8 mm. long, exceeding the nutlets
by about 4 mm.; corolla white, tube 3-4 mm. long, about as long
as the sepals, crests at the base of the tube evident, usually well
developed, fornices 0.5-1 mm. long, papillose, pale yellow, limb
7-8 mm. across, lobes and tube subequal, lobes united for about
lg their length, style exceeding the mature nutlets about 1.5
mm. ; nutlets ovate-lanceolate, acute, about 3.5 mm. long, margins
in contact, acute or obtuse, surfaces of nutlets somewhat glossy,
the dorsal distinctly rugose, usually with a few tubercles, indefi-
nitely and sparsely muriculate, or slightly roughened, scar
straight, narrow,margin not elevated.

[Vor. 14
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: Transition and Upper Sonoran Zones in western
Nebraska, southeastern Wyoming, eastern slope of the moun-
tains in Colorado and northern New Mexico. Туре: “Оп the
stony hills in southern Wyoming about Cheyenne, Laramie, etc.,
thence to middle Colorado,” Greene.

Specimens examined:

Nebraska: War Bonnet Canyon, June, 1890, Williams (Mo.);
Rush Creek, Deuel Co., July 2, 1891, Rydberg 255 (Minn., U.S.).

Wyoming: Rocky Mts., Platte, Nuttall (Phila.); Pine Bluffs,
June 27, 1889, Bodin (Minn.); Lower Pole Creek, July, 1856, H.
Engelmann (Мо.) ; Guernsey, June 26, 1901, Nelson 8263 (R.Mt.) ;
Cheyenne, June 30, 1896, Nelson 1990 (R.Mt., Minn.); 15 miles
west of Cheyenne, Aug. 24, 1925, Payson & Payson 4685 (В. Mt.) ;
Chug Creek, Albany Co., June 29, 1900, Nelson 7306 (R.Mt.,
Pomona, Minn., Mo., U.S., Gray); Table Mt., June 30, 1895,
Nelson 1362 (R.Mt.); Laramie, July 8, 1894, Nelson 418 (R.Mt.,
U.S., Gray); Sheep Mt., Albany Co., Sept. 2, 1903, Goodding 2076
(R.Mt., U.S.); Sheep Mt., June 24, 1924, Payson & Payson 4246
(R.Mt.).

Colorado: Pawnee Buttes, July 8, 1919, Osterhout 5934 (Gray) ;
Pawnee Buttes, July 1, 1906, Dodds 2077 (R.Mt.); Livermore,
Larimer Co., June 3, 1897, Osterhout (R.Mt.); Livermore, July
26, 1913, Osterhout 4979 (R.Mt.); near Florissant, Aug. 1-8,
1905, Ramaley 1448 (R.Mt.); Bear Canyon, Colorado Springs,
June 25, 1879, Jones 972 (R.Mt., Pomona, Calif., U.S.) ; Colorado
Springs, June 15, 1896, Knowlton 35 (U.S.); Pikes Peak, July,
1920, Johnston 2817 (Gray); Twin Lakes, July, 1873, Wolf &
Rothrock 700 (Phila., U.S.); Granite to Twin Lakes, Lake Co.,
Aug. 3, 1925, Smith 3916 (R.Mt.); Granite, July 23, 1889, Ever-
mann (U.S.) ; Canyon City, Sept. 26, 1874, G. Engelmann (Mo.);
Canyon City, 1877, Brandegee 4 (Mo.); Canyon City, June 27,
1896, Osterhout 629 (R.Mt., Osterh.); near Salida, June 27, 1917,
Payson 1014 (R.Mt.); Salida, June 19, 1898, Baker, Earle &
Tracy 13 (R.Mt., Pomona, Minn., Mo., U.S., Gray); South Park,
Aug. 3, 1884, Letterman (Field); Sangre de Cristo Creek, July 2,
1900, Rydberg & Vreeland 5704 (R.Mt.); Buena Vista, July 6,
1892, Sheldon 526 (U.S.); Buena Vista, Aug. 1-2, 1919, Eggleston
15879 (Field); Trinidad, June 28, 1898, Osterhout (В. Mt.);

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 285

Trinidad, July 20, 1918, Osterhout 5754 (R.Mt., Osterh., Gray);
near Trinidad, June 17, 1911, Standley 6036 (U.S.).

New Mexico: Canoncito, Santa Fe Co., June 18, 1897, A. A. &
E. С. Heller 3731 (Minn., Mo., U.S.); Glorieta, June, 1881,
Vasey (R.Mt., Mo., U.S.); Sierra Grande, Union Co., June 18,
1911, Standley 6056 (U.S.).

Early collections of this species were very generally referred
to C. Bradburiana. This was probably due in large part to the
fact that the two are rarely found growing together. In south-
eastern Wyoming where the ranges do overlap and they are seen
growing together it is at once evident that they are distinct
specifically. The broad inflorescence of thyrsiflora is the out-
standing distinction of the species, but in addition the flowers of
thyrsiflora are smaller and the blooming season later than іп
Bradburiana. Тһе author was at one time inclined to refer
certain Montana plants with a broad open inflorescence to this
species. Later, however, it has seemed evident that they are
forms of C. Bradburiana or of C. Sheldonit.

А careful examination of the type of Oreocarya dura makes it
quite certain that it is an immature specimen of C. thyrsiflora.

21. C. elata (Eastw.) new comb. Plate 28, figs. 56—58.

Oreocarya elata Eastw. Bull. Torr. Bot. Club 30: 241. 1903.

Perennial, probably short-lived; stem rather stout, one to
several, 3-5 dm. high, rather sparsely and loosely strigose as well
as hirsute-setose; radical leaves clustered at the base, spatulate,
1.5-3 em. long, blade often rather abruptly narrowed to the
petiole, acute or obtuse, rather coarsely strigose, densely pustulate
and appressed-setose on dorsal surface, ventral surface strigose
and pustulate, petioles densely covered with long white hairs;
cauline leaves linear-oblanceolate, acute, 2-4 cm. long, setae
more widely spreading; inflorescence extending over upper l4 or
15 the length of the stem, upper cymes elongating, the lower ге-
maining short, the inflorescence then with a tendency to have a
broad top, much narrower downwards, abundantly setose with
rather weak slender bristles, bracts inconspicuous; calyx abun-
dantly setose with rather weak bristles, sepals in anthesis lance-
olate, acute, about 4 mm. long, in fruit about 8 mm. long, ex-

[Vor. 14
286 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

ceeding the nutlets by 2-3 mm.; corolla white, tube about 4 mm.
long, equalling the sepals, сгезфз at the base of the tube well
developed, fornices nearly 1 mm. high, distinctly рарШове,
probably yellow, limb about 9 mm. broad, lobes and tube sub-
equal, lobes united for about 15 their length; fruit lance-ovoid,
all four nutlets usually maturing, style exceeding the nutlets 1-2
mm. ; nutlets ovate-lanceolate, acute or obtuse, about 5 mm. long,
margins acute, in contact, surfaces of nutlets slightly glossy,
the dorsal margined, densely tuberculate and more or less ru-
gose, entire surface minutely and densely papillose, ventral sur-
face similar but less evidently rugose, scar straight, closed, ex-
tending from near the base almost to the apex, slightly open and
forked at the base, no elevated margin present.

Distribution: Upper Sonoran Zone, near Grand Junction,
Colorado. Туре: “Collected by the author [Eastwood] near
Grand Junction, Colorado, on the road to the coal mines, growing
on the bare clay hills characteristic of the region, flowering
May 15, fruiting June 28, 1892.

ans examined:

Colorado: Grand Junction, Мау 15, 1892, Eastwood (Х.Ү.,
U.S., Gray); Grand Junction, June 10, 1920, Osterhout 5996
(R.Mt., Gray, Osterh.).

Although still imperfectly known, there can be no doubt but
that this species is very distinct and quite worthy of specific
rank. Іп habital characteristics it resembles C. insolita very
closely. "The nutlet characters of the two species are so different,
however, that one doubts any very close relationship between the
two. Excellent but immature specimens of the type collection
are in the National Herbarium and in the Gray Herbarium.

22. C. sericea (Gray) new comb.

Perennial from a tap root; stems 1-many from the base, 1.5-
4.5 dm. high, rather sparsely setose with slender, divaricate
bristles; leaves somewhat clustered at the base, spatulate to
oblanceolate, usually obtuse, 2.5-10 cm. long, dorsal surface
densely strigose and appressed-setose, abundantly pustulate,
ventral surface densely and uniformly silky-strigose, rarely, if at
all, pustulate; inflorescence extending over 24 to 34 the length

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 287

of the stem, in flower narrow, somewhat glomerate, abundantly
setose, foliar braets conspicuous, especially in the young in-
florescence; calyx setose, sepals in anthesis linear-lanceolate,
acute, 3—4 mm. long, in fruit 7-9 mm. long, exceeding the nutlets
by 3-5 mm.; corolla white, tube 2.5-3.5 mm. long, slightly shorter
than the sepals, crests at the base of the tube evident or con-
spicuous, fornices small (about 0.5 mm. high), rounded, papillose,
probably always pale yellow, limb 6-8 mm. broad, lobes and tube
subequal, lobes united for about 14 their length; fruit ovoid or
lance-ovoid, 3-4 nutlets commonly maturing, style exceeding
the mature nutlets by 0.25-1 mm.; nutlets lanceolate, acute,
2.75-3.5 mm. long, margins in contact, acute, surfaces of nutlets
dull, the dorsal sparsely tuberculate and somewhat rugose as well
as more or less muriculate, ventral surfaces at least somewhat
roughened, usually distinctly muriculate, scar straight, closed or
nearly so, extending from near the base to near the apex, no ele-
vated margin present.

Distribution: Upper Sonoran Zone, southern and southwestern
Wyoming, northern Colorado, northeastern Utah.

The varieties of C. sericea may be distinguished as follows:

Inflorescence becoming rather broad in age due to the elongation the cy-

mules; leaves oblanceolate; nutlets 3-3.5 mm. long.............. 7a. var. typica
Inflorescence usually narrow, even in age; leaves narrowly COR xcd nut-

Msz 10-0 mm. long............22 л КОСЕ eas saat 37b. var. perennis

22a. Var. typica n. var. Plate 28, figs. 59-61.

Eritrichium glomeratum var. humile Gray, Proc. Am. Acad. 10:
61. 1875, in large part.

Krynitzkia sericea Gray, Proc. Ат. Acad. 20: 279. 1885, in
part.

Oreocarya sericea Greene, Pittonia 1: 58. 1887.

O. argentea Rydb. Bull. Torr. Bot. Club 31:637. 1904. (Type:
Rifle, Garfield Co., Colo., 1900, Osterhout 2122).

O. procera Osterhout, Bull. Torr. Bot. Club 47:211. 1920.
(Type: Glenwood Springs, Garfield Co., Colorado, June 18, 1899,
Osterhout 1867).

Stems usually stout, 1.5-4 dm. high; leaves rather broadly ob-
lanceolate, obtuse, 4-10 cm. long; inflorescence becoming rather
broad in fruit due to the partial elongation of the cymules; sepals

[Vor. 14
288 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

about 7 mm. long in fruit, exceeding the nutlets by 3-4 mm.;
corolla tube about 3 mm. long, limb 6-8 mm. broad; fruit ovoid;
nutlets rather broadly lanceolate, 3-3.5 mm. long, rather incon-
spicuously margined, style scarcely exceeding the mature nutlets,
dorsal surface of the nutlets sparsely tuberculate, distinctly
rugose and more or less muriculate, ventral surface sparsely
muriculate, scar closed or nearly closed, straight.

Distribution: southern Wyoming, northwestern Colorado, and
northeastern Utah. Type: Bridger’s Pass (Wyoming), 1856,
Henry Engelmann.

Specimens examined:

Wyoming: Elk Mt., July 1, 1899, Little & Stanton 179 (Mo.);
Bridger Pass, 1856, H. Engelmann (Gray, түре, Mo.); Pass
Creek to Bridger’s Pass, 1859, H. Engelmann (N. Y., Gray); 11
miles north of Encampment, July 2, 1922, Payson & Payson
2528 (R.Mt., Pomona, Field, Mo., Gray).

Colorado: Hayden, Routt Co., July 10, 1913, Osterhout 4940
(R.Mt., Osterh.) ; Kremmling, Grand Co., June 22, 1907, Osterhout
3476 (R.Mt., Osterh.) ; Rifle, June 23, 1900, Osterhout 2122 (Pomo-
na, N. Y.); Middle Park, July 26, 1876, Patterson (Field) ; Glen-
wood Springs, June 8, 1920, Osterhout 5960 (R.Mt., U.S., Gray,
Osterh.); Glenwood Springs, June 29, 1895, Osterhout (R.Mt.);
Grand Junction, May 30, 1921, Osterhout 6094 (Osterh.).

Utah: Mill Fork, July 7, 1894, Jones 5607a (Pomona, Calif.,

.5.).

22b. Var. perennis (А. Nels.) new comb.

Oreocarya affinis perennis A. Nels. Erythea 7: 67. 1899.

O. perennis Rydb. Bull. Torr. Bot. Club 33: 150. 1906.

Stems rather slender, 1.5-4 dm. high; leaves narrowly oblance-
olate, acute or obtuse, 2.5-4.5 em. long; inflorescence narrow and
glomerate even in fruit, cymules scarcely elongating; sepals about
7 mm. long in fruit, exceeding the nutlets by 3.4 mm.; corolla
white with pale yellow fornices, tube 2.5 mm. long, limb about 6
mm. broad; fruit lance-ovoid; nutlets narrowly lanceolate, not
wing-margined, 2.75-3 mm. long, dorsal surface sparsely tuber-
culate, more or less rugose, somewhat muriculate, ventral surface
rather sparsely muriculate, scar straight, closed.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 289

Distribution: Upper Sonoran Zone, southwestern Wyoming.
Type: Green River, Wyoming, May 31, 1897, A. Nelson 3035.

Specimens examined:

Wyoming: between Eden and Big Piney, July 6, 1922, Payson
& Payson 2580 (R.Mt., Pomona, Field, Mo., Gray, Colo.);
Green River, June 25, 1895, Shear 4381 (U.S.); Green River,
June 23, 1896, Jones (Pomona); Green River, June 14, 1898,
Nelson 4715 (R.Mt., түре, Mo., Gray); Green River, Мау
30, 1897, Nelson 3035 (R.Mt., U.S.); 21 miles west of Green
River, June 19, 1923, Payson & Armstrong 3206 (R.Mt., Mo.,
Gray); between Opal.and Kemmerer, June 19, 1923, Payson &
Armstrong 3224 (R.Mt., Mo.); Ft. Bridger, July, 1873, Porter
(Herbarium not recorded); Evanston, July, 1869, Watson 852
(0.8.).

The name sericea has always been a stumbling block in the
way of any satisfactory treatment of this group of plants. At
the time when it was described (1885) only seven other species
had been proposed in this section. These were among the
most outstanding units in the genus and it is evident that sericea,
whatever it was, was different from any of these and so must be
retained as a valid species. Because of this necessity all botanists,
to treat these plants for the region assigned to sericea, had to
make some disposition of it. These interpretations have been
extremely varied, and herbarium material labeled “sericea” has
added to the confusion. During the course of the present study
it became probable that sericea, when it was located, would almost
certainly replace some more recently proposed name, since most
of the distinguishable forms were treated under recent names and
there was left no place for sericea—which was a name older than
these others.

The problem was evidently to be solved by a careful study of
the specimens available to Dr. Gray when he proposed the species
and a consideration of the evidence gathered from the published
description. The author believes he has reached a satisfactory
solution of the problem by following out these lines of research
and proposes to restrict the name sericea to the plant that has
recently been treated as Oreocarya argentea Rydb. Тһе evidence
on which this conclusion is based is presented in detail in the
following paragraphs.

ГУ ол. 14
290 ANNALS OF THE MISSOURI BOTANICAL GARDEN

First of all it is useful to summarize the criteria that have
governed the case in question.

1. It is necessary to select a type specimen on which to attach
the name in the final analysis, since it is evident that more than
one species was involved in Dr. Gray's concept. It must be
borne in mind that it is rather unfair to force a type specimen
concept on work that was done before this method was em-
phasized as it is at present. In case it can not be shown that
Dr. Gray had one plant especially in mind when he described
sericea then it might be better to discard the name altogether.

2. The type specimen, if one is to be located, must be selected
from the material Dr. Gray had at hand when he described
sericea. Since he based this species in large part, “тарпа parte,"
on his earlier variety humile, it would seem only reasonable to
select the type from specimens which he had previously considered
as representing that variety.

3. Тһе type should be represented by а mature and adequate
specimen in which the nutlets are developed since these were
described and are of the greatest importance in determining the
species of this genus.

4. Тһе description of sericea as given by Gray should be exam-
ined very carefully and the characters emphasized there should
belong to the type specimen. These may be summarized as
follows: sericea is said to differ from glomerata in being less hispid
and in being perennial rather than annual. It isa many-stemmed
plant. The indument is soft, strigose-sericeous, and the setae are
short, not very rigid and often appressed. The nutlets are
subrugose-tuberculate and oblong-ovate.

5. The range of the species sericea is given as follows: “ Alpine
or subalpine, on mountains from Colorado and Utah to Oregon
and Montana and probably in the British Possessions. There
are less canescent forms from the Saskatchewan region and also
from the higher Sierra Nevada (and uncertain as to the duration
of the root) which may belong either to this species or to dwarfed
forms of the foregoing.” It would seem that Dr. Gray thought
of the forms from Saskatchewan, the British Possessions, or from
the higher Sierra Nevada as aberrant. The type then should
come from some point in the range ‘‘alpine or subalpine, on

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 291

mountains from Colorado and Utah to Oregon and Montana."

6. Dr. Gray says, “І have adopted one of the two specific
names under which this species occurs in Nuttall's collections."
Nuttall's specimen should be examined and its identity consid-
ered in connection with the specimens and description of sericea
as given by Gray.

Тһе next point of attack 13 to examined the herbarium material
considered by Dr. Gray as representing his new species. There
are in the Gray Herbarium three sheets that bear the notation
“ Krynitzkia sericea Gray" in Dr. Gray's own handwriting.
These sheets also bear the designation “Ed. 2. Syn. Fl. N. Amer."
Each of these sheets bears several specimens. This material may
be described as follows:

Sheet 1 contains 4 specimens:

a. Bridger's Pass, 1856, H. Engelmann = 0. argentea Rydb.

b. W. side of Wasatch Mts., 1844, Frémont = О. humilis Greené, К. de-
pressa Jones, or some form
of O. nana Eastwood. Too
immature for certaih iden-

tion.
c. Clover Mts., Nevada, 1868, Watson 853 = О. humilis Greene, k. de-

immature for certain iden-
tification.
d. Mountain Hot Springs, Yellowstone Park, 1885,
Tweedy 816 = C. Bradburiana.

Sheet 2 contains 3 specimens:
а. Montana Terr. 1867 = C. Bradburiana.
It also bears the two separate notations in
Dr. Gray's handwriting “E. glomeratum” and
* Probably dwarf K. glomerata."

b. Summit, California, 1871, Bolander = humilis or possibly nubi-
gena. Specimens immature.
с. Grass Valley, Utah, 1875, Ward 49 — depressa

Sheet 3 contains 2 specimens at the present time and probably 5 (including frag-
ments) in Dr. Gray's time:
a. Shinberger's Canyon, S. Montana, 1880, Watson 287 = Sheldonii. | Specimen im-
mature but dentieation
practically certai
b. A specimen of Bradburiana without data. A note by Dr. Gray says “origin un-
certain."

[Vor. 14
292 ANNALS OF THE MISSOURI BOTANICAL GARDEN

c. Pine Creek, Baker Co., Oregon, 1879, Cusick. This is not now on this sheet but
the piece on which it is mounted fits the remains of apes 3, and evidently
it was attached to sheet 3 in Dr. Gray’s time. This is nubigena.

d. Another fragment has been located that was probably attached to sheet no. 3.
This fragment contains 2 fragmentary specimens and a in Dr. Gray’s
handwriting: “Two more dwarf and very sine forms in Herb.
Acad. Philad. Nuttall.” One of the fragmentary specimens is evidently a bit of
the plant we now know as caespitosa. It is from a specimen collected by
Nuttall.

e. The second specimen on the scrap described above is an immature piece of the
plant we now know as cana and was collected at Scott’s Bluff on May 30,
1858.

If we now list the specimens that may be considered to com-
pete as the type of sericea we see that the following species are
represented: argentea, humilis, depressa, nana, Bradburiana,
nubigena, Sheldonii, caespitosa, and cana. Five of these can be
discarded at once. Bradburiana must not be considered, since
Dr. Gray was separating his species from that and gave good
differences between the two, while nana must be discarded because
no specimen is represented that is mature enough to make iden-
tification certain. The species humilis can be eliminated because
it is represented by no mature specimen and because the speci-
mens from the higher Sierra Nevada were considered aberrant.
Sheldonii is represented by a single specimen in which the nutlets
are quite immature, and caespitosa is represented by fragments
so scanty that they are only recognizable by association with a
better specimen in the herbarium of the Philadelphia Academy.
It carries a manuscript name given it by Nuttall which was never
published and which is not sericea. We now have left for con-
sideration the species argentea, depressa, nubigena, and cana.

Earlier in this discussion it was suggested that since Gray’s
sericea was based in large part upon his variety humile it would
be reasonable to select a type that had been included in his variety
when it was published in 1875. If we now examine the specimens
of the remaining species we find that the mature specimen of
nubigena, and the only certain one of that species, was collected
in 1879, after the variety was published. We also find that the
only certain representative of depressa was collected in 1875, the
year of the publication of the variety, and so was evidently not
at hand when the variety was described since the papers by Dr.

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 203

Gray containing this description were presented to the Academy
“Мау 12 and Oct. 13, 1874." We find Dr. Gray’s notation
“Е. glomeratum, humile” on only two specimens of the series
that has been described. "These are on the one from Bridger's
Pass (argentea) and on the immature one from the “У. side of
the Wasatch Mts." This serves to confirm the selection of
argentea as the center around which Dr. Gray built his variety
humile and species sericea. We have left, however, two species
that need further consideration—argentea and cana—since
nubigena and depressa are eliminated because they did not form
a part of Gray’s variety humile.

The only reason for considering cana as a competitor for the
name of the species in question is because Dr. Gray said he chose
a name under which the species occurs in Nuttall’s collections.
The plant to which Nuttall gave the manuscript name of sericea
was undoubtedly the plant we now know as cana. Dr. Gray
believed this plant to be part of his species sericea, but it is evident
he did not describe it or have it particularly in mind. He said
sericea was the same as his var. humile in large part. He did not
consider Nuttall’s sericea as typical var. humile, as is shown by his
comment, “Nuttall collected and gave MSS. names to some
very dwarf and silky canescent forms, which appear to belong
here." These aberrant forms, we now know as caespitosa and
cana. The description of Gray’s sericea does not fit cana. The
pubescence in cana is practically uniform and the setae would
not have been described “short, not very rigid and often ap-
pressed." Тһе nutlets were unknown to Dr. Gray, and if they
had been known would not have been described as ''subrugose-
tubereulate." Тһе range given by Gray for sericea is scarcely
large enough to include cana. Scott's Bluffs, from where he had
а specimen of cana, is in Nebraska and so hardly to be described

“alpine and subalpine.”

By a process of elimination we obtain a type specimen for
sericea and select a specimen from Bridger’s Pass, collected by
H. Engelmann, to represent the species in the final analysis.
This selection, however, need not be made on a basis of elimina-
tion, since the specimen under consideration has many points
that argue for its being the plant that Dr. Gray had especially in

[Vor. 14
294 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

mind when the species was described. Since the matter is of so
much importance it may be well to recapitulate briefly the reasons
why the specimen from Bridger's Pass is held as the type of
sericea.

1. Of the 8 species that are probably represented on the sheets
that Gray had at hand when he described sericea there is no
numerical preponderance of any one species. Тһе species
argentea is represented by one specimen and another sheet was
undoubtedly present and considered as sericea, but in this case the
writing was not in Dr. Gray's hand and so was not considered
in the search for the type.

2. The specimen of argentea is the best one of the series and
shows mature fruits as well as flowers.

3. The published description of sericea fits the specimen that
has been selected as the type.

4. Тһе specimen under consideration formed part of Gray's
concept of the variety humile, and the species sericea is, he says,
the same as that, in large part.

5. The range given by Dr. Gray includes the locality in which
the specimen by Engelmann was collected.

6. No other species included in Gray's sericea has so good a
claim to be considered the typical form as does the plant that
has been known as Огеосатуа argentea.

It seems to have been generally assumed that Bridger's Pass
is in Montana, and so the effort has been made to search for
typical sericea in that state. Тһе Bridger’s Pass of Henry
Engelmann was surely in southern Wyoming, west of the Platte
River. This is near the northern limit of the range of О. argentea
Rydb.

It is possible that further collections of O. procera Osterhout
and of Jones' plant from Utah may prove them worthy of varietal
rank. Тһе author so designated them at one time.

C. sericea, as now understood, is chiefly distinguished by the
lack of pustulate hairs on the upper leaf surface and the con-
sequent sericeous appearance of the indument. Пт. Rydberg
evidently had quite an erroneous idea of the variety perennis
when he reported it from North Dakota and Idaho in his flora of
the Rocky Mountains.

1927]
PAYSON—SECTION OREOCARYA ОҒ СВУРТАМТНА 295

23. С. aperta (Eastw.) new comb. Plate 28, figs. 62-64.

Oreocarya aperta Eastw. Bull. Torr. Bot. Club 30: 241. 1903.

Perennial, caudex branched from a woody root; stems several,
rather slender, 1-2 dm. high; leaves densely clustered at base and
the dilated petioles imbricated, pubescence of upper and lower
leaf surfaces similar, sparsely strigose and abundantly pustulate-
setose, bristles conspicuous, spreading; radical leaves spatulate to
oblanceolate, 3 cm. long, 8 mm. wide; cauline leaves oblanceolate,
about as long as the radical leaves but narrower; inflorescence
branched from near the base with many spreading, simple or 2-
forked spikes, those of all the stems aggregated into a closely
branched thyrsus, spikes peduncled, longer than the subtending
leaves, pedicels very short, about 0.5 mm. long; calyx densely
setose, sepals in anthesis linear or linear-lanceolate, acute, about
3 mm. long, in fruit 8-10 mm. long, exceeding the nutlets about
5 mm.; corolla white, tube 2.5-3 mm. long, equalling or some-
what shorter than the sepals, crests at the base of the tube evident,
fornices conspicuously papillose, probably yellow, nearly 1 mm.
long, limb about 6 mm. broad, tube and limb subequal, the lobes
united for about !4 their length; fruit ovoid, all four nutlets com-
monly maturing, style exceeding the nutlets by about 2 mm.;
nutlets lanceolate, acute, 2.5-3 mm. long, margins in contact,
acute, surfaces of nutlets somewhat glossy, the dorsal tuberculate,
more or less rugose and somewhat muriculate, ventral surface ir-
regularly and ambiguously roughened, scar extending from near
the base to near the apex, straight, closed, no elevated margin
present.

Distribution: Upper Sonoran Zone, Grand Junction, Colorado,
June 27, 1892, Eastwood.

Specimen examined:

Colorado: Grand Junction, Мау 17, 1892, Eastwood (Calif.).

C. aperta 1з another species of this genus of which more material
is badly needed. Presumably it is distinct from all others but it
is so imperfectly known that it is impossible to know to what
other species to ally it.

24. C. rugulosa (Payson) new comb. Plate 28, figs. 65-67.
Oreocarya rugulosa Payson, Univ. Wyo. Publ. Bot. 1: 166.
1926.

[Vor. 14
296 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Short-lived perennial; stems few to many, unbranched above
the base, rather slender, 15-80 ст. high, rather sparsely but con-
spieuously setose; leaves narrowly oblanceolate or spatulate,
obtuse, 2-4 ст. long, strigose-canescent and setose, pustulate
hairs abundant on both surfaces, slightly more numerous on
dorsal, petioles ciliate with long, rather weak white hairs; in-
florescence mostly confined to the upper l4 or № of the stem,
cymes elongating, foliar bracts inconspicuous, inflorescence dense-
ly setose with long, white, slender hairs; calyx subtomentose and
abundantly setose, sepals in anthesis lanceolate, acute, 4 mm.
long, in fruit linear-lanceolate, 7-8 mm. long, exceeding the nut-
lets by about 5 mm.; corolla white, tube 3-4 mm. long, equalling
the sepals in anthesis, crests at the base of the tube well devel-
oped and conspicuous, fornices nearly closing the throat, well
developed, not over 0.5 mm. long, distinctly papillose, probably
yellow, limb about 6 mm. broad, tube and limb subequal, lobes
united for i4 their length; fruit ovoid, all four nutlets usually
maturing, style exceeding the mature nutlets 1-1.5 mm.; nutlets
lanceolate, subacute, 3 mm. long, margins in contact, acute,
surfaces of nutlets somewhat glossy, the dorsal distinctly rugose
with rather distant, low rugae, somewhat tuberculate also, ventral
surface only slightly uneven, scar straight, extending from near the
base almost to the apex, closed above, slightly open near the base,
no elevated margin present.

Distribution: Upper Sonoran Zone, western Utah. Туре:
Fish Springs, Utah, M. Е. Jones.

Specimen examined:

Utah: Fish Springs, June 4, 1891, Jones (R.Mt., TYPE, Pomona,
Calif.).

Тһе most outstanding characteristic of this plant is the smooth
inner surfaces of the nutlets. It is to be considered a member of
the spiculifera-interrupta-Sheldonii group of species.

25. C. interrupta (Greene) new comb. Plate 28, figs. 68-70.

Oreocarya interrupta Greene, Pittonia 3: 111. 1896.

Long-lived, caespitose perennial; stems few to many from the
branching caudex, rather slender, 3-9 dm. high, rather sparsely
strigose and setose with slender white trichomes; radical leaves

1927]
PAYSON—SECTION OREOCARYA ОЕ СВУРТАМТНА 297

tufted on the caudices, oblanceolate, obtuse or acute, 3-5 cm.
long, the blade tapering gradually to a hispid-ciliate petiole,
lower leaf-surfaces rather densely strigose or subtomentose,
setose with subappressed and rather inconspicuous bristles,
pustulate, upper surfaces more finely strigose and very incon-
spicuously setose, pustules minute; stem-leaves similar, reduced
and narrower upwards; inflorescence densely setose with white
hairs, more or less interrupted, narrow cymes somewhat elongat-
ing at the top, stems bearing cymes for 1%-% of their length;
calyx setose, sepals in anthesis about 3 mm. long, linear or
linear-lanceolate, in fruit 6-8 mm. long, exceeding the nutlets
by 4-5 mm.; corolla white, tube З mm. long, equalling the sepals
in anthesis, crests at the base of the tube well developed, fornices
probably pale yellow, low, scarcely papillose, limb 5-7 mm. broad,
tube and limb subequal or the tube slightly longer, lobes united
for 14 their length; fruit lanceolate-ovoid, 1-4 nutlets maturing,
style exceeding the mature nutlets by less than 1 mm.; nutlets
lanceolate, obtuse or subacute, 3 mm. long, margins in contact
or nearly so, acute, surfaces of nutlets dull or slightly glossy, the
dorsal definitely tuberculate with small tubercles, a few of these
elongated on some nutlets, but the nutlets only rarely rugose,
indefinitely muriculate, especially toward the apex and the mar-
gins, ventral surfaces similar but with fewer tubercles and muricu-
lations, scar straight, extending from near the base to near the
apex, closed or nearly so, margin not elevated.

Distribution: Upper Sonoran Zone in northeastern Nevada.
Type: “іп open woods some miles east of Wells," Greene.

Specimens examined:

Nevada: Park’s Station, 25 miles north of Elko, Aug. 4, 1913,
Hitchcock 1005 (U.S.); Elko, Aug. 2, 1913, Hitchcock 929 (U.S.);
Humboldt Wells, Elko Co., July 28, 1908, Heller 9185 (Mo.,
N.Y., U.S., Phila.).

This plant is imperfectly known but is apparently most closely
related to C. spiculifera. From that species it differs in the less
conspicuously setose, broader radical leaves, the tuberculate and
not rugose nutlets, and the shorter style. It has not been possible
to locate the type but specimens from near the type locality agree
very well with the original description and there is probably no

[Vor. 14
298 ANNALS OF THE MISSOURI BOTANICAL GARDEN

doubt but that the species has been correctly identified. Хо
specimens have been seen that were so high as the plants de-
scribed by Greene. Не says “11% to 3 feet high." The speci-
mens at hand are all about one foot high.

26. C. spiculifera (Piper) new comb. Plate 28, figs. 71-73.
Oreocarya spiculifera Piper, Contr. U. S. Nat. Herb. 11: 481.
1906.

О. cilio-hirsuta Nels. & Macbr. Bot. Gaz. 55: 378. 1918.
(Type: Nelson & Macbride 1799, Minidoka, Idaho, June 23, 1912.)

Long-lived, caespitose perennial; stems few to many from the
base, slender, 15-30 cm. high, strigose, hirsute and setose with
long slender white hairs; leaves linear-oblanceolate or linear-
spatulate, acute or obtuse, 3-7 ст. long, dorsal surface densely
strigose and with rather numerous pustulate, spreading setae,
ventral surface densely strigose, sparsely setose with somewhat
weaker and more closely appressed pustulate hairs, petioles con-
spicuously long-ciliate; inflorescence оп the upper 1% or 24 of the
stem, densely setose with long, white, slender hairs, cymules
somewhat elongating, foliar bracts mostly inconspicuous; calyx
strigose and densely setose, sepals in anthesis 4-5 mm. long,
linear-lanceolate, in fruit 8-10 mm. long, exceeding the nutlets
by 6-7 mm.; corolla white, tube 3-4 mm. long, equalling the
sepals in anthesis, crests at the base of the tube evident, fornices
well developed, at least 0.5 mm. high, scarcely papillose, pale
yellow, limb about 7 mm. broad, tube and limb subequal or the
tube slightly longer, lobes united for 14 their length; fruit ovoid,
all four nutlets usually maturing, style exceeding the mature
nutlets by about 2 mm.; nutlets lanceolate, acute, 3-4 mm. long,
margins usually in contact or nearly so, acute or obtuse, surfaces
of nutlets dull, the dorsal definitely but narrowly margined,
densely muriculate, rugose and sparsely tuberculate, ventral sur-
faces densely muriculate, sparsely tuberculate and with a few
short rugae, scar straight, extending from near the base to near
the apex, narrowly open, no elevated margin present.

Distribution: Upper Sonoran Zone of western Washington
and south-central Idaho. Type: “collected at Ritzville, Adams
County, by Sandberg & Leiberg (No. 164), June 6, 1893.”

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 299

Specimens examined:

Idaho: Salmon, June 23, 1920, Payson & Payson 1768 (R.Mt.,
Mo., Gray); Arco, June 19, 1893, Palmer 189 (U.S.); Challis,
July 15, 1916, Macbride & Payson 3223 (R.Mt., Gray); Martin,
July 5, 1916, Macbride & Payson 3039 (R.Mt., Mo., U.S., Gray);
Minidoka, June 23, 1912, Nelson & Macbride 1799 (R. Mt., Mo.,
U.S., Gray); Shoshone, May 27, 1899, Saunders 4875 (Mo.);
Boise-Payette Project, June 2, 1911, Macbride 875 (Pomona, R.
Mt., Minn., Field, Mo., U.S., Gray); New Plymouth, May 21,
1910, Macbride 93 (R.Mt.); Weiser, April 26, 1900, Jones 6673
(Pomona).

Washington: Wilson Creek, eastern Washington, June, 1893,
Sandberg & Leiberg (Pomona, Minn.); Ritzville, June 6, 1893,
Sandberg & Leiberg 164 (Wash., Calif., Mo., U.S., түре, Gray).

C. spiculifera is а fairly well-defined unit characterized by
narrow radical leaves with conspicuously setose petioles. The
caudex is multicipital and the leaves are very numerous in typi-
cal forms. It is likely to be confused with C. Macounit, and at one
time the author was inclined to unite the two as varieties under
one specific name. This process of submergence would lead to
such wholesale reductions, however—Sheldonii, Macounti, spicu-
lifera, interrupta, rugulosa, Bradburiana, sobolifera—that it
seemed unwise. C. spiculifera has some contact, too, with
C. nana var. Shantzii. А specimen from southeastern Idaho
was examined that looked remarkably like spiculifera but the
nutlets were uniformly muricate as in the var. Shantzit. This
intermediate plant might, of course, have been a hybrid.

27. C. celosioides (Eastw.) new comb. | Plate 28, figs. 74-76.

Oreocarya celosioides Eastw. Bull. Torr. Bot. Club 30: 240.
1903.

Long-lived perennial from a stout, woody root; stems 1-several
from a leafy caudex which is densely clothed xii the broad im-
bricated petioles of former leaves, stout, 2-4 dm. high, densely
setose with stiff divaricate bristles; leaves clustered at the base
of the stem—or only the petioles of former leaves remaining to
show the basal tuft—spatulate to oblanceolate, usually obtuse,
2-5 cm. long, densely setose with spreading white bristles and

[Уог. 14
300 ANNALS OF THE MISSOURI BOTANICAL GARDEN

subtomentose, densely pustulate on both surfaces, petioles con-
spicuously ciliate; inflorescence extending over upper 1% to 24
of stem, becoming rather broad due to the elongation of the
cymules, densely setose, foliar bracts inconspicuous; calyx densely
setose with spreading bristles, sepals in anthesis about 5 mm. long,
linear-lanceolate, acute in fruit, about 12 mm. long, exceeding
the nutlets by about 8 mm.; corolla white, tube 4-5 mm. long,
about as long as the sepals, crests at the base of the tube well-
developed, conspicuous, fornices nearly 1 mm. high, probably
yellow, slightly papillose, limb about 8 mm. across, lobes and
tube subequal, lobes united for 14 to V4 their length; fruit ovoid,
2-4 nutlets maturing, style exceeding the mature nutlets by
2.5-3 mm.; nutlets 4-5 mm. long, ovate-lanceolate, acute,
margins acute, almost winged, in contact, surfaces of nutlets dull
or slightly glossy, the dorsal conspicuously rugose toward the
middle and muriculate toward the edge, somewhat tuberculate
also, in the type collection slightly keeled, the ventral surfaces
rugose or tuberculate and somewhat muriculate, scar straight,
closed, extending from near the base to near the apex, margin not
elevated.

Distribution: Upper Sonoran Zone along the Columbia River
in Washington and in the drainage of the John Day River in
Oregon. Туре: “Thos. J. Howell, from the banks of the Colum-
bia River, eastern Washington, July, 1881."

Specimens examined:

Washington: Washington Terr. 1883, Brandegee 996 (Calif.,
Gray); Klickitat Hills, May, 1888, Howell 431 (Wash., Calif.) ;
banks of Columbia River, July, 1881, Howell (Calif.); Rock
Island, Kittitas Co., July 12, 1893, Sandberg & Leiberg 440
(Wash., Calif., Minn., U.S., Gray); near Columbus, June 10,
1886, Suksdorf 888 (Calif., Field, Mo., U.S., Gray); Wenatchee,
1895, Whited (Wash.); Wenatchee, May 28, 1899, Whited 1099
(Wash., U.S., Gray).

Oregon: Dalles, May, 1882, Howell (Phila.); Mitchell, Aug. 1,
1917, Lawrence 1030 (U.S.); near Fossil, Gilliam Co., Мау 29,
1894, Leiberg 125 (U.S.); Canyon City, July, 1902, Griffiths &
Hunter 170 (U.S.); dry hills near Fossil, Wheeler Co., June 2,
1925, Henderson 5212 (Mo., Gray); Kimberly, Grant Co., June

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 301

24, 1925, Henderson 5212 (Mo., Gray); dry slope, Clarno, south-
east Wasco Co., July 3, 1921, Peck 10020 (N.Y.).

I would limit this plant to the Upper Sonoran Zone of central
Washington and Oregon. This is in essential agreement with
Piper in his ‘Flora of Washington’ except that he includes one
plant (Cotton 359 from the Rattlesnake Hills) which I refer, rather
dubiously, to C. Масоити. He says that celosioides comes from
the “Arid Transition" Zone but according to the map of life
zones in his ‘Flora,’ the localities in which celosioides are found
are all in the Upper Sonoran Zone. Тһе large nutlets and stout
stems are the best characters of celosioides that may be used to
separate it from its nearest allies—C. Sheldonii and C. spiculifera.

28. C. Sheldonii (Brand) new comb. Plate 28, figs. 77-79.

Oreocarya sericea Piper, Contr. U.S. Nat. Herb. 11: 482. 1906,
not Krynitzkia sericea Gray.

O. celosioides Macbr. Contr. Gray Herb. 48: 29. 1916, in part,
as to specimens cited, not О. celosioides Eastw.

О. glomerata Standley, Contr. U. S. Nat. Herb. 22: 401. 1921,
not Cynoglossum glomeratum Pursh.

O. Sheldonii Brand, Fedde, Rep. Sp. Nov. 19: 73. 1923.

Definitely perennial, rather loosely caespitose; stem rather
stout, 2-3 ст. high, abundantly setose with rather slender, di-
varicate hairs; basal leaves numerous, spatulate to oblanceolate,
usually obtuse, 2-4 em. long, 5-8 mm. broad, abundantly setose
with rather weak subappressed hairs, the older leaves distinctly
tomentose, pustules present on both leaf surfaces; cauline leaves
similar, reduced upwards; inflorescence setose, rather narrow or
the branches elongating in age, mainly uninterrupted, extending
over 15 to 34 of the stem, foliar bracts inconspicuous; calyx
densely setose, sepals in anthesis linear-lanceolate, acute, 3-5
mm. long, in fruit about 8 mm. long, exceeding the nutlets by
4-5 mm.; corolla white, tube 4 mm. long, subequal to the sepals
in anthesis, crests at the base of the tube evident, fornices 0.5
mm. high, probably yellow, papillose, limb 5-10 mm. broad,
limb and tube subequal or the tube slightly longer than the limb,
lobes united for about 15 their length; fruit ovoid or lanceolate-
ovoid, all four nutlets commonly maturing, style exceeding the

[Vor. 14
302 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

nutlets by about 2 mm.; nutlets lanceolate, acute or subacute,
3-4 mm. long, margins jn contact, acute, surfaces of nutlets
slightly glossy, the dorsal tuberculate, usually more or less rugose
and obscurely muricate, ventral surface more or less tuberculate;
scar straight, extending from near the base to near the apex, closed,
no elevated margin present.

Distribution: Transition Zone of western Montana, northern
Idaho, eastern Washington, and northeastern Oregon. Туре:
Deep Creek, Wallowa Co., Oregon, Sheldon 8315.

Specimens examined:

Montana: St. Mary Lake, Glacier Nat’l. Park, Aug. 8, 1919,
Standley 17423 (U.S.); Glacier Park Station, July 6, 1919,
Standley 15107 (U.S.); Glacier Park Station, Aug. 15, 1919,
Standley 17594 (U.S.); Great Falls, June 16, 1891, Williams 109
(Minn., U.S.); Helena, 1891, Е. М. Brandegee (Calif.); Hallett's
Ranch, Aug. 14, 1892, Е. №. Brandegee 17 (Calif.); Helena, June
& July, 1892, Aiton 64 (Minn.); Deer Lodge Valley, July 19,
1905, Jones (Pomona); Missoula, June 10, 1922, Kirkwood 1262
(Mont.); near Missoula, June 12, 1901, MacDougal 169, this
specimen is intermediate to No. 31 (C. Bradburiana) (U.S.,
Mont., Gray); upland benches, Hamilton, June 16, 1906, Blankin-
ship 734 (Field, U.S., Mont.); on Madison River, 4300 ft.,
June 28, 1860, Hayden (Mo.); 10 miles east of Monida, June 18,
1899, А. & Е. Nelson 5429 (R.Mt., Mo.); Shinberger’s Canyon,
southern Montana, July 15, 1880, Watson 287 (Gray).

Idaho: Kootenai Co., July, 1880, Sandberg (Minn.); in the
Palouse country and about Lake Coeur d’Alene, June—July,
1892, Aiton 31 (Minn.).

Washington: Spokane, May 16, 1896, Piper 2294 (Wash.,
Gray); Hangman Creek, Spokane, May 23, 1897, Piper (Wash.);
Spokane River, June 2, 1892, Henderson 2563 (Wash., Gray);
Spokane, June 19, 1913, T'uresson (R.Mt.); between mouth of
Spokane and Colville, 1838-42, Wilkes 437 (61), (Phila.).

Oregon: Deep Creek, Wallowa Co., June 16, 1897, Sheldon 8315
(N.Y., Mo., 0.5.).

This species as here defined is composed of rather different in-
dividuals from a large range. Mature specimens are still rather
few, and the specific limits are sometimes difficult of location.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 303

The plant is obviously a relative of C. Bradburiana in spite of
Brand's suggestion that it is closely related to leucophaea. From
Bradburiana it is separated because of the perennial, caespitose
base and the western range. Macbride confused it, in part,
with celosioides. From that species Sheldonii may be distin-
guished by the smaller, differently marked nutlets, the more
slender stems, and the more easterly range. It is also likely to
be confused at times with spiculifera. That plant has narrower
leaves that are strongly ciliate-hirsute, a stronger tendency to a
multicipital caudex, somewhat different nutlets, and a more
southern and eastern range. С. Macounii is the other perennial
relative of C. Bradburiana. "That plant is characterized by nar-
row, strongly setose leaves, while Sheldon?i has broad, tomentose
ones. The range of Macounii is more northern than the others.
Piper recognized the species as distinct from celosioides (and
glomerata, of course) but supposed it to be sericea. То this
Macbride makes an interesting comment, ‘‘ Piper may be justified
in distinguishing two species here, but if the material from
eastern Washington [Sheldonii] represents a species distinct from
that of the Columbia Valley [celosioides], it cannot bear the name
O. sericea which must be used to designate a very different plant
of the Rocky Mountains." With sericea now identified with
Rydberg's argentea, Macbride's comment is particularly pertinent.

29. C. Macounii (Eastw.) new comb. Plate 29, figs. 80-82.

Eritrichium glomeratum Macoun, Cat. Canad. Pl. 1: 337-338.
1883, in part.

Oreocarya Macounii Eastw. Bull. Torr. Bot. Club 40: 480.

13

Caespitose, long-lived perennial from a rather slender tap-root;
stems slender, 10-20 cm. high, conspicuously setose with divari-
cate bristles; leaves linear to linear-oblanceolate, 2-5 cm. long,
acute or obtuse, conspicuously setose with spreading bristles
(partieularly on the petioles), leaf-surfaces strigose, more densely
pustulate below than above; inflorescence narrow, mainly
limited to upper 14 ог 16 of the stem, densely setose, foliar bracts
sometimes rather conspicuous near the base of the inflorescence;
calyx densely setose as well as strigose, sepals linear-lanceolate,

[Vor. 14
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN

acute, about 3 mm. long in anthesis, in fruit 6-7 mm. long, ex-
ceeding the nutlets 3-4 mm.; corolla white, tube about 3 mm.
long, equalling the sepals, crests at the base of the tube evident
but not conspicuous, fornices about 0.5 mm. high, probably
yellow, limb 7-9 mm. broad, tube and limb subequal, lobes
united for about 15 their length; fruit lanceolate-ovoid, all four
nutlets commonly maturing, style exceeding the nutlets 1.5-2
mm.; nutlets lanceolate, acute or subacute, 2.5-3 mm. long,
margins in contact, acute, surfaces of nutlets dull or slightly
glossy, the dorsal rather indeterminately rugose, tuberculate and
muriculate, the ventral surfaces indeterminately muriculate and
tuberculate, scarcely rugose, scar extending from the base to near
the apex, straight, closed except for а very small area at the base,
margin not elevated.

Distribution: rocky hills, from southern Saskatchewan, Al-
berta, and British Columbia into Montana, Wyoming, and
Washington. Туре: Moose Mountain Creek, Saskatchewan,
John Macoun.

Specimens examined:

Saskatchewan: Saskatchewan, 1858, Bourgeau (Gray) ; Carlton
House, 1827 (?), Drummond (Gray) ; Cypress Hills, June 8, 1884,
Macoun 85007 (Canada); Moose Jaw, June 14, 1903, Barber 325
(Gray); Moose Jaw, July 7, 1880, Macoun (Minn.); Chaplin,
June 18, 1896, Macoun 12800 (Pomona, Field, Canada); Moose
Mt. Creek, July 6, 1880, Macoun (N.Y., түре, U.S.); Wood Mt.,
June 5, 1895, Macoun 11819 (Canada); Spy Hill, July 1, 1906,
Macoun 78500 (Canada); Parkby, June 10, 1905, Palmer (U.S.);
South Saskatchewan, Aug. 14, 1872, Macoun 17089 (Canada);
File Hills, July 2, 1879, Macoun 17089 (Canada).

Alberta: Medicine Hat, May 31, 1894, Macoun 5801 (Canada,
Gray); Macleod, May 12, 1913, Moodie (U.S.); Old Man's
River, June 20, 1881, Dawson 17088 (Canada); Milk River,
July 13, 1895, Macoun 11820 (Pomona, Canada).

British Columbia: Similkamen River, June 8, 1905, Macoun
76741: (Wash., Field, Canada).

Montana: Baltic, May 30, 1900, Wilcox 79 (U.S.); Browning,

1 This specimen is not a typical C. Macounii but is intermediate to C. Sheldonit.

1927] Ё
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 305

Sept. 10, 1909, Jones! (Pomona); Lewis & Clark Nat’l. Forest on
Beaver Creek, Aug. 6, 1912, Saunders! (U.S.) ; near Helena, June,
1921, Wooton! (U.S.); Helena, June, 1893, Starz! (Mo.); Helena,
1886, Kelsey 102! (Pomona); Helena, June, 1888, Kelsey! (U.S.);
plains about Helena, June, 1887, Anderson! (Calif., Mo.); Mid-
vale, June 24, 1903, Umbach 149* (U.S.) ; Garrison, July 10, 1909,
Jones! (Pomona); Armstead, June 20, 1920, Payson & Payson
1746? (R.Mt., Mo., Gray).

Wyoming: between Sheridan and Buffalo, June 15-July 15,
1900, Tweedy 3570 (R.Mt.); West Hurlbut Creek, Big Horn
Mts., June 15, 1909, Willits 92 (R.Mt.).

Washington: north side Rattlesnake Mts., May 11, 1901,
Cotton 3591 (R.Mt., Wash., Mo., Phila., U.S., Gray).

C. Macounii, although readily distinguishable in its extreme
form as it is found on the Canadian plains, has rather doubtful
specific limits in the south where it comes in contact with the
ranges of related species. It is likely to be confused with C.
spiculifera, but the author believes that it is in reality more closely
related to C. Bradburiana and C. Sheldonii. It is certain, how-
ever, that all four species are too close for entire convenience and
all may be found to merge. Its right to specific recognition
rests on its narrow leaves, abundant and conspicuous setae, and
small, rather ambiguously marked nutlets.

30. C. sobolifera Payson? Plate 29, figs. 83-85.

Long-lived multicipital perennial; stems branching from the

1 These specimens are not typical C. Macounii. They are from a unified area in
western Montana and are distinctly aberrant. In some ways they are intermediate
between Macounii and Sheldonii, in others they are more like C. Bradburiana.
Further study may make it seem desirable to separate them varietally from Масоитії
on the basis of а more robust habit.

? This is ann a typical C. Macounii, but is intermediate to C. Sheldonii.

sobolifera sp. nov., perennis; caulibus sterilibus gh TI ad apicem

foliatis, i floriferis 15-18 cm. altis hirsutis et setosis; foliis obovato-spathulatis
obtusis 1.5-3.5 cm. longis dense strigosis aut subtomentosis et ны setoso-
hirsutis supra et uicta pustulosis, basis in petiolas gracilibus gradatim attenuatis;
thyrsis angustis longo-setosis; sepalis lineari-lanceolatis 3.5 mm. longis, fructiferis
6-8 mm. longis quam nuculis 4 mm. Mess vetula alba, як 3-4 mm. longo,
calycem non superante, limbo 7 m 1.5-2 mm. superante; nuculis
non nitidis ovato-lanceolatis ыы 3 mm. pe facie exteriore tuberculosa aut
leviter rugosa, faciebus ventralibus fere laevibus, sulco angusto recto, margine non
edita.—Collected by M. E. Jones, Upper Marias Pass, Montana, Sept. 10, 1909
(Pomona, TYPE).

[Vor. 14
306 ANNALS OF THE MISSOURI BOTANICAL GARDEN

root and apparently more or less prostrate or creeping, some of
them sterile and terminating in a cluster of leaves (soboles),
fertile stems 15-18 cm. high, hirsute and strigose; leaves clustered
on the ends of the sterile branches and scattered on the stem,
obovate-spatulate, obtuse, 1.5-3.5 em. long, gradually narrowed
to a slender petiole, dorsal and ventral surfaces similar, densely
strigose or subtomentose and conspicuously setose-hirsute with
long, slender, white bristles, both surfaces pustulate, petioles
densely setose-ciliate; inflorescence on the upper 14 or № of the
stem, cymules slightly elongating, foliar bracts inconspicuous,
inflorescence densely long-setose; calyx strigose as well as setose,
sepals in anthesis 3.5 mm. long, linear-lanceolate, in fruit 6-8
mm. long, exceeding the nutlets by about 4 mm.; corolla white,
tube 3-4 mm. long, equalling the sepals i in айдады crests at the
base of the tube forming a united ring, conspicuous, fornices
nearly 1 mm. long, yellow, emarginate, papillose, limb about
7 mm. broad, tube and limb subequal, lobes united for 4-4
their length; fruit ovoid, all four nutlets commonly maturing,
style exceeding the mature nutlets by 1.5-2 mm.; nutlets ovate-
lanceolate, subacute, 3 mm. long, margins in contact, acute,
surfaces of nutlets dull, the dorsal rather sparsely tuberculate,
sometimes more or less rugose, indistinctly muricate, the ventral
nearly smooth or with a few indefinite tubercles, scar straight, ex-
tending from near the base to near the apex, closed, margins not
elevated.

Distribution: Canadian Zone, northwestern Montana. Туре:
Upper Marias Pass, Montana, M. E. Jones.

Specimens examined:

Montana: Cottonwood Creek, Central Montana, July 30,
1896, Flodman 748 (N.Y.); Upper Marias Pass, 6,000 ft. alt.,
Sept. 10, 1909, Jones (Pomona, TYPE).

It is with much hesitation that the author describes new species
in the Bradburiana group. Тһе specimens cited for this species
are so aberrant, however, that it has been impossible to assign
them to any known species. It is very probable that further
collections in the mountains of Montana will show this plant to
be an important and distinct unit of this perplexing alliance.
The curious development of sterile, creeping stems or soboles

1927]
PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA 307

indicates a habitat on loose earth or gravel slides. "The label on
the type collection bears the statement, “Upper Temperate Life
Zone." This is defined by Jones as the spruce belt. Тһе plant
is evidently one of the very few members of the genus that in-
habits a subarctic region.

31. C. Bradburiana Payson new name. Plate 29, figs. 86-88.
Cynoglossum glomeratum Pursh, Fl. Am. Sept. 2: 729. 1814,
not Cryptantha glomerata Lehm.
M yosotis glomerata (Pursh) Nutt. Fl. Am. Sept. 2: 729. 1814.
Rochelia glomerata (Pursh) Torr. Ann. Lyc. М. У. 2: 226. 1828.
Eritrichium glomeratum (Pursh) DC. Prodr. 10: 131. 1846.
Krymitzkia glomerata (Pursh) Gray, Proc. Am. Acad. 20: 279.
885.

Oreocarya glomerata (Pursh) Greene, Pittonia 1: 58. 1887.

O. affinis Greene, Pittonia 3: 110. 1896 (Type: ‘‘sandy hills
near Red Buttes, Wyoming, 5 July, 1896,” Greene), not Cryp-
tantha affinis (Gray) Greene.

Krynitzkia pustulata Blankinship, Montana Agr. Coll. Sci.
Studies, Bot. 1:96. 1905, not Cryptantha pustulosa (Rydb.) Pay-
son.

Oreocarya perennis Rydb. Fl. Rocky Mountains, 722. 1917,
in part, not O. affinis perennis A. Nels.

Biennial, or short-lived perennial (?) from a tap root; stems
rather stout, simple or branched from the base, if branched from
the base then one stem usually exceeding the others, stems 1.5-
3.5 dm. high, coarsely setose with divaricate bristles; leaves
forming a rosette at the base the first year, radical leaves spatu-
late to oblanceolate or the outermost almost obovate, obtuse,
2-5 ст. long, cauline leaves somewhat narrower and longer,
pubescence similar on both leaf surfaces, setose with spreading
bristles and subtomentose, abundantly pustulate; inflorescence
rather narrow, somewhat glomerate, extending over upper % or
34 of the stem, foliar bracts near the base of the inflorescence
conspicuous and longer than the cymules in the young inflores-
cence; calyx densely setose and hirsute, sepals linear-lanceolate,
acute, in anthesis about 4 mm. long, in fruit 8-10 mm. long, ex-
ceeding the nutlets about 5 mm.; corolla white, tube 3-4 mm.

[Vor. 14
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long, equalling the sepals, crests at the base of the tube con-
spicuous and well developed, fornices low, rounded or emarginate,
about 0.5 mm. long, papillose, pale yellow, limb about 10 mm.
broad, lobes and tube subequal or the lobes slightly longer than
the tube, lobes united for about 15 their length; fruit ovoid, all
four nutlets usually maturing; style exceeding the mature nut-
lets by about 1.5 mm.; nutlets dull or slightly glossy, margins
acute, in contact, the dorsal surface of nutlets more or less rugose-
tuberculate and with at least a few muriculations, ventral surface
indefinitely tuberculate or rugose and somewhat muriculate, scar
straight, closed, extending from near the base to near the apex,
no elevated margin present.

Distribution: Upper Sonoran and Transition Zones, southern
Alberta, North Dakota, western South Dakota and Nebraska,
Montana, Wyoming, and northeastern Colorado. Type: “іп
Upper Louisiana," Bradbury. Bradbury collected this plant at
the Big Bend of the Missouri in what is now South Dakota.

Specimens examined:

Alberta: Lethbridge, June 5, 1894, Macoun 5802 (Mo., Canada,
Gray).

North Dakota: Pembina, Havard (U.S.) ; Minot, June 5, 1908,
Lunell (U.S.); Glen Ullin, June, 1891, Holzinger 6 (U.S.);
Medora, July 17, 1891, Waldron (R.Mt.); N. W. Territory,
Nicollet 374 (Phila.); Marmarth, June 15, 1914, Moyer 453
(Minn.).

South Dakota: bluffs of Missouri River, Walworth Co., June
15, 1909, Moyer (Minn.); Missouri Valley near Ft. Pierre, June
25, 1839, Geyer (U.S.); Ft. Meade, June 9, 1887, Forwood 272
(U.S.); “Louisiana,” Bradbury (Phila.); Interior, Stanley Co.,
June 5, 1914, Over 5188 (U.S.); White River, Washington Co.,
Aug. 5, 1914, Over 6185 (U.S.); Date, Perkins Co., June 1, 1912,
Visher 587 (R.Mt.); Short Pines, Harding Co., June 9, 1911,
Visher 481 (R.Mt.); Cone Hills, Harding Co., July 8, 1920, Over
(U.S.); Crook, July 23, 1910, Visher 295 (R.Mt.); 8 miles north of
Belle Fourche, Butte Co., June 11, 1916, Eggleston 12520 (0.5.);
Belle Fourche River, June 2, 1894, Bailey 14 (U.S.); Piedmont,
June, 1895,* Pratt (R.Mt., Minn.); Rockerville, Black Hills,
June 15-30, 1909, White (Mo.); Hot Springs, June 13, 1892,
Rydberg 893 (0.8.).

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 309

Nebraska: Lavaea, July 14, 1898, Bates (R.Mt.); Hay Springs,
June 6-7, 1901, MacDougal 60 (Mont.); Ft. Robinson, July,
1891, Bates (Minn.); Belmont, July 24, 1889, Webber (Mo.);
Warbonnet Canyon, Williams (U.S.).

Montana: Custer, May 16, 1890, Blankinship (R.Mt., Mo.);
Livingston, June 8, 1906, Blankinship 372a (Pomona, Mo.);
Wreck Creek, Sweet Grass Co., June 15-19, 1912, Eggleston 7959
(U.S.); Greycliff, Sweet Grass Co., Мау 15-31, 1912, Eggleston
7888 (U.S.); Livingston, June 7, 1901, Scheuber (Minn., U.S.);
Bozeman, May 19, 1900, Moore (Calif.); Bozeman, June 16,
1905, Blankinship 373 (0.8.); Sedan, June 11, 1901, Jones
(R.Mt.); mouth of Shield's River, June 6, 1883, Scribner 174
(Phila., Gray); near Jefferson City, June 27, 1883, Scribner 174
(U.S.); Park Co., 1889, Tweedy (U.S.); near Pony, July 8, 1897,
Rydberg & Bessey 48821 (R.Mt., Wash., Minn., U.S., Mont.,
Gray); Spanish Basin, Gallatin Co., June 23, 1897, Rydberg &
Bessey 48831 (R.Mt., Wash., Minn., U.S., Gray); Park Co.,
1889, Tweedy (U.S.).

Wyoming: Cruse Creek, Sheridan Co., July 6, 1909, Willits
236 (R.Mt.); hills east of Sheridan, June 17, 1912, Sharp 132
(R.Mt.); between Sheridan and Buffalo, June 15-July 15, 1900,
Tweedy 3569 (R.Mt.); south of Buffalo, July 24, 1926, Nelson
10741 (R.Mt.); 40 miles west of Cody, Aug. 31, 1922, von
Schrenk (Мо.); Mammoth Hot Springs, June, 1887, Tweedy
816 (Field, U.S.); Cheyenne River, Williams (R.Mt., Gray);
Wind River, July 14, 1881-82, Forwood (U.S.); head of Middle
Fork of Powder River, July 18, 1901, Goodding 287 (Pomona,
R.Mt., U.S., Gray); well on Salt Creek Road, Natrona Co.,
July 9, 1901, Goodding 230 (R.Mt., Mo., U.S., Gray); Birds Eye,
June 24, 1910, Nelson 9355 (R.Mt., Minn., Gray); Birds Eye,
June 20, 1910, Nelson 9411 (R. Mt., Minn., Gray); Hartville,
June 30, 1901, Nelson 8819 (Pomona, R.Mt., Mo.) ; Uva, July 10,
1894, Nelson 388 (Pomona, R.Mt., Minn., Mo., U.S., Gray);
Chug Creek, Albany Co., July, 1900, Nelson (R.Mt.); Laramie
Hills, June 16, 1894, Nelson 255 (U.S., Gray); Sheep Mt., Albany
Co., June 24, 1925, Payson & Payson 4247 (R.Mt.); Laramie

1 Rydberg & Bessey 4882 and 4883 are not typical C. Bradburiana but are inter-
mediate to C. Sheldonii.

[Vor. 14
310 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

Hills, June 18, 1896, Nelson 1956 (Pomona, R.Mt., Minn., Mo.);
Rock River, June 18, 1901, Goodding 29 (Pomona, R.Mt., Field,
U.S., Gray); Red Buttes, June 18, 1891, Buffum 640 (R.Mt.);
Sand Creek, Albany Co., May 31, 1900, Nelson 6961 (Pomona,
R.Mt., Mo., U.S.); Ft. Steele, June 16, 1907, Nelson 9045 (R.Mt.,
Minn., Mo., Gray); Ft. Steele, June 18, 1898, Nelson 4837
(R.Mt.); Ft. Steele, June 16, 1900, Nelson 7248 (Pomona, R.Mt.,
Minn., U.S., Gray); Ft. Steele, May 25-June 10, 1901, Tweedy
4261 (U.S., Gray); Tipton, June 17, 1898, Nelson 4788 (R.Mt.);
near Table Rock in Red Desert, July 5, 1926, Nelson 10700
(R.Mt.).

Colorado: Pawnee Buttes, July 8, 1920, Osterhout 5927 (R.Mt.,
U.S., Gray, Osterh.).

C. Bradburiana was the first species of this genus to be described
and probably also the first species to find its way into herbaria.
This was due very largely to its wide distribution in an accessible
part of the western United States. Because of the priority of
this species and the homogeneous nature of the genus as a whole,
very many specimens have been determined as glomerata (1.е.,
Bradburiana) that are in reality quite different plants. А number
of varieties of this species have been described from time to time
that are equally far removed from glomerata as it is here under-
stood. Indeed the plants most closely related have been de-
scribed in recent years and have been proposed and maintained
as species since their introduction into scientific literature.

The specific concept of Bradburiana and its allies that has been
adopted in the present paper is too small for entire satisfaction.
With a more inclusive concept the following units would be recog-
nized as varieties only: sobolifera, celosioides, Macounii, inter-
rupta, spiculifera, Sheldon, and possibly rugulosa. This reduc-
tion of specific units is not accomplished in the present paper for
a number of reasons. In the first place the smaller units are
mostly satisfactory in the sense that the intermediates are few
and the geographical ranges isolated and consistent. In the
second place the binomials are somewhat more convenient and
the segregates have been previously maintained by other authors
as species. In the third place if aggregation proceeded to the
point indicated above, there would be good reason to extend the

1927]
PAYSON—SECTION OREOCARYA OF СВУРТАМТНА 311

specific boundaries further and include humilis, sericea, and
aperia. From that it would be only a step to include nana and
its allies, and so the one species would come to contain a large
proportion of the units in the genus. In other words, by main-
taining а smaller specific concept than is entirely satisfactory,
the various units in the genus are kept more nearly uniform in
degree of difference from one another and when one commences `
to aggregate species, it is difficult to stop and the result is not
satisfactory because the species retained are of such very different
values.

The greatest difficulty comes in delimiting Bradburiana,
Sheldonii,and Macounii. In western Montana these forms are
very perplexing and much work yet remains to be done before
an entirely satisfactory solution can be reached. Sheldon is
distinctly and conspicuously perennial and caespitose, while
Bradburiana is typically biennial and solitary. The compara-
tively few intermediates that have come to hand have been cited
with the species they seemed most to resemble. Macounii is
distinctly perennial but has very narrow leaves. This is in
great contrast with Sheldoni but in a few cases it has been dif-
ficult to place a particular plant in one species or the other.
Only one segregate of Bradburiana, in the limited sense, has ever
been proposed, namely, O. affinis Greene. The present author
can see no reason for maintaining this. Macbride (Contr.
Gray Herb. 48: 30. 1916) quotes Dr. Nelson concerning the
validity of affinis over glomerata (i.e., Bradburiana). It has
become evident that Nelson was contrasting C. Bradburiana
and C. thyrsiflora and so the differences given do not relate to the
present question. From Macbride’s correspondence, which is
at hand, it is certain that he maintained affinis almost entirely
on Nelson's judgment.

It is unfortunate to have to substitute an unfamiliar name for
а familiar one but since Cryptantha glomerata Lehm., which is а
valid South American species and the type of the genus Cryp-
tantha, was described long ago, there remains no other alternative
if Oreocarya is merged in Cryptantha.

Тһе name is given in honor of the first collector of the species,
who, it seems, fared rather badly in the matter of scientific recog-
nition for his many hardships and privations.

[Vor. 14
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Series 4. NaNaE. Nutlets conspicuously muricate, in some
species also somewhat rugose. Scar with at least a tendency to
be open and so triangular at the base. Species 32-38.

32. C. nana (Eastw.) new comb.

Oreocarya nana Eastw. Bull. Torr. Bot. Club 30: 243. 1903.

Perennial, more or less densely caespitose; stems 1-many from
а branching caudex, rather slender, 5-25 cm. high; leaves spatu-
late to oblanceolate, usually obtuse, 1-5 cm. long, ventral surface
more or less pustulate but somewhat less abundantly so than the
dorsal surface; inflorescence usually rather narrow, cymules
usually not elongating, foliar bracts not conspicuous; sepals in
anthesis linear to linear-lanceolate, acute, 3.5-5 mm. long, in
fruit 7-10 mm. long, exceeding the nutlets by 3-4 mm.; corolla
white, tube about 3 mm. long, usually distinctly durior than
the calyx lobes in anthesis, crests at base of tube conspicuous or
nearly obsolete, fornices low (less than 1 mm.), rounded, more
or less papillose, probably yellow, limb 7-10 mm. broad, lobes
united for nearly 14 their length; fruit lance-ovoid to ovoid, 1-4
nutlets maturing, margins regularly in contact, acute or obtuse;
nutlets lanceolate to ovate-lanceolate, acute, dorsal, and usually
the ventral surfaces also, densely and uniformly muriculate, not
rugose nor tuberculate, scar slightly open at least, and so more
or less triangular, no elevated margin evident.

Distribution: Upper Sonoran Zone of western Colorado, south-
eastern Idaho, Utah, and eastern and southern Nevada.

Тһе varieties of C. nana may be distinguished as follows:
Stems 15-25 cm. high; leaves rather sparsely subtomentose and setose;

styles 0.5-1 mm. longer than the mature nutlets; central and western

Utah and Santen NOVA Е. x ona tht n 82а. var. commizta
Stems 10-20 em. high; leaves densely tomentose and appressed-setose; styles

scarcely exceeding the mature nutlets; north-central Utah and south-

ТОВ ot eibi ai е oe ETERNA AX өре 825. var. Shantzii
Stems 10-15 cm. high; leaves densely tomentose and appressed-setose; calyx

very densely setose and tomentose; styles exceeding the mature nutlets by

0.5-1 mm.; south-central Utah to southeastern Nevada.......... 82с. var. ovina
Stems 5-12 cm. high; leaves coarsely strigose and setose; calyx densely setose

but scarcely tomentose; styles not exceeding the mature nutlets; western

Colorado to northeastern (Лаһ.............................. 32d. var. typica

32a. Var. commixta (Macbride) new comb.

Oreocarya commixta Macbride, Contr. Gray Herb. 48: 33. 1916.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 313

Perennial; stems 15-25 ст. high; leaves spatulate, obtuse, 3—5
em. long, setose with subappressed or spreading bristles, strigose
or subtomentose; inflorescence rather broad, cymules somewhat
elongating, foliar bracts not conspicuous; calyx densely setose
but scarcely tomentose, in fruit showing some tendency to break
away together with the pedicel; style exceeding the mature
nutlets by 0.5-1.0 mm.

Distribution: Upper Sonoran Zone, central Utah and eastern
Nevada. Туре: sandy slides, Juab, Utah, June 9, 1902, Goodding
1074.

Specimens examined:

Utah: Price, June 11, 1900, Stokes (Calif.); Belknap, June 12,
1900, Stokes (Calif.); Juab, June 9, 1902, Goodding 1074 (R.Mt.,
Mo., U.S., Gray, түре); Vermilion, June 4, 1901, Jones (Pomona);
Monroe, May 12, 1899, Jones (Pomona); Detroit, near Oasis,
May 26, 1891, Jones 6698 (Pomona); Clifton, June 19, 1891,
Jones 6699 (Pomona); Desert Mts., n. w. of Oasis, May 20, 1891,
Jones 6700 (Pomona); Detroit, May 26, 1891, Jones (Pomona);
Granite Mts., Tooele Co., June 7, 1900, Jones 6705 (Pomona).

Nevada: Furber, e. Nevada, June 9, 1891, Jones (Pomona);
Ferguson Spring, June 14, 1900, Jones 6706 (Pomona).

32b. Var. Shantzii (Tidestr.) new comb. Plate 29, figs. 89-91.

Oreocarya Shantzii Tidestr. Proc. Biol. Soc. Wash. 26: 122.
1913.

O. dolosa Macbr. Contr. Gray Herb. 48: 32. 1916. (Type:
College Bench, Logan, Utah, June 4, 1909, Charles Piper Smith
1605).

Long-lived, caespitose perennial; stem 10-20 cm. high; leaves
spatulate, obtuse, 2-5 cm. long, tomentose and appressed-setose
with slender more or less appressed bristles; inflorescence usually
narrow, cymules only slightly elongating, lower foliar bracts
rather conspicuous in the young inflorescence; calyx densely
setose and subtomentose; styles scarcely exceeding the mature
nutlets.

Distribution: Upper Sonoran Zone, central to northern Utah
and southeastern Idaho. Type: College Bench, Logan, Cache
County, Utah, June 4, 1909, Charles Piper Smith 1605.

[Vor. 14
314 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:

Idaho: Pocatello, June 15, 1893, Palmer 598 (U.S.); Pocatello,
spring of 1921, Soth P-9 (R.Mt.); Pocatello, Мау 17, 1909,
Slaughter 7 (R.Mt.); Soda Springs, June 22, 1892, Mulford
(Minn., Gray); Nouman, June 5, 1913, Schoper 28 (R.Mt.);
Montpelier, May 15, 1910, Macbride 2 (R.Mt., Wash., Calif.,
Minn., Field, U.S., Gray).

Utah: Logan, May 12, 1909, Smith 1573 (R.Mt.); Logan, June
4, 1909, Smith 1605 (R.Mt.); Robinson, June 26, 1909, Jones
(Pomona); Salt Lake Co., May 15, 1903, Garrett 266 (R.Mt.,
Gray); Salt Lake City, May 14, 1906, Garrett 1737 (U.S.);
benches near Salt Lake, May 15, 1900, Stokes (Calif.); Salt Lake
City, May 20, 1900, Stokes (Minn., U.S.) ; Tintic Junction, May 9,
1910, Jones (Pomona); City Creek Canyon, April 28, 1889,
Jones (Pomona); City Creek Canyon, May 19, 1880, Jones 6719
(Pomona); Garfield, June 30, 1889, Jones 6709 (Pomona); City
Creek, Salt Lake Co., April 20, 1880, Jones (R.Mt.); City Creek
Canyon, Мау 5, 1880, Jones 1692 (R.Mt., Calif., U.S.); Grants
Station, south of Great Salt Lake, Kearney & Shantz 3098 (U.S.,
TYPE); Ft. Douglas, Oct. 12, 1909, Clemens (R.Mt.); near
Emigration Canyon, April 22, 1908, Clemens (Gray) ; Springville,
4400-5000 feet, May 17, 1913, Hill (Mo.) ; Cottonwood, Johnson's
Pass, Tooele Co., June 6, 1900, Jones (Pomona); Nephi, April 25,
1910, Jones (Pomona); 3 miles south of Redmond, Sevier Co.,
May 31, 1915, Eggleston 11132 (U.S.).

32c. Var. ovina new var.!

Densely caespitose, long-lived perennial; stems 10-15 cm.
high; leaves spatulate or oblanceolate, obtuse, 2-4 cm. long,
tomentose and appressed-setose with rather weak bristles;inflores-
cence narrow, or in age somewhat open due to the slight elongation
of the cymules, lower foliar bracts scarcely conspicuous; calyx
densely setose and tomentose; styles exceeding the mature nutlets
by 0.5-1 mm.

! Cryptantha nana var. ovina Payson var. nov., perennis caespitosa; caulibus 10-
15 cm. altis; foliis spathulatis aut oblanceolatis tomentosis et adpresse setulosis;
sepalis dense tomentosis; nuculis muriculatis; stylo nuculas 0.5-1 mm. superante.
—Collected by Georgia Н. Bentley, in vicinity of Currant, Nye County, Nevada,
June, 1916, (В. Mt., түре).

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 315

Distribution: Upper Sonoran Zone, southern Utah, and
southern Nevada. Туре: vicinity of Currant, Nye County,
Nevada, June, 1916, Georgia Н. Bentley.

Specimens examined:

Nevada: Wa Wa, June 25, 1906, Jones (Pomona); near Currant,
June, 1915, Bentley (R.Mt., TYPE, Mo.); Charleston Mts., 1898,
Purpus 6070 (U.S.).

The variety ovina is most nearly related to the variety Shantzii
and is distinguished from it chiefly by the more woolly leaves and
calyces and the lower, more caespitose habit. From material at
hand it would seem that the styles exceed the mature nutlets
more in ovina than Shantzü.

32d. Var. typica n. var.

Oreocarya nana Eastw. Bull. Torr. Bot. Club 30: 243. 1903.

Long-lived, densely caespitose perennial; stems 5-12 cm. high;
leaves narrowly oblanceolate, 1-3 cm. long, setose with spreading
bristles, rather coarsely strigose also; inflorescence narrow, cy-
mules not elongating, foliar bracts inconspicuous; calyx densely
setose and hirsute but scarcely tomentose; styles not exceeding
the mature nutlets.

Distribution: Upper Sonoran Zone, western Colorado, and
northeastern Utah. Type: “Collected by the author (A. East-
wood) near Grand Junction, Colorado, on the mesa above the
Gunnison River, May 17, 1892.”

Specimens examined:

Colorado: DeBeque, June 9, 1920, Osterhout 5980 (R.Mt., U.S.,
Gray, Osterh.); Grand Junction, June 11, 1920, Osterhout 6010
(Gray, Osterh.); near Grand Junction, May—Oct., 1898, Purpus
(Calif.); Grand Junction, May 17, 1892, Eastwood (R.Mt., Calif.,
Gray).

Utah: north of Duchesne, June 22, 1922, Osterhout 6200
(R.Mt.); Theodore, May 14, 1908, Jones (Pomona).

The aggregate treated here as C. nana is distinguished chiefly
by the short corollas and the uniformly muricate nutlets. These
are characters possessed also by C. cana and C. breviflora but
those species have conspicuously sericeous, minutely pustulate
leaves, while in C. nana the leaves always have the two kinds of

[Vor. 14
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN

trichomes and are either distinctly tomentulose or at least con-
spicuously pustulate.

In any general floristic work, the author would have по hesi-
tation in submerging the varieties recognized here and would
use only the specific name for the assemblage—so close are the
varieties and so numerous the intermediates. Тһеу seem to be
geographie units and so are worthy of discrimination in а more
technical treatment.

33. C. cana (A. Nels.) new comb. Plate 29, figs. 92-94.

Oreocarya cana А. Nels. Bot. Gaz. 34: 30. 1902.

Krynitzkia sericea Nutt., fide Rydb. Fl. Rocky Mountains, 723.
1917. This combination was neither published nor suggested by
Nuttall since he never used the generic name Krynitzkia.

Very densely caespitose long-lived perennial; leaves narrowly
oblanceolate, usually acute, clustered on the crowns of the
caudices, 2—6 ст. long, densely and, to the naked eye, uniformly
silky strigose, older leaves under the lens inconspicuously pustu-
late below and setose with setae that are scarcely differentiated
from the strigose hairs, upper surface obscurely pustulate;
inflorescence narrow, mainly restricted to the upper 1% of the
stem, densely but rather weakly setose, foliar bracts inconspicu-
ous; calyx rather finely setose, sepals in anthesis linear or linear-
lanceolate, acute, 3 mm. long, in fruit 5.5-6 mm. long, exceeding
the mature nutlets by 3-4 mm.; corolla white, tube about 3 mm.
long, as long as the calyx lobes, crests at the base of the tube
evident, fornices distinctly papillose, low, rounded, about 0.5
mm. high, probably yellow, limb about 6 mm. broad, tube and
corolla lobes subequal, lobes united for about 14 their length;
fruit usually unsymmetrical, tending to be lance-ovoid, only one
nutlet usually maturing, style shorter than the mature nutlet;
nutlets lanceolate, acute, 3-3.5 mm. long, margins in contact
(when more than one develops), acute or obtuse, surfaces of nut-
lets dull, the dorsal densely muricate with elongated and some-
times confluent papillae, the ventral surfaces similar but papillae
less elevated, scar open toward the base, rather narrowly triangu-
lar, no elevated margin present.

Distribution: Upper Sonoran Zone, central to southeastern

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 317

Wyoming, northeastern Colorado, and western Nebraska.
Type: gravelly open hilltops, Ft. Laramie, Wyoming, A. Nelson
8309.

Specimens examined:

Nebraska: Ft. Robinson, June 1, 1890, Bates (N.Y.); War
Bonnet, June 24, 1890, Williams (N.Y.); McColligan Canyon,
Deuel Co., June 26, 1891, Rydberg 256 (U.S., N.Y.); Sidney,
May 23, 1922, Nelson (R.Mt.).

Wyoming: Birds Eye, June 25, 1910, Nelson 9367 (R.Mt.);
Pine Bluffs, June 27, 1889, Войт (Minn.); Pine Bluffs, May 14,
1897, Nelson 2876 (R.Mt., Calif., Field); Uva, July 10, 1894,
Nelson 389 (R.Mt.); Ft. Laramie, June 29, 1901, Nelson 8309
(R.Mt., түре, Gray); Platte Hills, Nuttall (Phila.).

Colorado: Pawnee Buttes, Weld Co., June 17, 1919, Osterhout
5902 (R.Mt., U.S., Gray, Osterh.); Tuttle, May 16, 1909, Cary
276 (U.S.).

This is a most distinct species by virtue of its densely caespitose
habit, its silky strigose indument, and muricate nutlets. It has
no near relatives with which it might be confused in the region
where it grows.

34. C. propria (Nels. & Macbr.) new comb.
Plate 29, figs. 95-97.

Krynitzkia fulvocanescens var. idahoensis Jones, Contr. West.
Bot. 13:6. 1910. (Туре: Jones No. 6474, near Weiser, Idaho,
April 28, 1900.)

Oreocarya propria Nels. & Macbr. Bot. Gaz. 62: 145. 1916.

Densely caespitose, long-lived perennial; stems few to many
from the branched, woody caudex, 15-25 cm. high, rather slender,
sparsely and weakly setose, caudex densely clothed with the leaf-
bases of previous years; leaves clustered on the crowns of the
caudex, spatulate, obtuse, 4-8 ст. long, dorsal surface finely
strigose and abundantly appressed-setulose with short pustulate
bristles, ventral surface densely and finely strigose with a few
pustulate hairs, petioles ciliate near the base with long white
hairs; inflorescence narrow, continuous or nearly so, confined to
the upper 16 or 24 of the stem, densely but rather weakly setose,
foliar bracts not conspicuous; calyx densely and weakly setose

[Vor. 14
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and hirsute, sepals in anthesis linear or linear-lanceolate, 4—6
mm. long, acute, in fruit 8-10 mm. long, exceeding the nutlets by
about 5 mm.; corolla white, tube 3.5-5 mm. long, tube and sepals
subequal, crests at base of tube conspicuous, fornices nearly 1 mm.
high, probably yellow, finely papillose, limb about 7 mm. broad,
lobes and tube subequal or the tube slightly longer, lobes united
for about l4 their length; fruit ovoid, four nutlets commonly
maturing, style exceeding the mature nutlets 2-3 mm.; nutlets
lanceolate, acute or nearly so, 3-4 mm. long, margins acute,
surfaces of nutlets dull, the dorsal densely and sinuously rugulose
with narrow but relatively high ridges, conspicuously muricate
near the margins, the ventral surfaces densely but not uniformly
muriculate, these elevations sometimes irregularly confluent,
scar narrow but slightly open, extending from the base nearly to
the apex, no elevated margin evident.

Distribution: clay hillsides in the Upper Sonoran Zone of
western Idaho and adjacent Oregon. Туре: Vale, Malheur
County (Oregon), May 14, 1896, J. B. Leiberg 2049.

Specimens examined:

Idaho: 18 miles below Weiser, April 28, 1900, Jones 6474
(Pomona, Mo.).

Oregon: Malheur Valley near Harper Ranch, June 8, 1896,
Leiberg 2223 (СаШ., U.S., Gray, түре); Vale, Malheur Co., Мау
14, 1896, Leiberg 2049 (Pomona, Calif., Field, U.S., Gray).

It is difficult to say to what species this plant is most nearly
related. It would perhaps be most easily confused with C. cana
or C. breviflora but those are plants far removed from it geograph-
ically, and C. propria may not be very closely related to either.
C. propria must stand as a very distinct specific unit.

35. C. breviflora (Osterh.) new comb.
Oreocarya breviflora Osterhout, Univ. Wyo. Publ. Bot. 1: 169.
1926.

Long-lived perennial from a woody root; stems rather slender,
solitary or few, 15-25 cm. high, white-hairy at the base with
long, straight, appressed trichomes, strigose upwards and hirsute
near the inflorescence; leaves conspicuously clustered near the
base, oblanceolate to spatulate, obtuse, 2.5-5 em. long, uniformly

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 319

silky strigose, pustulate hairs scarcely distinguishable from the
others and difficult of observation in dried material, pustules
very numerous on dorsal surface, few and small on ventral surface;
inflorescence rather narrow, nearly continuous, on upper 16-34
of stem, foliar bracts small, inconspicuous, inflorescence setose-
hispid with divaricate bristles that (sometimes at least) become
yellowish in age; calyx setose, sepals in anthesis linear or nearly
so, acute, 4-5 mm. long, in fruit 8-10 mm. long, exceeding the
nutlets by about 4 mm.; corolla white, tube 3-4 mm. long,
slightly shorter than the sepals, crests evident at the base of the
tube, fornices small, rounded, nearly closing the throat, 1 mm.
or less long, yellow, limb 7-9 mm. in diameter, limb and tube
subequal, lobes united for about 14 their length; fruit ovoid,
usually less than 4 nutlets maturing, style very slightly (0.5
mm.) exceeding the mature nutlets; nutlets lanceolate in outline,
acute or obtuse, 4 mm. long, margins in contact, acute, apices
somewhat spreading, surfaces of nutlets dull, the dorsal uniformly
muriculate, the ventral similar, scar open, triangular, extending
from the base to near the middle of the nutlet, margin not elevated.

Distribution: northeastern Utah, probably in the upper part
of the Upper Sonoran Zone. Type: 616 miles north of Jensen
(Utah), Osterhout 6414.

Specimens examined:

Utah: 61% miles north of Jensen, June 19, 1925, Osterhout 6414
(R.Mt., түрЕ, Osterh.); Uinta Mts., 1902, Langille 113 (N.Y.,
U.S.); Ft. Duchesne, Мау 22, 1908, Jones (Pomona).

C. breviflora is evidently closely related to C. fulvocanescens.
It differs from that species chiefly by the short corolla tube and
the low fornices. The ranges of the two seem consistently
separated.

36. C. fulvocanescens (Gray) new comb.

Eritrichium fulvocanescens Gray, Proc. Am. Acad. 10:61. 1875.

Krynitzkia fulvocanescens Gray, Proc. Am. Acad. 20: 280. 1885.

Oreocarya fulvocanescens (Gray) Greene, Pittonia 1: 58. 1887.

О. nitida Greene, Pl. Baker. 3:21. 1901. (Туре: ір dry stony
ground at Deer Run, Colorado, 11 June, C. F. Baker 95.)

Perennial and caespitose from a woody root; stems few to

[Vor. 14
320 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

many from the root, rather slender, 12-25 cm. high, white-hairy
at the base with long, straight-appressed trichomes, strigose and
hirsute upwards; leaves conspicuously clustered near the base,
oblanceolate to linear-oblanceolate, usually acute, 2-8 ст. long,
uniformly silky strigose, pustulate hairs scarcely differentiated
and difficult of observation in dried material, pustules numerous
on dorsal surface, fèw and small on ventral surface; inflorescence
rather narrow, nearly continuous, on upper 14 or 34 of the stem,
softly setose-silky at first, later becoming more hispid, bristles
usually turning yellowish with age, foliar bracts small, incon-
spicuous; calyx weakly but densely setose, sepals in anthesis
linear or nearly so, acute, 6-8 mm. long, in fruit about 12 mm.
long, exceeding the nutlets by 6-8 mm.; corolla white, tube 9-11
mm. long, exceeding the sepals by 2-4 mm., crests lacking at the
base of the tube, fornices conspicuous, yellow, elongated, over
1 mm. long, spreading, limb 7-9 mm. in diameter, about l4 as
long as the tube, lobes united for 14 their length or less; fruit
ovoid, 1-2 nutlets only usually maturing, style exceeding the
mature nutlets by 2-5 mm. (species conspicuously heterostyled) ;
nutlets lanceolate in outline, acute or obtuse, 4 mm. long, margins
in contact (when more than 1 nutlet matures), apices somewhat
spreading, margins acute or nearly obtuse, surfaces of nutlets dull,
the dorsal densely and uniformly muriculate, the ventral similar,
scar short, nearly closed, extending from the base about 14 the
distance to the apex, margins not elevated.

Distribution: Upper Sonoran Zone in southwestern Colorado,
northeastern New Mexico, southeastern Utah, and northeastern
Arizona. Type: near Santa Fe, New Mexico, Fendler 632.

Specimens examined:

Colorado: Grand Junction, June 11, 1920, Osterhout 6003
(R.Mt., U.S., Gray, Osterh.); Deer Run, June 11, 1901, Baker 95
(Pomona, R.Mt., Calif., Mo., U.S., Gray); Grand Junction,
May 17, 1892, Eastwood (R.Mt., U.S., Gray); Westwater, May 6,
1891, Jones (Pomona); Naturita, May 4, 1914, Payson 271
(R.Mt., Minn., Field, Gray, Colo.); Naturita, June 26, 1924,
Payson & Payson 3876 (R.Mt., Mo., Gray); Paradox, June 13,
1912, Walker 85 (R.Mt., Pomona, Minn., Mo., U.S., Gray);
Paradox, June 21, 1912, Walker 156 (R.Mt., Minn., U.S., Gray).

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 321

New Mexico: Gallup, June 14, 1916, Eastwood 5596 (U.S.,
Gray); Aztec, Aug. 1899, Baker 561 (Pomona, R.Mt., Mo., U.S.,
Gray); Bloomfield, San Juan Co., 1892, Waring 15 (Phila.);
near Santa Fe, 1847, Fendler 632 (Field, Phila., Mo., U.S., Gray,
TYPE); hills at Santa Fe, May 13, 1897, A. A. & E. G. Heller
3517 (Minn., Mo., 0.8.).

Utah: Cisco, May 2, 1890, Jones (Pomona, R.Mt., Calif.).

Arizona: 18 miles north of Holbrook, May 21, 1901, Hough 39
(U.S.); Holbrook, May 6, 1899, Zuck (U.S.).

This is a very distinct species characterized by the long corollas
with the elongated fornices, the silky indument, and the nutlets
that are densely and uniformly muriculate. It is most likely to
be confused with C. echinoides and C. breviflora. From the
former it differs in the muriculations of the nutlets which may
be acute but are never truly setose; from the latter in the long
corolla and elongated fornices.

37. C. echinoides (Jones) new comb.

Krynitzkia echinoides Jones, Proc. Calif. Acad. Sci. II. 5: 709.
1895.

Oreocarya echinoides (Jones) Macbr. Contr. Gray Herb. 48: 31.
1916, as to synonymy, not as to specimens cited.

Caespitose perennial very similar in general appearance to C.
fulvocanescens; stems 12—20 cm. tall; leaves crowded at the base,
oblanceolate to spatulate, usually acute, 3-6 cm. long, dorsal
surface strigose and densely appressed-setose with numerous
pustulate hairs, ventral surface densely strigose, setae and
pustules nearly or quite lacking, petioles somewhat ciliate at the
base; inflorescence extending over upper 34 of the stem, narrow,
continuous, fulvous-setose, foliar bracts inconspicuous, calyx
densely setose with spreading or divaricate, yellowish (in age)
hairs, sepals in anthesis about 5 mm. long, linear or nearly so,
acute, in fruit about 10 mm. long, exceeding the nutlets by about
6 mm.; corolla white or cream-colored, tube 8-9 mm. long, ex-
ceeding the sepals by 1.5-3 mm., crests at base of tube sometimes
evident, fornices conspicuous, probably yellow, elongated, erect
or spreading, 1-1.5 mm. long, limb 7-9 mm. in diameter, lobes
nearly half as long as the tube, united for 14-15 their length;

ГУ оз. 14
322 ANNALS OF THE MISSOURI BOTANICAL GARDEN

fruit lance-ovoid, fewer than 4 nutlets maturing (usually 1 or 2?),
style exceeding mature nutlets 4-6 mm.; nutlets lanceolate in
outline, nearly acute, about 4 mm. long, margins acute, probably
in contact, surfaces of nutlets not glossy, the dorsal densely and
uniformly papillose, each papilla usually bearing a short seta from
the apex, sometimes several setae from a single large papilla,
median dorsal ridge often evident, ventral surface similar, scar
narrow, often unsymmetrical, extending from the base to about
the middle of the nutlet, margin not elevated.

Distribution: Upper Sonoran Zone in southern Utah and ad-
jacent Arizona. Туре: Pahria Canyon, Utah, 5300 ft. alt., in
red sand, M. Е. Jones 5279p.

Specimens examined:

Utah: Pahria Canyon, Мау 26, 1894, Jones 5297р (Pomona,
TYPE); Cannonville, May 28, 1894, Jones 5312 ac (Pomona,
Calif., U.S.).

Arizona: Navajo Spring, near Johnson City, June 19, 1890,
Jones (Pomona).

C. echinoides was entirely misunderstood and misinterpreted
by Macbride in Proc. Am. Acad. 51: 547. 1916, and in Contr.
Gray Herb. 48: 31. 1916. From the specimens cited by Mac-
bride it 13 evident that he never saw a specimen of true echinoides
and his interpretation of the identity of the species was apparently
obtained from Jones' discussion of the identity of fulvocanescens.
With Jones' specimens at hand there can be no doubt as to the
plant that should bear the name of echinoides. Two collections
are cited by Jones as representing his new species, and these two
are essentially identical. His description very clearly applies to
the collections cited and so the plant with hedgehog-like nutlets
from Pahria Canyon and from Cannonville must be taken as
true echinoides. Jones took Watson’s plant from Utah (Егі-
trichium glomeratum var. fulvocanescens Wats. Bot. King's Exp.
243. 1871) as being the plant that must bear the specific name
fulvocanescens. Тһе first legal publication of this name 13 this
publication of Watson's, and the type of his variety is the Utah
plant collected by Fremont rather than the Fendlerian collection
designated by Gray in manuscript. Whatever the type of
Watson's variety may be, it does not bear on the case in hand.

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 323

The first specific use of the name fulvocanescens is in connection
with Fendler's plant from Santa Fe, and so that plant becomes
the type of the species fulvocanescens of Gray. It is not surprising
that Jones considered his plant from Pahria Canyon and Fendler's
plant from Santa Fe as conspecific; the two are certainly very
closely related.

Although available material of echinoides is somewhat scanty
it seems that the flowers in this species are only slightly dimorphic.
The difference between the point of attachment in stamens from
flowers assumed to represent the two forms is not over 1.5 mm.
In C. fulvocanescens the difference is distinctly greater.

38. C. Jonesiana (Payson) new comb. Plate 29, figs. 98-100.

Oreocarya Jonesiana Payson, Univ. Wyo. Publ. Bot. 1: 168.
1926.

Densely caespitose perennial from a woody root; caudex much
branched, the branches clothed with the remains of leaves of
previous years; stems 6-8 cm. tall, rather slender, sparsely setose
and coarsely strigose; basal leaves numerous, thick, spatulate to
obovate-spatulate, 1.5-3.5 ст. long, obtuse, blade abruptly ог
gradually reduced to the slender petiole, cauline leaves few,
spatulate, smaller, leaf surfaces appressed-setose and coarsely
strigose, the petioles not ciliate, pustulate hairs present and con-
spicuous on both surfaces, somewhat more numerous on dorsal;
inflorescence mostly confined to upper Lé of stem, narrow, cymules
nearly sessile, foliar bracts inconspicuous, inflorescence densely
and coarsely fulvous-setose; calyx densely setose, sepals in
anthesis linear, acute, about 7 mm. long, in fruit about 10 mm.
long, exceeding the nutlets 4-5 mm.; corolla probably white with
yellow fornices, tube 13-15 mm. long, 6-8 mm. longer than the
sepals, rather conspicuously flaring in the throat, crests at the
base of the tube obsolete, fornices very broad and low, spreading,
limb 10-12 mm. broad, lobes not over 1 as long as the tube,
united for about 14 their length; fruit lance-ovoid, apparently
only 2-3 nutlets usually maturing, style exceeding the mature
nutlets 4—5 mm. (species apparently only slightly heterostyled) ;
nutlets lanceolate in outline, obtuse or acute, about 4 mm. long,
margins in contact, acute, apices scarcely spreading, surfaces dull,

[Vor. 14
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN

the dorsal muricate and sometimes distinctly rugose due to the
confluence of the murications, slightly keeled, the elevations
usually acute and often tipped with a distinct seta, ventral surface
similar to the dorsal but elevations somewhat fewer and lower,
scar narrow, straight, open, extending from the base 4/; of distance
to apex, no elevated margin.

Distribution: Upper Sonoran Zone in south-central Utah.
Туре: San Rafael Swell, Jones.

Specimens examined:

Utah: San Rafael Swell, May 15, 1914, Jones (Pomona,
TYPE); San Rafael Swell, May 19, 1914, Jones (Pomona).

This species seems most closely related to C. fulvocanescens and
C. echinoides but is much more distinct from them than they are
from one another. It differs from them by its more densely
caespitose habit, lower stems, harsher pubescence, less conspicu-
ously dimorphic flowers, lower and broader fornices, and slightly
rugose nutlets.

Series 5. FLAVOCULATAE. Nutlets usually deeply and соп-
spicuously rugose and tuberculate, sometimes muricate or foveo-
late also. Scar, except in C. Bakeri, at least slightly open and
then showing a tendency to be constricted above the base, margin
usually elevated. Species 39-46.

39. C. Wetherillii (Eastw.) new comb. Plate 30, figs. 101-103.

Krynitzkia glomerata var. acuta Jones, Zoe 2: 250. 1891.
(Type: “Collected May 2, 1890, at Cisco, Utah.’’)

Oreocarya Wetherillii Eastw. Bull. Torr. Bot. Club 30: 242.
1903.

Short-lived perennial or biennial; stem stout, branched from
the base with one stout stem and often with one or more low
slender ones, 1.5-3 dm. high, densely strigose and conspicuously
setose with long white divaricate bristles; radical leaves spatulate
or oblanceolate, obtuse or acute, 1-3 cm. long, rather coarsely
strigose dorsally and rather shortly setose with spreading but
scarcely divaricate bristles, ventral surface more finely strigose,
not setose, pustules abundant on dorsal surface, nearly or entirely
absent on ventral (sometimes more abundant in age), petioles
broad, scarcely ciliate; cauline leaves narrower, scarcely petioled,

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 325

usually acute, 1.5-5 cm. long; inflorescence on upper 34 of the
stem, rather broad and open, not interrupted, cymules elongat-
ing, setose, foliar bracts rather conspicuous below, diminishing
upwards; calyx abundantly setose, sepals linear-lanceolate, acute,
5-6 mm. long in anthesis, in fruit about 1 cm. long, exceeding the
mature nutlets by about 6 mm.; corolla white, tube 6-7 mm. long,
exceeding the calyx lobes by about 2 mm., crests at the base of
the tube absent or inconspicuous, fornices probably yellow,
conspicuously long-papillose, emarginate, about 1 mm. long,
limb 10-12 mm. broad, tube about twice as long as the lobes,
lobes united for about 1% their length; fruit ovoid, 3-4 nutlets
usually maturing, style exceeding the mature nutlets 3-4 mm.;
nutlets rather broadly lanceolate, obtuse or subacute, about 4 mm.
long, margins acute, in contact, surfaces of nutlets slightly glossy,
dorsal with numerous conspicuous rounded tubercles that are at
times somewhat confluent and so tend to form rugae, not muricu-
late, ventral surface similar but tubercles lower, some confluent,
scar narrow but open, surrounded by a distinct elevated margin.

Distribution: Upper Sonoran Zone, east-central Utah. Туре:
“Collected by the author [Eastwood] in flower and fruit, May 25,
1892, in Court House Wash, near Moab, southeastern Utah."

Specimens examined:

Utah: Court House Wash, near Moab, May 25, 1892, Eastwood
(Calif., part of түре); Cisco, Мау 2, 1891, Jones (Calif., Mo.,
U.S., Gray); Green River, May 7, 1891, Jones (R.Mt., Calif.,
Pomona); Price, 1900, Stokes (Calif.); Price, June 29, 1898,
Jones (Pomona).

C. Wetherillài is most closely related to C. longiflora, and at one
time the author felt that they could not be separated specifically.
However, more careful study has seemed to show that they are
definitely distinct. Тһе differences between the two species
may be contrasted as follows:

С. Wetherillii C. longiflora
Radical leaves distinctly pustu- Pustulate hairs about equally
late below, scarcely so above. distributed on both surfaces.
Species not known to have di- Species with evidently dimor-
morphic flowers (this may be phic flowers.

due to lack of specimens).

326

Fornices long-papillose or ‘‘his-
id."
Corolla tube 6-7 mm. long.
Nutlets tuberculate and more
or less rugose.
Sear open but narrow, slightly

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Forniees scarcely papillose at
all

Corolla tube 12-14 mm. long.

Nutlets distinctly
scarcely tuberculate.
Sear very narrow, sides nearly

rugose,

parallel, elevated margin
not evident or very slight.

broader near the middle,
surrounded by а distinct
elevated margin.

40. C. longiflora (А. Nels.) new comb. Plate 30, figs. 104-106.

Oreocarya longiflora A. Nels. Erythea 7: 67. 1899.

O. horridula Greene, Pl. Baker. 3: 20. 1901. (Type: Deer
Run, Colorado, 11 June, on a dry bank, C. F. Baker 133.)

А short-lived perennial (possibly at times a biennial); stems
several (1-5), unbranched, rather stout, 10-30 cm. tall, conspicu-
ously setose with long, divaricate bristles, hirsute; leaves clustered
near the base of the stem, oblanceolate, spatulate or nearly obo-
vate, obtuse, 2.5-7 ст. long, blade narrowed to a broad, setose
petiole, blade setose with ascending rather than strongly divari-
cate bristles, sparingly hirsute, some smaller contorted hairs also
present on the surface, pustulate hairs about equally distributed
on the two surfaces; inflorescence occupying 34 or more of the
stem, rather broad and loose, bracts not exceeding the flowers,
whole inflorescence strikingly setose with long straight bristles;
calyx strongly setose-hairy, sepals in anthesis linear-lanceolate to
nearly linear, acute, 8-10 mm. long, in fruit 10-12 mm. long,
erect, exceeding the nutlets by 7-10 mm.; corolla white, conspicu-
ously dimorphic, tube 12-14 mm. long, usually exceeding the
calyx lobes by 2-4 mm., crests at the base lacking, fornices not
over 1 mm. long, yellow, broad, rounded, or emarginate, they and
the upper part of the tube papillose, limb about 10mm. in diameter,
lobes 14 to 14 as long as the tube, united about м their length
from throat; fruit ovoid, 2, 3 or 4 nutlets maturing (apparently
usually less than 4), styles exceeding mature nutlets 3-6 mm.;
nutlet margins nearly in contact, acute, surfaces of nutlets pale,
somewhat glossy, the dorsal surface densely tuberculate and more
or less rugose, tubercles and rugae usually rounded on the top, the

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 321

ventral surface similar, scar straight, closed or narrowly open,
surrounded by a slightly elevated tuberculate margin.

Distribution: Upper Sonoran Zone in west-central Colorado
and probably adjacent Utah. Type: “Collected by Prof. C. S.
Crandall, at Palisades, Mesa Co., Colo., May 14, 1898."

Specimens examined:

Colorado: DeBeque, Mesa Co., May 19, 1911, Osterhout 4468
(R.Mt.); Delta, May 20, 1911, Osterhout 4497 (R.Mt.) ; DeBeque,
June 9, 1920, Osterhout 5975 (R.Mt., U.S., Gray); Grand Junc-
tion, May 14, 1892, Eastwood (Pomona, U.S., Gray); Palisades,
Mesa Co., May 14, 1898, Crandall (R.Mt., түре); Westwater,
May 6, 1891, Jones (Pomona); Montrose, May 23, 1912, Payson
35 (R.Mt.); Montrose, June 14, 1914, Payson 659 (R.Mt., Gray).

C. longiflora is perhaps most closely related to C. Wetherillii
and is most likely to be confused with that species. For a
comparison of these two species see the discussion under C.
Wetherilli.

O. horridula Greene has not been seen except for a few frag-
ments from the type. А comparison of mature nutlets with those
of C. longiflora shows them to be most similar; a comparison of
longiflora with the description of horridula reveals no important
differences; the type locality of horridula at Deer Run makes it
very probable that the two are identical.

41. C. tenuis (Eastw.) new comb. Plate 30, figs. 107—109.

Oreocarya tenuis Eastw. Bull. Torr. Bot. Club 30: 244. 1903.

Perennial, more or less caespitose; stems several to many from
the base, slender, 15-25 cm. high, strigose and divaricate-setose
with rather short, weak bristles; leaves clustered at the base,
rather few on the stem, linear-spatulate, obtuse or subacute,
2—4 cm. long, strigose and weakly appressed-setose on the dorsal
surface, uniformly strigose on ventral surface, densely pustulate
dorsally, sparsely, if at all, pustulate ventrally, petioles scarcely
ciliate; inflorescence extending over upper 16 to 34 of the stem,
rather narrow, uninterrupted, rather weakly but abundantly
setose, foliar bracts inconspicuous; calyx strigose and abundantly
short-setose, sepals in anthesis narrowly lanceolate, acute, 4—5
mm. long, in fruit 6-7 mm. long, exceeding the nutlets by 3-4 mm. ;

[Vor. 14
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN

corolla white, tube 6 mm. long, slightly exceeding the sepals in
anthesis (not over 1 mm.), crests at the base of the corolla tube
present or absent, fornices probably yellow, broad, truncate,
emarginate, about 0.5 mm. high, distinctly papillose, limb unusu-
ally campanulate for the genus, about 5 mm. across, tube 1.5-2
times as long as the limb, lobes united for about 14 their length,
anthers with their apices about on a level with the tops of the
fornices; fruit ovoid, 3-4 nutlets usually maturing (?), style ex-
ceeding the mature nutlets by about 4 mm.; nutlets lanceolate in
outline, nearly acute, about 4 mm. long, margins in contact, acute,
surfaces of nutlets dull, the dorsal sharply and deeply rugose, with
an evident but sinuous or flexuous dorsal ridge, sparingly tuber-
culate, not at all muriculate, the ventral surface sharply rugose,
scar narrow but evidently open, surrounded by an elevated
margin. Хо evidence is at hand to show that this species is di-
morphie in flower structure. With so few collections available,
however, it is possible that they may all represent one form.

Distribution: Upper Sonoran Zone in southeastern Utah.
Type: “near Moab, in Court House Wash, southeastern Utah,
May 25, 1892," Alice Eastwood.

Specimens examined:

Utah: without locality, May-Oet. 1898, Purpus (Calif.) ; Court
House Wash., в.е. Utah, May 25, 1892, Eastwood (R.Mt., Cahf.,
Gray).

C. tenuis, although known from but two collections, possesses
characters that seem to prove its right to specific recognition.
These outstanding characteristics are (1) the corolla tube that
slightly, but definitely, exceeds the calyx lobes, (2) the cam-
panulate rather than spreading limb of the corolla, (3) the an-
thers whose apices are on a level with the fornices, (4) the sharply
and deeply rugose nutlets, (5) the narrow leaves and slender erect
stems. As is the case in most species of the section Oreocarya, it
is difficult to say to what species it is most closely related. For
one reason and another it seems nearest to Osterhoutii (from which
it differs by the tall stems and longer corolla tube), to longiflora
(from which it differs in the shorter corolla tube, sharply rugose
nutlets, shorter setae), and to Wetherillit (from which it differs
by the narrower leaves, sharply rugose nutlets, and much smaller
flowers).

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 329

42. C. Osterhoutii (Payson) new comb. Plate 30, figs. 110-112.
Oreocarya Osterhoutii Payson, Univ. Wyo. Publ. Bot. 1: 167.
1926.

Densely caespitose perennial; caudices much branched and
clothed with the leaf-bases of previous years; stems slender, 2-6
ст. high, near the base covered with long, white, mostly ap-
pressed hairs, strigose and weakly setose upwards; leaves spatu-
late or oblanceolate, usually obtuse, 1-1.5 em. long, strigose and
appressed-setose on dorsal surface, almost uniformly strigose on
ventral surface, pustulate hairs lacking on ventral surface, well
developed on dorsal, petioles somewhat ciliate; inflorescence
reduced but open, on upper 34 of the stem, rather softly and
shortly setose, foliar bracts inconspicuous; calyx strigose-hirsute
and sparsely and weakly setose, sepals in anthesis linear-lanceo-
late, acute, 2-4 mm. long, in fruit 5-6 mm. long, exceeding the
mature nutlets by 2-3 mm.; corolla white, tube about 3 mm. long,
equalling the sepals in anthesis, crests at the base of the tube
usually evident, poorly developed, fornices yellow, broad and
low (0.5 mm. long), emarginate, distinctly papillose, limb 5-7
mm. broad, tube and limb subequal, lobes united for about Yg
their length; fruit broadly ovoid, in the material examined only
Т or 2 nutlets have matured, style exceeding the mature nutlets
by about 0.5 mm.; nutlets turgid, lanceolate in outline, acute,
3 mm. long, somewhat incurved margins usually not in contact,
obtuse, surfaces of nutlets somewhat glossy, the dorsal indefinitely
carinate, sharply tuberculate and somewhat rugose, the ventral
tuberculate with fewer tubercles, scar open, extending at least 1%
the length of the nutlet, evident, elevated and tuberculate margin
present.

Distribution: Upper Sonoran Zone in western Colorado and
eastern Utah. Туре: Monument Park, near Grand Junction,
Colorado, Geo. E. Osterhout 6138.

Specimens examined:

Colorado: Grand Junction, Monument Park, June 3, 1921,
Osterhout 6138 (R.Mt., түре, Colo., Osterh.).

Utah: Court House Wash, June 16, 1913, Jones (Pomona).

This species is one of the most densely caespitose members of
the genus. For this reason and because of the short stems it

[Vor. 14
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN

might be confused with C. nana. The resemblance, however,
between the two species is quite superficial. It seems to be most
closely related to C. paradoxa. The short corollas separate it
definitely from that species.

43. C. paradoxa (А. Nels.) new comb. Plate 30, figs. 113-115.

Oreocarya paradoxa A. Nelson, Bot. Gaz. 56: 69. 1918.

O. gypsophila Payson, Bot. Gaz. 60: 380. 1915. (Type: "on
а dry gypsum hill in Paradox Valley, Colorado, alt. slightly over
5000 ft., Payson 458, June 18, 1914.”)

Densely caespitose perennial; caudices much branched and
clothed with the leaf-bases of previous years; stems slender, 5-12
ст. high, somewhat white-hairy near the base, more or less stri-
gose and weakly setose above; leaves crowded on the end of the
branches of the caudex, spatulate or oblanceolate, usually folded,
obtuse, 1.5-3 em. long, strigose and appressed-setose on the dorsal
surface, uniformly strigose on the ventral surface, pustulate hairs
lacking on the ventral surface, well developed on the dorsal,
petioles somewhat ciliate; inflorescence reduced but not con-
gested, on upper 16-34 of stem, hirsute or weakly setose, foliar
bracts inconspicuous; calyx weakly setose with spreading hairs,
sepals in anthesis linear-lanceolate, acute, 4-5 mm. long, in fruit
7-8 mm. long, exceeding the mature nutlets by about 5 mm.;
corolla white, tube 10-12 mm. long, 2-3 times as long as the
sepals, crests at base of tube lacking, fornices yellow, broad and
low (0.5 mm. high), slightly emarginate, distinctly papillose,
limb 10-12 mm. broad, tube about 215 times as long as the lobes,
lobes united for about 14 their length; fruit oblate-ovoid, all 4
nutlets usually maturing, style exceeding the mature nutlets by
4-8 mm. (species conspicuously heterostyled); nutlets lanceolate
in outline, acute or obtuse, turgid, 2-3 mm. long, not incurved,
margins not in contact, obtuse, surfaces of nutlets slightly glossy,
dorsal surface not carinate, densely tuberculate and rugose, defin-
itely margined, the ventral tuberculate, scar open, extending at
least 14 the length of the nutlet, margin slightly elevated.

Distribution: Upper Sonoran Zone, western Colorado, and
eastern Utah. Туре: on dry “gyp” hills, in Paradox Valley,
Colorado, Е. Р. Walker 91.

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 331

Specimens examined:

Colorado: Paradox, Montrose Co., June 17, 1912, Walker 91
(R.Mt., түре, Pomona, Minn., Mo., U.S., Gray); East Paradox
Valley, June 20, 1924, Payson & Payson 4223 (R.Mt., Mo., Gray);
Paradox Valley, June 18, 1914, Payson 458 (R.Mt., Mo., Gray).

Utah: Myton, May 20, 1908, Jones (Pomona).

This species is conspicuously dimorphic in flower structure.
The stamens in the two forms are attached 4-5 mm. apart and
the styles differ in length by about the same amount. In the
original description of paradora the anthers are described as
being “just below the crests.” Оп the type sheet in the Rocky
Mountain Herbarium, three plants are fastened; on two of these
the anthers are as described, on the third plant they are attached
about half way up the tube. In the original description of
gypsophila the anthers are said to be “attached about midway
on the corolla tube." All the specimens preserved on the type
sheet in the Rocky Mountain Herbarium have anthers attached
just below the fornices. This is to be explained by supposing
that the specimens from which the description was drawn were
distributed and other plants were kept to represent the type and
at the time these two specific names were proposed neither Dr.
Nelson nor the writer suspected the dimorphic nature of the
flowers. Due to the fact that the flowers had discolored some-
what in fading, Dr. Nelson described them as yellow. The
author knew his plant to have white flowers. Unfortunately the
specimens of paradoxa had not yet been mounted and so were not
available when gypsophila was described. However, there can
be no doubt but that the two names are completely synonymous.

C. paradoxa is unique in the possession of a long corolla tube,
densely caespitose habit, and conspicuously rugose and tuber-
culate nutlets that are not in contact on the margins. Its nearest
relative is perhaps C. Osterhoutii. 'That species may be distin-
guished at once by the short corollas.

44. C. Bakeri (Greene) new comb. Plate 30, figs. 116—118.

Oreocarya Bakeri Greene, Pittonia 4: 92. 1899.

O. eulophus Rydb. Bull. Torr. Bot. Club 31: 637. 1904.
(Type: Dolores, Colorado, 1892, Crandall.)

[Vor. 14
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Evidently perennial, probably short-lived, from a stout root;
caudex with few to several rather short branches; stems 1-8
(rarely more), rather slender, 12-30 ст. high, densely setose with
horizontal bristles, hirsute; leaves somewhat clustered at the base,
oblanceolate, blade narrowed to a distinct, setose petiole, obtuse,
4-8 cm. long, dorsal surface appressed and rather finely setose with
numerous pustulate hairs and coarsely strigose, ventral surface
densely strigose and with a few inconspicuous, appressed, pus-
tulate setae; inflorescence occupying upper */;, or more, of stem,
somewhat glomerate, foliar bracts few, linear-oblanceolate, ex-
ceeding the cymules, often reflexed; calyx setose and hirsute,
sepals in anthesis lanceolate-acuminate, about 6 mm. long, in
fruit ovate-lanceolate, shortly acuminate, about 9 mm. long, ex-
ceeding mature nutlets 4-5 mm.; corolla white, tube 6-7 mm.
long, usually very slightly longer than the sepals in anthesis,
crests at base of tube evident, fornices yellow, conspicuous, linear,
emarginate, about 2 mm. long, papillose, limb 7-8 mm. broad,
nearly twice as long as the radius of the limb, united portion 1%
as long as the lobes or less; fruit broadly ovoid, 2-3 or (usually)
4 nutlets maturing, style 1-2 mm. longer than nutlets; nutlets
ovate-lanceolate in outline, obtuse, turgid, 3-3.5 mm. long,
margins not closely in contact, obtuse, surfaces of nutlets rather
dull, the dorsal densely and irregularly rugose and tuberculate,
sometimes more or less foveolate, rugae and tubercles usually
acute, ventral surface mainly tuberculate, scar, in dried nutlets,
usually eurved or twisted, closed, surrounded by an elevated,
tuberculate margin.

Distribution: Transition or higher Upper Sonoran Zone in
southwestern Colorado and southeastern Utah. Туре: Collected
on the Мапсоз River sage plains in southern Colorado, by Messrs.
Baker, Earle & Tracy, 8 July, 1898, and distributed under No.
827."

Specimens examined:

Colorado: Meeker, June 8, 1902, Osterhout 2610 (Osterh.) ;
Gunnison, July, 1897, Bethel (Colo.); Gunnison Co., June, 1888,
Eastwood (0.5.); 3 miles south of Ridgway, July 16, 1917, Payson
1079 (R.Mt., Mo., Gray); Ridgway, June 18, 1924, Payson &
Payson 3834 (R.Mt., Mo., Gray); Dolores, June 16, 1892,

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 333

Crandall (N.Y.); Dolores, June 15, 1892, Herb. Colo. Agric. Coll.
(N.Y.); Мапсоз, Aug. 11, 1925, Smith 3936 (R.Mt.); sage plains,
Mancos, July 8, 1898, Baker, Earle & Tracy 827 (Pomona);
Mesa Verde, Aug. 2, 1917, Payson 1135 (R.Mt.); Chimney Rock
Mesa, Piedra, June 24, 1924, Schmoll 1281 (R.Mt., Colo.);
Chimney Rock Mesa, Piedra, June 30, 1924, Schmoll 1346
(R.Mt., Colo.).

Utah: meadow south of Monticello, 1911, Rydberg & Garrett
9130 (N.Y.).

So far as the material at hand shows, the flowers of this species
have only one form. Тһеу have the anthers fastened about
half way up the tube and the stigma reaches at least to the base
of the anthers and possibly higher. Тһе very low position of the
anthers makes it seem probable that another form with anthers
near the top of the tube will be found. Тһе position of the
stigma, however, is that which usually occurs in the species with
only one kind of flower.

This species is а very distinct one characterized by the leaves
that lack pustulate hairs above, the corolla tube that slightly
exceeds the sepals, and the nutlets with a closed scar and an
elevated margin.

45. C. mensana (Jones) new comb. | Plate 30, figs. 119-121.

Krynitzkia mensana Jones, Contr. West. Bot. 13: 4. 1910.

Oreocarya mensana (Jones) Payson, Univ. Wyo. Publ. Bot. 1:
171. 1926.

Rather loosely caespitose perennial; stems several from the
base, 8-12 cm. high, sparsely strigose and abundantly white-
setose with rather slender bristles; leaves rather broadly spatulate
to oblanceolate, 2-7 cm. long, obtuse, lower surface rather
sparsely strigose, setose with spreading, slender bristles that are
pustulate at the base, upper surface more finely strigose, setae
inconspicuous, pustules fewer; inflorescence not greatly exceeding
the radical leaves, somewhat open and paniculate, extending over
3/„-4/s of the stem, foliar bracts large and well-developed; calyx
strigose and conspicuously setose, sepals in anthesis linear-
lanceolate, acute, about 4 mm. long, in fruit 7-8 mm. long, ex-
ceeding the nutlets by 4-5 mm.; corolla white, tube about 4 mm.

[Vor. 14
334 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

long, equalling or shorter than the sepals in anthesis, crests at
the base of the tube nearly but not quite obsolete, fornices prob-
ably yellow, less than 1 mm. long, slightly papillose, limb 6-8
mm. broad, lobes united for about 14 their length, nearly or
quite as long as the tube; fruit very broadly ovoid, 2-4 nutlets
maturing (in the material examined), style exceeding the nutlets
1.5-2 mm.; nutlets lanceolate, obtuse, 3-3.5 mm. long, margins
usually obtuse, definitely separated by an interval in the material
examined, surfaces of nutlets dull or somewhat glossy, the dorsal
conspicuously but rather openly muricate, tuberculate and rugose,
somewhat ridged down the median line, not foveolate, the ventral
surface conspicuously tuberculate, scarcely muricate or rugose,
scar open and very conspicuous, surrounded by a high elevated
margin.

Distribution: Upper Sonoran Zone in south-central Utah.
Type: Emery, Мау 16, 1894, М. E. Jones 5445р.

Specimens examined:

Utah: Emery, June 16, 1894, Jones 5445p (Pomona, ТҮРЕ,
U.S.); San Rafael Swell, May 15, 1914, Jones (Pomona).

С. mensana is certainly most closely related to С. flavoculata
but is probably well worthy of specific rank. The short corolla
tubes serve to distinguish it definitely from that species. In ad-
dition it is of characteristic habit due to the short stems.

46. С. flavoculata (A. Nels.) new comb. Plate 30, figs. 122-124.

Oreocarya flavoculata A. Nels. Erythea 7: 66. 1899.

O. flavoculata spatulata A. Nels. Erythea 7:67. 1899. (Type:
from gravelly hill tops near Evanston, Wyoming, Nelson 2977,
May 29, 1897.)

О. cristata Eastw. Bull. Torr. Bot. Club 30: 244. 1903. (Type:
Grand Junction, Colo., May 17, 1892, Eastwood.)

О. Shockleyi Eastw. Bull. Torr. Bot. Club 30: 245. 1903.
(Type: Miller Mountain, Esmeralda Co., Nevada, elev. 7500 ft.,
Shockley 244.)

О. Eastwoodae Nels. & Kennedy, Muhlenbergia 3: 141. 1908.
(Type: Mormon Mts., Lincoln Co., Nevada, P. B. Kennedy &
L. N. Goodding 146.)

Caespitose perennial from a woody root; stems few to many

1927]
PAYSON—SECTION OREOCARYA ОҒ CRYPTANTHA 335

from the base, rather slender, 10-30 cm. high, strigose and densely
setose with rather slender bristles; leaves linear-oblanceolate to
spatulate, usually obtuse but sometimes acute, 3-8 cm. long,
densely strigose and, usually, rather weakly setose, abundantly
pustulate on dorsal surface, ventral surface sometimes silky-
strigose and quite lacking in pustulate setae, sometimes as densely
pustulate аз on the dorsal surface; inflorescence continuous or
somewhat interrupted, rather narrow, usually extending over
upper 14 or 24 of the stem, densely but rather weakly and softly
setose, foliar bracts not conspicuous; calyx densely setose and
strigose, sepals in anthesis linear-lanceolate, acute, about 5 mm.
long, in fruit lanceolate or broader, acuminate, often spreading
in age, 8-10 mm. long, exceeding the nutlets 5-7 mm.; corolla
white or pale yellow, tube 7-10 mm. long, exceeding the sepals
in anthesis by 1.5-3 mm., crests at the base of the tube lacking,
fornices conspicuously yellow, rather narrow, long (about 1.5
mm.), slightly papillose, limb 8-12 mm. broad, lobes united for
about l4 their length, 1% to 24 as long as the tube; fruit very
broadly ovoid, all four nutlets commonly maturing, style exceed-
ing the nutlets 4-8 mm. (species moderately heterostyled);
nutlets lanceolate to ovate, mostly obtuse, 2.5-3.5 mm. long,
margins usually obtuse, in contact or separated by a definite
interval; surfaces of nutlets somewhat glossy, the dorsal muricu-
late, tuberculate and usually more or less rugose, sometimes
almost foveolate, the ventral surface conspicuously tuberculate,
scarcely muriculate, rarely rugose, scar open and very conspicuous,
surrounded by a high elevated margin.

Distribution: Upper Sonoran and Transition Zones, Wyoming,
western Colorado, Utah, Nevada, except the northern third, and
southeastern California. Type: Piedmont, Wyoming, June 7,
1898, A. Nelson 4572.

Specimens examined:

Wyoming: Trapper Canyon, Big Horn Co., April 20, 1926,
Finley 2 (R.Mt.); Cummins, Albany Co., July 29, 1895, Nelson
1511 (Mo.); Cooper Creek, June 6, 1898, E. Nelson 4337 (R.Mt.,
Mo., U.S., Gray); hills of the Platte in the Rocky Mountains,
Nuttall (Phila.); Cooper Creek, Albany Co., June 18, 1892,
Buffum 642 (R.Mt.); Hanna, Carbon Co., June 17, 1920, Payson

[Vor. 14
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN

& Payson 1689 (R.Mt., Mo., Gray); Ft. Steele, June 18, 1898,
Nelson 4815 (R.Mt., Mo., U.S.); Ft. Steele, May 25-June 10,
1901, T'weedy 4259 (0.8.); Point of Rocks, June 17, 1901, Merrill
& Wilcox 457 (R.Mt., U.S., Gray); Point of Rocks, June 15,
1898, Nelson 4750 (R.Mt.) ; Steamboat Mt., June 9, 1900, Nelson
7090 (R.Mt., Pomona, Minn., Mo., U.S., Gray); plains between
Eden and Big Piney, July 6, 1922, Payson & Payson 2583 (R.Mt.,
Mo.); Green River, June 23, 1896, Jones 6726 (Pomona); between
Evanston and Rock Springs, June 28, 1922, Osterhout 6252
(R.Mt.); Granger, June 13, 1898, Nelson 4749а (R.Mt.); Kem-
merer, June 1, 1907, Nelson 9028 (R.Mt., Minn., Mo., Gray);
Cokeville, June 11, 1898, Nelson 4665 (R.Mt.); Carter, June 9,
1898, Nelson 4616 (R.Mt.); Piedmont, June 7, 1898, Nelson 4572
(R.Mt., түре, Field); Ft. Bridger, 1873, Parry 49 (Gray);
Evanston, June 4, 1898, Nelson 4513 (R.Mt., U.S., Gray).

Colorado: south of Yampa River, Moffat Co., June 21, 1922,
Osterhout 6194 (R.Mt.); MeCoys, Eagle Co., June 14, 1903,
Osterhout 2850 (R.Mt., Osterh.); Glenwood Springs, June 8,
1920, Osterhout 5956 (R.Mt., U.S., Gray, Osterh.); Glenwood
Springs, June 23, 1912, Osterhout 4696 (R.Mt., Osterh.) ; Atchee,
May 27, 1908, Jones (Pomona); Mack, May 27, 1908, Jones
(Pomona); Grand Junction, June 11, 1920, Osterhout 6006 (R.Mt.,
U.S., Gray, Osterh.); Westwater, Мау 6, 1891, Jones 6715
(Pomona); Grand Junction, May 17, 1892, Eastwood (Calif.);
Naturita, June 20, 1924, Payson & Payson 3865 (R.Mt., Мо.,
Gray) ; 10 miles northeast of Nucla, Montrose Co., June 22, 1924,
Payson & Payson 3874, 3875 (R.Mt., Mo., Gray); near mouth of
Tabeguache Creek, Montrose Co., June 6, 1917, Payson 997
(R.Mt.); Naturita, April 22, 1914, Payson 246 (R.Mt., Minn.,
Field, Mo., Gray); east of Montrose, June 15, 1915, Payson 670
(R.Mt., Gray); near Montrose, July 10, 1917, Payson 1046
(R.Mt.,); Naturita, May 22, 1914, Payson 337 (R.Mt., Minn.,
Field, Mo., Gray, Colo.).

Utah: Uinta Mts., July, 1869, Watson 852 (Gray); Brush Creek
Canyon, Uinta Mts., July 7, 1902, Goodding 1282 (R.Mt., Gray);
low hills east of Bear Lake, June 16, 1922, Gilbert (R.Mt.);
north of Duchesne, June 22, 1922, Osterhout 6197 (R.Mt.);
Theodore, May 12, 1908, Jones (Pomona); Robinson, June 26,

1927]
PAYSON—SECTION OREOCARYA OF CRYPTANTHA 337

1909, Jones (Pomona); Dragon, May 25, 1908, Jones (Pomona);
Frisco, June 25, 1880, Jones 6720 (Pomona); Milford, June 19,
1880, Jones 6721 (Pomona); Thistle Junction, June 10, 1900,
Stokes (Calif.); bluffs near Price, June 12, 1900, Stokes (Minn.,
U.S.); Cottonwood, Johnson's Pass, Tooele Co., June 6, 1900,
Jones 6677 (Pomona); Glenwood, 5500 ft., May 22, 1875, Ward
87 (U.S., Gray); Fish Springs, June 4, 1891, Jones (Calif., U.S.);
Sink Valley, Kane Co., June 23, 1890, Jones 6678 (Pomona);
Deep Creek, western Utah, June 6, 1891, Jones (Pomona);
Salina, Exp. Station, Fishlake Forest, May 26, 1915, Eggleston
11118 (U.S.); Cedar City, May 8, 1894, Jones 5204l (Pomona,
U.S., in part); San Pitch Mts., June 24, 1908, T'idestrom 1293
(U.S.).

Nevada: Sprucemont, July 21, 1891, Jones 6715 (Pomona);
Aurum, June 20, 1893, Jones (Pomona, Mo.); Aurum, July 5,
1905, Jones (Pomona); Furber, June 9, 1891, Jones 6717? (Po-
mona); Ferguson Spring, June 14, 1900, Jones (Pomona); Os-
ceola, June 26, 1906, Jones (Pomona); between Austin and Big
Creek ranch, Lander Co., July 26, 1913, Kennedy 4519 (Mo.,
Phila.); between Austin and Carter’s ranch, July 27, 1913,
Hitchcock 755 (0.8.); Mormon Mts., Lincoln Co., July, 1906,
Kennedy & Goodding 146 (R.Mt., Mo., U.S.); east of Carson
Lake, June 4, Lieut. Bryan's Exp. (Mo.); White Mts., near
Sunland, Mineral Co., June 25, 1912, Heller 10508 (Mo., U.S.,
Gray); Miller Mt., Esmeralda Co., 7500 %., May & June,
Shockley 244 (Calif.); rocky soil, Palmetto Range, May-Oct.
1898, Purpus 5869 (Calif., U.S.); Kyle Canyon, Charleston Mts.,
May 25, 1919, Tidestrom 9610 (Мо.).

California: Summit, Owen's Valley, May 22, 1897, Jones 6729
(Pomona, U.S.); Inyo Mts., June 25, 1891, Coville & Funston
2151 (U.S.).

This species certainly possesses a number of different forms but
so far they have seemed unworthy of named segregation from the
main specifico complex. Further study may develop some
characters that will serve for varietal distinctions but at present
these have not been found.

In western Colorado, near Naturita, the author is familiar
with two forms, on а basis of setose-hispid and silky-strigose

[Vor. 14
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN
indument. For a limited locality it would seem that these two
forms are even worthy of specific rank but on examination of a
large series of specimens it appears that the differentiation is
only local. Oreocarya cristata Eastwood has very narrow leaves
and so has a slightly different aspect. Osterhout’s 6006 seems
to be а good match for the type of cristata. O. Shockleyt is also
characterized by narrow, acute leaves. In general appearance
the collection from the Mormon Mts. by Kennedy and Goodding
is very similar to the type collection of O. Shockleyi, nor can the
author see that the nutlets are different.
The difference between the stamen insertion in the two forms
of the flowers is probably never more than 3 mm. and is usually
less than that.

INDEX TO SPECIMENS CITED

In the following index the numbers in italics outside the parenthesis are the col-

lector’s numbers.
to the species in the present monograph.

Aiton, G. B.

31, 64 (28) C. Sheldonit.
Anderson, Е. W

— (29) С. Macouniüi.
Anect, Bro.

164 (1a) C. Jamesii var. ет
Austin, Mrs. В. M. & Bru

2270 с C. nubigena.
Austin, S

— (5) ^ vivide:

Bailey, V.

14 (31) C. о

"m V., F. V. Coville & F. Funston.

— (11) C. Анал 1888 (15) С.
virginensis
Baker, C. F.

563 (1а) C. Jamesii var. multicaulis;
455 (1c) C. Jamesii var. cinerea; 562
(6) C. flava; — (13) C. virgata; 95, 561
(36) C. fulvocanescens.
Baker, C. F., F. S. Earle & S. M. Tracy.
18 (20) C. thyrsiflora; 827 (44) C. Bakeri.
Barber, Н. 8.
118 (1c) С. Jamesti var. cinerea.

Those in black type in the parenthesis are the numbers assigned

Barber, M
325 (29) н Манай.

Bates, Ј. М.
5768, 5797 (le) С. Jamesit var.
typica; — (31) C. Bradburiana; — (33)

. cana.
Beals, Mrs. W. G.
— (4) C. oblata.
Beattie, R. Kent.
3921 (7) C. leucophaea.
Bentley, Georgia H.
— (32c) C. nana var. ovina.

E.
— (6) C. flava; — (44) C. Bakeri.
Bethel, E., F. S. Nielley & I. W. Clokey.
4258 (1e) C. Jamesti var. typica.
Blankinship, J. W.
734 (28) C. — ет 878, —
(31) С. Bradbu
Bodin, J. E.
— (le) C. Jamesii var. per — (20)
C. thyrsiflora; — (33) C. cana.
Bourgeau, E.
— (29) C. Macounii.

1927]

PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA

Bradbury, John.
— (31) C. Bradburiana.
Lj" na E. N.
— (28) C. Sheldonit.
ан ам К.
— (5) С. confertiflora; — (14) C. in-
solita; — (15) C. virginensis; — (16)
C. tumulosa.
Brandegee, T. 8.
— (1с) C. Jamesii var. cinerea; 997
2 leucophaea; 898 (13) C. virgata;
16) C. tumulosa; 4 (20) С. thyrsi-
3d; 996 (27) C. celosioides.
Brewer, W. H.
T ч С. confertiflora; 1887 (10) C.

Bryan, Lieut. ES.
ai C. |
Buffum
640 i e Bradburiana; 642 (46) C.
flavocula

Carlton, I a.

246 (1e) C. Jamesii var. typica.
Carlson, John I.

— (4) C. oblata.

Chestnut, V. K. & E. R. Drew
10) C. nubigena.
Clemens, Mrs. Joseph
lemensae; — (32b) C. nana
zii.

Clements, F. E. & E. S.

102 (13) C. virgata.
Clifton, R. L.

3039 (1a) C. Jamesii var. multicaulis.
Clute, W. N.

37 (1d) C. Jamesii var. disticha.
Cockerell, T. D. A.

— (le) C. Jamesii var. typica.
Colo. Agr. College Herb.

44 Bakeri.

— (le) C. Jamesii var. typica
Cooper, W. В.

117 (13) C. virgata.
Copeland, E. B.

52 (11) C. Clemensae.

339

Cotton, J.
1027 (7) C. leucophaea; 859 (29) C.
Macounit.
Coville, F. V.
1514 (10) C. nubigena.
Coville, F. V. & F. Funston
2151 (46) С. flavoculata.
Coville, F. V. & J. B. Leiberg
91 (18) C. humilis.
Cowen, J. H.
— (1c) C. Jamesii var. cinerea.
Crandall, C. S
— (le) C. Jamesii var. typica; — іш
C. flava; — (13) C. virgata; — (40) С
longiflora; — (44) C. Bakeri.
Cusick, Wm
2028, — (10) C. nubigena.
Davidson, A
608 (1a) C. Jamesii var. multicaulis;
2722 (5) C. confertiflora.
Dawson, G.
17088 (29) C. Macounii.
Dodds, С. 8.
1832 (13) C. virgata; 2077 (20) C.

8366 2 C. Jamesii var. multicaulis;
— (14) C. Jamesii var. ми: -—
(1е) С. н var. typica; — (6) С.
flava; — (21) C. elata; —(23) С. aperta;
— (32d) C. nana var. typica; 5596, —
(36) C. fulvocanescens; — (39) C.
Wetherillii; — (40) C. longiflora; —
(41) С. tenuis; — (44) С. Bakeri; —
(46) C. flavoculata.

Eggert, H.

— (la) C. ген var. multicaulis;

— (3) C. Palm

Eggleston, W. wo
16184 (1а) C. Jamesii var. multicaulis;
11194a, 15168 (1e) С. Jamesii var.
typica; 15379 (20) C. thyrsiflora; 7838,
7959, 12520 (31) C. buriana;
11182 (32b) C. nana var. Браги;
11118 (46) C. flavoculata.

340

Ellis, Charlotte C.
866, 463 (1а) C. Jamesii var. multi-
caulis.

Elmer, A. D. E.
1056 (7) C. leucophaea.

Engelmann, G.
— (10) C. nubigena; — (20) C. thyrsi-
flora

| MN Н.
— (20) С. е — (22) С. sericea.
Еуегтапп, В.
— (20) С. Ба
Беп ег, A.
636 (1a) С. Jamesii var. multicaulis;
632 (36) C. fulvocanescens.
Ferris, R. & R. Duthie.
784 (10) C. nubigena.
Finley, C.
2 (46) C. flavoculata.
Flodman
748 (30) C. sobolifera
n d
272, — a C. Bradburiana;

rrett, A. O
266, ү 737 (32b) С. nana var. Shantzii.
Geyer, C. A

— (31) С, жағынды
Gilbert, H.
— (46) C. flavoculata.
Goldman, E. A.
407 (1a) C. Jamesii var. multicaulis.
Goodding, L. N.
1264, 2349 (1a) C. Jamesii var. mul-
ticaulis; 208 (le) C. Jamesii var.
typica; 814, 2221 (5) C. confertiflora;
53,164 (6) C. flava; 589 (12) C. setosis-
sima; 2286 (14) C. insolita; 830 (15)
C. virginensis; 996 (17) C. modesta;
60 (19) C. caespilosa; 2076 (20) C.
MN 29, 230, 287 (31) C. Brad-
buriana; 1074 (32a) C. mama var.
commixta; 1282 (46) C. flavoculata.
Greene, E
32 (1a) C. Jamesii var. multicaulis;
— (1c) C. Jamesii var. cinerea; 300
(le) C. Jamesii var. typica; — (12)
Б, setosissima;—(18) C. humilis.
Griffiths, David.
4268, 4912 (1а) C. Jamesii var. multi-
caulis; 5300 (12) C. setosissima.

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Griffiths, David & B. Hunter.
170 (27) C. celosioides.
Hall, H. M. & H. D. Babcock.
5270 (5) C. confertiflora.
Hall, H. M. & H. P. Chandler.
7234 (5) C. confertiflora.
Hanson, й.
А 140 (1a) С. Jamesii var. multicaulis;
$99 (4) C. oblata.
Havard, V.
— (1a) C. Jamesii var. multicaulis; —
(1b) C. Jamesii var. laxa; — (4) C
oblata; — (31) C. Bradburiana.
Hayden, F. V.
— (19) C. caespitosa; — (28) C. Shel-
donii.
Heller, A. A.
11016 (1f) C. Jamesii var. abortiva;
8211, 9619 (5) C. confertiflora; 13435
(10) C. nubigena; — ( . humilis;
9185 (25) C. interrupta; 10508 (46) C.

Heller, A. A. & E. G.
8577 (ic) C. James var. cinerea;
8781 (20) C. thyrsiflora; 3517 (36) C.
fulvocanescens.
Henderson, L. F.
5212 (27) C. celosioides; 2563 (28) C.
Sheldonüi.
Hess, W. L.
106 (1e) C. Jamesii var. typica.
Hicks, Mr. & Mrs. G. H
178 (1c) C. Jamesii var. cinerea.
Hill, G. R., Jr.
— (32b) C. nana var. браги.
Hindshaw, H. H.
2 (7) C. leucophaea.
Hitchcock, A. E.
1193 (1c) C. Jamesii var. cinerea;
7 (6) C. flava; 929, 1005 (25) C. in-
terrupta; 755 (46) C. flavoculata.
Hitchcock, А. S
847 (Le) C. Jamesii var. typica.
Holzinger, G. A.
6 (31) C. Bradburiana.
Hough, W.
8 (1c) C. Jamesii var. cinerea; 88 (12)
C. setosissima; 39 (36) C. fulvocanes-
cens

1927]

PAYSON—SECTION OREOCARYA OF CRYPTANTHA

Howell, Tho
481, — (27) C. celosioides.
Tecate, Е. С.
— (1а) C. Jamesii var. multicaulis;
1113 (15) 4 virginensis
James, Edw
— (1e) С. a var. typica.
Johnston, E. L
845 (1e) C. Jamesii var. typica.
Johnston, I. M.
2809 (1e) C. Jamesii var. typica; 2818
(13) C. virgata; 2817 (20) C. thyrsiflora.
Jones, B. J.
— (31) C. Bradburiana.

Jones, M. E.
4007, 4047, 58974, Via 5300, 6685,
6687, — (1a) C. Jamesii var. multi-
Am 54 4042, 6328, 481. 6688, — “ad
С. Jamesii var. cinerea; 66, — (1e) C.

Jamesii var. typica; 5315f, — (1f) С.
Jamesii var. abortiva; 3759 (4) C. ob-
lata; 5144, 52611, 52891, — (5) C. con-
Је оға; 5455c, 5464, 6725, — (6) С.
flava; 56954, 5790т, 5812, 6015al,
60548, 6686 (12) C. setosissima; 296,
(13) C. virgata; — (14) C. insolita; 5125,
5195а, 6728, — (15) С. virginensis;
6692, 6713 (17) C. modesta; 6732, —
(18) С. p 6726 (19) C. caespi-
losa; 972 (20) C. dido yer
(22a) C. sericea var. typica (22b)
C. sericea var. perennis; — “о 4) С.
rugulosa; 6673 (26) С. spiculifera; —
(28) C. Sheldonii; — (29) C. Macounii;
— (30) C. sobolifera; 6698, 6699, 6700,
a) C.

6705, 6706, — (32 nana var.
commixta; 1692, 6709, 6719, — (32b)
C. nana var. Shanizii; (32е) G

nana var. ovina; — (32d) С. nana var.
typica; 6474 (34) C. propria; — (35)
C. breviflora; — (36) C. fulvocanescens;
5297p, 5312ac, — (37) С. echinoides;
— (38) C. Jonesiana; — (39) C
Wetherillii; — (40) C. longiflora; —
(42) C. Osterhoutii; — (43) C. para-
dora; 5445р, (45) C. mensana;
52041, 6677, 6673, 6715, 6716, 6717,
6720, 6721, 6726, 2729, — (46) C
flavoculata.

қ» ас”; А

341

Jones, W. W.

$39 (1а) C. Jamesii var. E
Kearney, T. H. & H. L. Shan

3098 (32b) C. nana var. aom
Kellerman, W. A

6 (1e) C. Jamesii var. typica.
Kellogg, A

102, — (29) C. Macounii.
Kennedy, P. B
1664 (18) C. humilis; 4519 (46) C.
flavoculata.
Kennedy, P. B. & L. N. Goodding.
110 (5) C. confertiflora; 78 (15) C. vir-
ginensis; d (46) C. flavoculata.
Kirkwood, J.
1808 (28) ^ Sheldonii.
Kirtley, C. L
— (8) C. salmonensis.
Knowlton, F. H.
$5 (20) C. thyrsiflora.
Kunze, E.
— (12) C. setosissima.
Langille, H. D.
117 Es C. flava; 113 (35) C. brevi-

erar Wm
1030 (27) C. o
Lechiner, H. J.
3 (7) C. leucophaea.
Leckenby, A. B.
— (7) C. leucophaea.
Leiberg, J. B.
5545 (1a) C. Jamesii var. multicaulis;
18

27) C. celosioides;
2049, 2223 (34) C. propria.

Lemmon, J. G.
165 (18) C. humilis.

Lemmon, á G. & wife.
— (1а) C. Jamesii var. multicaulis; —
(12) C. setosissima.

179 (22a) С. sericea var. typica.
Lunell, J.
— (31) C. Bradburiana.

342

Lyall, David.
— (7) С. =

Macbride, J
93, 875 d C. spiculifera; 2 (32b) C.
nana var. Shantzii

Macbride, J. F. & E. B. Payson.

C. Jamesii var. cinerea; 705

(le) C. Jamesii var. typica; 165 (12)
C. setosissima; 169 (28) C. Sheldonii;
60 (31) C. Bradburiana.

Macoun, John.
5801, 11819, 11820, 12800, 17087,
17089, 76741, 78500, 85007, — (29)
C. Macounii; 5802 (31) C. Bradburi-

ana.
Mearns, E. A.

111, 859, 360, 2091,

Jamesii var. multicauli
Merrill, E. D. & E. N. Wileox.

486, 497, 726 (6) C. flava; 617 (19) C

caespitosa; 457 (46) C. flavoculata.
Metcalfe, O. B.

70, 431, 863, 1061 (1a) C. Jamesii var.

multicaulis; 1576 (4) C. oblata.
т M. E.

(29) сі Macounii.

Moore, Е А

— (31) Hi Bradburiana.
Moyer, L. R.

458, — wi C. Bradburiana.
Mulford, A

— (32b) С. nana var. Shantzii.
Munz, P. A.

5721 (1f) С. Jamesii var. abortiva.
Munz, P. A., I. M. Johnston & R. D.

2522 (la) C.

arwood.
4209 (16) C. tumulosa.
Nealley, G. C
— (la) C. Jamesii var. multicaulis;
167 (4) С. oblata.
Nelson, А.
477, 2567, 2882, 8305, 9379, — (1e) С.
Jamesii var. typica; 22, 3074, 3098,

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

4771, 7067, 10693, — (6) C. flava
231, 1967, 1937, 7338 (13) C. virgula;
3072, 3120, 4671, 4749, 2772, 7078,
7255, 10696 (19) C. caespitosa; €
4715 (22b) C. sericea var.
418, 1362, 1990, 7306, 8263 Q0 с
thyrsiflora; 255, 888, 1956, 4788, 4837,
69 ‚ 8 45, 9356, 9411,
9045, 10700, 10741, — (31) C. Brad-
buriana; 389, 2876, 8309, 9367, —
(33) C. cana; 1511, 4518, 4572, 4616,
4665, 4749a, 4750, 4815, 7090, 9028,
(46) C. flavoculata

Nelson, A. & J. Е. Macbride.
1960, 1980, — (18) C. humilis; 1799
(26) C. spiculifera.

Nelson, A. & E. Nelson.
5429 (28) C. Sheldonit.

6350 (1b) C. Jamesii var. (аха.
Nelson, E.
4906 (1e) C. Jamesii var. typica; 4397
(6) C. flava; 4338, 4497 (19) C. caes-
pitosa; 4337 (46) C. flavoculata.
Nicollet, Г. М.
374 (31) С. Bradburiana.
Nuttall, Т.
-- (6) C. flava; — (19) С. caespitosa;
— (20) C. thyrsiflora; — (33) C. cana;
— (46) C. flavoculata.
Osterhout, G. E.
628, 2087 (1c) C. Jamesii var. cinerea;
627, 629, 3992, 4979, 5754, — (1e) С.
Jamesii var. typica; 6389, 6406 (6) C
flava; 6195 (9) C. stricta; 2463, — (13)
C. virgata; 6248 (19) C. caespitosa; —
4979, 5934 (20) C. thyrsiflora; 5996
(21) C. elata; 2122, 3476, 4940, 5960,
6094, — (22a) C. sericea var. typica;
5927 (31) C. Bradburiana; 5902 (33) C.
cana; 5980, 6010, 6200 (32d) C. nana
var. typica; 6416 (35) C. breviflora; 6003
(36) C. fulvocanescens; 4463, 4497,
5975 (40) C. longiflora; 6138 (42) C.
Osterhoutii; 2610 (44) C. Bakeri; 2750,
4696, 5956, 6006, 6194, 6197, 6252 (46)

W.H.
5188, 6185, — (31) C. Bradburiana.

1927]

PAYSON—SECTION OREOCARYA OF CRYPTANTHA

Overholts, L. O.
10, 155 (13) C. virgata.

Palmer, Ed.
591 (1a) C. Jamesii var. multicaulis;
375 (1c) C. Jamesii var. cinerea; 895
(3) C. Palmeri; —
(12) C. setosissima; 189 (26) C. spicu-
lifera; 598 (32b) C. nana var. Shanizit.

Palmer, Wm.
— (29) C. Macounii.

Parish, 8. B.
1480, 3238, 3694 (1f) C. Jamesii var.
abortiva; 1316, 1319, 3240, 4887, 19167
(5) C. confertiflora; 10243 (16) C.

tumulosa.

Parish, S. В. & W. Е. Parish.
1316 (5) C. confertiflora.

Parry, C. C.
166 (5) C. confertiflora; 173 (15) C.
virginensis; 49 (46) C. flavoculata.

Patterson, H.
110 (13) C. virgata;—(22a) С. sericea
var. typica.

Payson, E. B.
1019 (1c) C. James var. cinerea;
1027 (le) C. James var. typica;
405 (6) C. flava; 1014 (20) C. thyrsi-
flora; 271 (36) C. fulvocanescens;
85, 659 (40) C. longiflora; 458 (43) C.
ратадота; 1079, 1135 (44) C. Bakeri;
d 337, 670, 997, 1046 (46) C. flavo-

Ns Е. B. & G. M. Armstrong.
3223 (19) C. caespitosa; 0а 8224
(22b) C. sericea var. perennis.

Payson, E. B. & E. Bethel.

1608 (1e) C. Jamesii var. typica.

Payson же L В.

2533, ' 2557, 4307 (6) C. flava; 1880 (8)

C. salmonensis; 4248, 4253, 4291 (13)

C. virgata; 2578 (19) C. caespitosa;

4246, 4685 (20) C. thyrsiflora; 2528

(22a) C. sericea var. typica; 2580 (22b)

C. sericea var. perennis; 1768 (26) C.

spiculifera; 1746 (29) C. Macounii;

4247 (31) C. рав 3876 (36)

С. pee 4223 (43) C. para-

doza; 883. C. Bakeri; 1689, 2588,

3865, 3874, nag (46) С. flavoculata.

343

Pearson, G
210 (1a) Ag J amesii var. multicaulis.

LM.
10020 (27) C. celosioides.
Pierson, F. E.
3103 (14) C. Јатезії var. abortiva.
Piper, C. V.
2987, — C. leucophaea; 2294 (28)
C. Sheldoni
Porter, T. o.
ma C. sericea var. perennis.
Pratt, A
-- (31) E Bradburiana.
Pringle, C. G.
776 (1b) C. Jamesii var. lara; — (1а)
C. Jamesii var. multicaulis.
Purpus, C. A.
6068, 7067, 8048a, 8195 (1c) C. Jamestt
var. cinerea; ipi 5802 (5) C. con-
poras 8048 (12) C. setosissima; —
15) C. virginensis; 6070 (32e) C. nana
var. wina; — С. папа var.
typica; — (41) C. tenuis; 5869 (46) С.
flavoculata.

maley, F.
A. 107, 1078 (13) C. virgata; 1448 (20)
C. thyrsiflora.
Reverchon, J.
2120 (3) C. Palmeri.
Rothrock, J
700 (20) C. оға
Rusby, Н. Н.
283, 749 (1a) С. Jamesii var. multi-
caulis; 748 (12) C. setosissima.
Rydberg, P. A
254, 1514 (1e) C. Jamesii var. typica;
255 (20) C. thyrsiflora; 893 (31) C.
aire 256 (33) C. cana.
Rydbe * . А. Bessey.
js y (31) C. Bradburiana.
Rydberg, P. A. & E. C. Carlton.
7041 (1c) C. Jamesii var. cinerea.
552% P. А. & A. О. Garrett.
9 (2) С. желді b 8481 (6) С.
wine 9130 (44) С. В
Rydberg, P. A. & Е. к Pu
5702 (1c) C. Jamesii var. cinerea; 5704
(20) C. thyrsiflora.

344

Sandberg, J. H.
— (28) C. Sheldonii.
Sandberg, J. H. & J. B. Leiberg
373 (7) C. leucophaea; 164,—(26) C.
spiculifera; 440 (27) C. celosioides.
Saunders, A. A
— (29) C. Macounii.
Saunders, D. A.
4875 (26) C. spiculifera.
Scheuber, E. W.
— (31) C. Bradburiana.
Schmoll, H. M.
1281, 1346 (44) C. Bakeri.
Schoper, M.
28 (32b) C. nana var. Shantzii.
Scribner, G. L.
174 (31) C. Bradburiana.
Shantz, H. L
671 (1e) C. Jamesii var. typica.
Sharp, 8. 8.
182 (31) С. Bradburiana.
Shear, С. L.
4381 (22b) C. sericea var. perennis.
Sheldon, C. 8.
527 (1c) C. Jamesii var. cinerea; 216
(13) C. virgata; 526 (20) C. thyrsiflora.
Sheldon, E. P.
8815 (28) C. Sheldonii.
Shockley, W. H.
— (8) C. confertiflora; 244 (46) C.
flavoculata.
Slaughter, I. W.
7 (32b) C. nana var. Shantzii.
Smith, C. P.
7680 (1c) C. Jamesii var. cinerea;
3912 (1e) C. Jamesii var. typica;
3916 (20) C. thyrsiflora; 1573, 1605
(32b) C. nana var. Shantzii; 3936 (44)
C. Bakeri.
Sonne, C. F
— (18) C. humilis.
Soth, Mrs. M. E.
P-9 pe C. nana var. Shantzii.
Standley,
4899, E 7830, 7967 (1a) C. Jamesii
var. multicaulis; — (4) C. oblata; 6036,
6056 (20) C. thyrsiflora; 15107, 17428,
17594 (28) C. Sheldonii.

\

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Starz, E.
— (29) C. Macounii.

Stevens, G. W.
ние 3024, 3039 (1а) C. Jamesii var.
multicaulis
UE G.
— (6) C. flava; — (17) C. humilis; —
(32a) C. nana var. commixta; — (32 уд
C na var. Shantzii; — (39) C
Wetherillii; — (46) С. flavoculata.
Suksdorf, W. N.
407 (7) C. leucophaea; 888 (27) C.
celosioides.
Taylor, W. P. & H. C. Bryant.
— (10) C. nubigena.
Thompson, C. H.
97 (1e) C. Jamesii var. typica.
Thurber, Geo.
278, 293 (1a) C. Jamesii var. multi-
caulis; 147 (4) C. oblata.
— Ivar
1 (6) C. pe 1293, 9610 (46) C.
4.
ошпеу, J. W.
"n (12) C. setosissima.
Trac :
7835 (1a) С: —€— эд multicaulis.
Tracy, SS M. &F. S. E
424 (3) C. Palmeri.

ue, G. H.
$65 (18) C. humilis.

Turesson, G. W.
— (28) C. Sheldonii.

Tweedy, Frank.
576 (la) C. Jamesii var. multicaulis;
5219 (1e) C. Jamesii var. typica; 4262
(6) C. flava; 5222, 5223, 5676 (13) C.
virgata; 4260 (19) C. caespitosa; 3570
(29) С. Macounii; 816, 3569, 4261, —
(31) C. Bradburiana; 4259 (46) C.

149 (29) C. Macounü.
Vasey, G. R.
— (4) C. oblata; — (20) C. thyrsiflora.
Visher

295, 481, 587 (31) С. Bradburiana.
von бы Ж,

— (31) С. Bradburiana.

1927]

PAYSON——SECTION OREOCARYA OF CRYPTANTHA

15, 108 (6) C. flava.
Waldron, C. B.
— (31) C. Bradburiana.
Walker, E. P
85, 156 (36) C. fulvocanescens; 91
(43) C. paradoxa.

Ward, L. F.
393, 557 (1d) С. Jamesii var. disticha;

108, — бў я confertiflora; — (6) С
AE 12) C. setosissima; 49 (17)
Mu gp (46) C. flavoculata.

"usd M. G.
15 (36) C. fulvocanescens.
Watson, 8.
853 (18) C. humilis; 852 e: ©;
sericea var. perennis; 287 (28) C
Sheldonii; 852 (A6) C. pelos.
Webber, H. J.
— (31) C. Bradburiana.
White, O. E.
— (31) C. Bradburiana.
Whited, Kirk
1099, — (27) C. celosioides.

345

Wilcox, E.
79 (29) М Мой

Wilkes, Charles.
(61) 437 (28) C. Sheldonii.

Williams, R. S
109 (28) C. Shéldandt.

Williams, T. A
— (1a) C. Jamesii var. typica; — (20
C. thyrsiflora; — (31) C. Bradburiana;
— (33) C. cana.

Willits, Vie.
92 (29) C. Macounii; 236 (31) C.
Bradburiana.

Wolf, John, & Rothrock, J. T.
700 (20) C. thyrsiflora.

Wooton, E. O

1a) C. Jamesii var.

2)

C. setosissima; — (29) C. Macounii.
Wright, C.

E @) C. Palmeri; 1566 (4) C. oblata.
Zuck

— bes C. fulvocanescens.

INDEX TO SPECIES AND VARIETIES

Synonyms are printed in #ас8; species or genera maintained in this paper
are printed in Roman type; new species, varieties, or combinations are indicated

in bold face type.

Page
o er са

нр ors Eie 307
e { ANREDE ЛАН, neret ua Ad 295
CT. COEM Lr: 331
he Gh Sle ED 307
ПЕ O E 318
саев Фояа................... 281
газ A A 316
О В... 299
ihe r0 E 207

anaa araale. O
еп... 821
elatag. ема 285
UAVS Ж. Peo he Е e 259
flavoculata . ая 334
fulvocanescens . И WART. 319
етте Аба d er ОСЕ 307

Page

Cms ЕЕ. ак, 270
АИТ БЫН ЗЕН 278
N о ЕЕЕ ТИ 278
Eei on i Ои. rere e 296
HDTV. LC Р: NC 242
ао РИО ИР 250
VEE. o o a EO 246
ТЕГЕП Уг... 248
ОАЕ ҒИ ао к.» 246
ань пои ОО 244
Col polo MEC АЛАН UNES 248

а E MELLE 323
ООО... 262
ОН 326
МАСИН т.н... 303
тере ИЕ c d gs ii dae oo ВА 333
а. H

346
Page
о e soa IP AES EN 312
Var. GOmUIDIXÜA codes 312
ы; Жы ЖОЛГОЛОТ 314
var. Shantzii............... 313
var. бурісе.,................ 315
MUBIQONA: ......... ets 265
BIBUA o Е 254
Osterhoutii.................. 329
P eri. . 253
релдова.................... 330
от ии 317
рвегосагуа.................... 270
pustulosa................... 252
NEN. ISO aT 295
salmonensis................. 263
re ТЕРГ 286
var. perennis 288
var ЕС 28
setosissima.................. 268
йһедопн.................... 301
sobolifera................/.. 305
spiculifera................... 298
э RO OTE 264
PERDAN аж 242
E ауар т 327
ботата 225. 283
— Vel Dor T РАР ОН ea 276
4, РРР ЕРИ 270
а 270
a ha n O orana reia 274
Wetherillii 324
Cynoglossum
діотеғаійт................... 307
Eritrichium
шітосатевсепв................ 319
ПОЕТО. о 303, 307
var. fulvocanescens........... 278
var. изрийзяатиит.......... 283
var. һитїе.............. 278, 287
var. virgatum Spans 270
О РТУ е 248
Ре ГО Cans 262
ПИН о cU ave 244
зе{боаїзїтитї.............,.... 268
CHOCO РУР ТЕА 270
Krynitzkia
ПВО ТТТ” 277
echinoides . 321

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Page
fulsocanescena. oes enn 319
VOL. ЗОО: So ee 317
ук SET ee errr fee 307
var. 324
УЫТ, УГОН. (сео. 274
с 12 x resa ee ex e 248
МОЛ ОВО «eser ice rre t 262
орала 256
але 333
dci Love cue TERT SIRE is
с 2’, ПАРА 250
ТАТ, 86086» 2,25%... 60-2... 244
; rS PT E 254
г О PE nade asan 253
РИМЕ +. Ea we ae 307
DEG: ca rovro nine уук 287, 316
әсісггевіта,..............“.... 26
PEO. sn biden gc казу ув ркы 270
Myosotis
HOMO. eee eee ea aes 307
LOUCODNOED eios os гй дуз»
в! тиесова................ 242, 248
Огеосатуа
ОА СЕЧЕ 250
Encl АКРЕТ 307
afinis PENE 30k Ke seks en 288
NECI OS VIR сақы» 256
DC. o cvs vip S SOIRS Үз 295
DIRIGE Lei corr rc I CCS 287
1 os 5 eas Shaw enna DU 331
ВО с ives eo kh rr rin 318
савериова.,..>5.......55>%%... 281
о Е 316
се[ооїйев................. 299, 301
сііш-Мтяша................... 298
с CE AEE EEA ET 246
os ERROR LE EE ee ens 312
сочуевгайоға................... 256
НИН 22222. 334
Grane. eee UNES 277
ү га cae EE E 248
BL eee И ee eee T үн 313
о 283
и О О 334
schinoides ен 278, 321
HODE. 331

Е ОТТАР te
у

1927]

PAYSON—SECTION OREOCARYA OF CRYPTANTHA 347
Page Page
о. ЖАРОВА 334 PORNO Tes аралы сау, 253
ЕКЫ РИ 334 г SE СРОКА 330
Лаефотекене................. 319 СВОТ. a RETE es 255
ou ОНР 301, 307 SUE. и СИИ 288, 307
о од ОМИРИ TETTE EE 330 рыла ПРЕ, Зое 287
OECD OPE PIE PRIN SERIE Ыы 278 UD. Loco do PERREXIT 317
мәракента............... 254, 283 Рим. . Еа; 252
КРЕ. 522222 270 РЕДИ: ао ота REL IRA 295
SL ee ee ЗА 326 м тоттан. os a vs oa Е 263
sos ЫЕ Se ee ae 278 ONG. i nce ere es 287, 301
200 ИИ T И 273 р . N Е ес 268
s ЧОРООН 296 е corr co ПОТЕРИ 313
^^ ^^ ic Gi 5 a ss skew ONES 323 О т 301
БЕРМЕУ, IU EA 246 Р-Р oo. icc as c RECS 334
20 РРР 262 ПОЮ ТК 270
var. соп/етЕЙоға............. 256 о АКВА 298
о T A RE 326 or s ИИ D UM 264
0 ава 256 РЕК, с. 248
s LIT ATESTERTERETECEROITT S 259 ПШ НИНИ... СК 250
о СОАТ 278 Р РЕ 246
| s О у;...............+ 303 var, mullicaule............. 244
КЕ vous EST 333 ED A O RES 327
ot 6.0... 283 с ТЕР АР ТТТ сана 283
O LL E 244 о Sis «sak cs wae on 276
U Eha D 246 Qo LL A Ls EP PEE 283
Se 246 БИКЕ ERE Eae 270
БЕРЕСІ ....... 312, 315 forma spicata... 270
D АОН 319 БЕРЕКЕГЕ, 274
о. „РАСА 265 2s o АВР 324

‚о SLICE C 255 | Rochelia
^^ АРЕН 329 ААВ ЗОРУ 307

348 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
Figures illustrating nutlets of Cryptantha, section Oreocarya. Each one is illus-
trated by three figures—a dorsal, lateral, and ventral view. All are drawn to scale,
times 10, except fig. 4 of pl. 25 (complete fruit), which is times 71%

PLATE 25

—4. С. Jamesii var. multicaulis. Drawn from Wooton 401. Тһе nutlets
of all the varieties of C. Jamesii, C тек and С. Palmeri are very similar, апа
these figures will serve to illustrate them

Figs. 5-7 .oblata. Drawn from Jones 3759.

Figs. 8-10. С. confertiflora. Drawn from Goodding 814.

Figs. 11-13. С. flava. Drawn from Goodding 164.

Figs. 14-16. С. leucophaea. Drawn from Sandberg & Leiberg 373.

Figs. 17-19. С. salmonensis. Drawn from Payson & Payson 1880.

Figs. 1

[Vor. 14, 1927]

ANN. Мо. Вот. GARD., VoL. 14, 1927 PLATE 25

PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA

COCKAYNE, BOSTON

[Vor. 14, 1927]
350 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 26

Figs. 20-22. С. stricta. Drawn from Osterhout 6195.

Figs. 23-25. C. nubigena. Drawn from specimen collected by Cusick, Pine
Creek, Oregon.

Figs. 26-28. С. Clemensae. Drawn from type, Clemens, Glenn’s Pass, California.

Figs. 29-31. С. setosissima. Drawn from specimen collected by М. Е. Jones,
Flagstaff, Arizona, August 4,

Figs. 32-34. С. virgata. ‘Drawn from specimens collected near Laramie, Wyo-
ming, by the author in 1925.

Figs. 35-37. С. insolita. Drawn from specimen collected by К. Brandegee,
Las Vegas, Nevada.

ANN. Mo. Bor. GARD., VoL. 14, 1927

PAYSON—SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

PLATE 26

[Vor. 14, 1927
352 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 27
Figs. 38-40. С. virginensis. Drawn from M. E. Jones 5195 a.
Figs. 41-43. C. tumulosa. Drawn from type, T. S. Brandegee, Providence
Mountains, California.
1 C. modesta. Drawn from type, Jones 6692, Aurum, Nevada.

Figs. 47-49. C. humilis. Drawn from specimen collected by Sonne, along the
Truckee River, at Verdi, California.

Figs. 50-52. С. caespitosa. Drawn from Payson & Payson 2578.
Figs. 53-55. C. thyrsiflora. Drawn from Osterhout 4979.

ANN. Мо. Bor. GARD., VoL. 14, 1927 PLATE 27

mm - 3

wat fae

---

$
е اج‎

ОЕ СКҮРТАМТНА

PAYSON—SECTION OREOCARYA

COCKAYNE, BOSTON

[Vor. 14, 1927]
354 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 28

Figs. 56-58. C. elata. Drawn from Osterhout 5996.

Figs. 59-61. С. sericea var. typica. Drawn from Payson & Payson 2528.

Figs. 62-64. C. aperta. Drawn from type, Eastwood, Grand Junction, Colorado.
Figs. 65-67. С. rugulosa. Drawn from type, М. Е. Jones, Fish Springs, Utah.
Figs. 68-70. С. interrupta. Drawn from Heller 9185.

Figs. 71-73. С. spiculifera. Drawn from type, Sandberg & Leiberg 164.

Figs. 74-76. С. celosioides. Drawn from Suksdorf 888.

Figs. 77-79. С. Sheldonii. Drawn from type, Sheldon 8315.

com o ga ARES S

ANN. Mo. Вот. GARD., VoL. 14, 1927

PAYSON—SECTION OREOCARYA

COCKAYNE, BOSTON

ج

ОЕ СКҮРТАХТНА

PLATE 28

[Vor. 14, 1927]
356 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION оғ PLATE
PLATE 29

Figs. 80-82. C. Macounii. Drawn from specimen collected by Macoun, at
Moose Jaw, Saskatchewan.

Figs. 83-85. С. sobolifera. Drawn from type, M. E. Jones, Upper Marias
Pass, Montana

Figs. 86-88. 0 Bradburiana.

Figs. 89-91. С. nana var. Shantzii. Drawn from type of Oreocarya dolosa, С. P.
Smith 1605. Тһе nutlets of C. breviflora, C. fulvocanescens, and C. echinoides are
2” to those of C. nana var. Shantzii. The differences are noted in the descrip-

E 92-94. С. сапа. Drawn from type, A. Nelson 8309.
5-97. C. propria. Drawn from Leiberg 2223.
Figs. 98-100. C. Jonesiana. Drawn from type, Jones, San Rafael Swell, Utah.

ANN. Mo. Вот. GARD., VoL. 14, 1927

£
к
м.
>
б»)
8

22
Уч

A 77.

PAYSON—SECTION OREOCARYA ОЕ CRYPTANTHA

COCKAYNE, BOSTON

PLATE 29

358

[Vor. 14, 1927]
ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 30

. 101-103. С. Wetherillit. Drawn from type, Eastwood, Court House Wash,

. 104-106. С. longiflora. Drawn from Payson 659.

. 107-109. С. tenuis. Drawn from specimen collected by Purpus.
. 110-112. С. Osterhoutii. Drawn from type, Osterhout 6138.

. 113-115. С. paradoza. Drawn from Payson & Payson 4223.

. 116-118. С. Bakeri. Drawn from Payson & Payson 3834.

‚ 119-121. С. mensana. Drawn from type, Jones 5445

. 122-124. С. flavoculata. Drawn from Payson & Payson 2583,

-——— ———— — GÀ 7

жаны айы... каза шалы CEP

ANN. Мо. Bor. GARD., VoL. 14, 1927

#

PAYSON—SECTION OREOCARYA OF СКҮРТАХТНА

COCKAYNE, BOSTON

^
&i

ғ”
a
у ср
B be

PLATE 30

РЛ ce `

Annals

of the
Missouri Botanical Garden

Vol. 14 NOVEMBER, 1927 No. 4

COTTON WILT: A PATHOLOGICAL AND
PHYSIOLOGICAL INVESTIGATION!
DAVID C. NEAL
Plant Pathologist, Mississippi Agricultural Experiment Station
Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany of
Washington University,
and Special Research Fellow of the National Fertilizer Association, 1925-27

INTRODUCTION

Cotton wilt is undoubtedly the most important disease of the
cotton crop in the United States and causes the largest annual
loss. According to records of the Plant Disease Survey of the
U. S. Department of Agriculture compiled for several years, the
disease is widely distributed through the cotton belt from Virginia
to Texas, and is apparently spreading from year to year to new
localities.

'The damage caused by this important disease of cotton varies
in different localities each season, but frequently large areas are
involved and, without question, the annual loss to the entire
cotton belt must reach into the millions of dollars. Тһе esti-
mated reduction in yield in the United States because of wilt was
350,000 bales in 1925. In Mississippi, one of the leading cotton-

. producing states of the mid-south, the annual loss from this

disease is fully 50,000 bales, and it reaches even larger proportions
during unusually dry seasons.

In view of the wide prevalence of wilt in Mississippi and other
southern states, an investigation of this problem was begun by
the writer at the Missouri Botanical Garden in the fall of 1925,
where а year was spent in а laboratory and greenhouse investi-

! An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University and sub-
mitted as a thesis in partial fulfillment of the requirements for the degree of doctor
of philosophy in the Henry Shaw School of Botany of Washington University.

ANN. Mo. Bor. Garp., Vor. 14, 1927 (359)

[Vor. 14
360 ANNALS OF THE MISSOURI BOTANICAL GARDEN

gation of this disease and of the causal organism. The study of
the problem has been continued since this time at the Mississippi
Agricultural Experiment Station. Briefly, the objects of the in-
vestigations reported in this paper are: (1) to ascertain some of
the physiological relations of the fungus causing the disease, such
as growth-temperature relations in pure culture, effects of H-ion
concentration upon growth, and the factors involved in producing
wilt by the fungus; (2) to determine the pathogenicity of the
fungus; (3) to determine the effects of varying conditions of
nutrition upon the development of the disease under greenhouse
and field conditions; (4) to study differences in varietal suscepti-
bility and resistance in both upland and staple cottons.

PREVIOUS INVESTIGATIONS ON COTTON WILT

One of the first investigators to establish the identity and
probable relationship of the fungus associated with cotton wilt
was Atkinson (92). He considered the wilting of the plant to
be due to the plugging of the vessels of the plant near the base of
the stem by the mycelium of a fungus. He made isolations from
the tissues of affected plants and described the fungus found as a
new species under the name Fusarium vasinfectum Atk. Atkinson
also found a species of Fusarium on the surface of some of the
infected plants, but considered this a saprophyte, distinct from
the internal fungus. Some years later, Smith (799) made a
more extensive study of the Fusaria found associated with wilt
in cotton, in cowpea, and in watermelon. This investigator was
under the impression that he had developed the ascigerous stage
of the fungus on cotton, and this he designated Neocosmospora
vasinfecta (Atk.) Smith. He stated that the fungus probably
also occurred on okra, but that its parasitism and the genetic
connection of the various spore forms were not proved and that
chlamydospores were not observed. At the time these investiga-
tions were made, Smith seems to have had no doubt as to the
parasitic nature of the cotton or cowpea wilt fungus, or as to the
genetic relationship of the various spore forms found on cotton.
He admitted that these points had not been definitely settled
through satisfactory infection experiments, by deriving one spore
form from the other in pure cultures. Тһе field evidence, how-

EE eR Ups

1927]
NEAL—COTTON WILT 361

ever, of the parasitic nature of the cotton and cowpea fungus was
thought by Smith to be most convincing, since the fungus was
always present in the vessels of the diseased plants and the
disease occurred year after year on the same soils. More recent
studies by Higgins (09) and Butler (710) indicate that in all
probability there is no genetic relationship between the Neocos-
mospora referred to by Smith and the internal fungus Fusarium
vasinfectum which Atkinson associated with cotton wilt. Neither
did Higgins regard the Neocosmospora as having any connection
with the wilt of cowpea. In fact, a study of the literature in-
dicates that with cultures of Fusarium vasinfectum Atk., isolated
from wilted plants and accordingly capable of infecting cotton,
perithecia have never been produced. It appears, therefore,
that the correct identity of the fungus associated with the wilt
disease of cotton should properly date back to Atkinson’s work
in Alabama in 1892.

Orton (’00) apparently accepted Smith’s determination for
cotton wilt as Neocosmospora vasinfecta and was under the im-
pression that the wilt of okra was caused by the same fungus.
He states: “Хо inoculation experiments have been tried, but in
the experience of the writer, okra has never failed to contract
the disease when planted in fields infected with the cotton wilt
disease."

Orton (700) studied the period of incubation, or the interval
elapsing between the time the young seedlings of cotton were
exposed to the attacks of the fungus and the first visible external
symptoms. He found that a time interval of forty days or
longer was required. Не noted the conditions which favored the
progress of the fungus through the plant and although these were
not fully understood, he states: ‘‘From observations that have
been made, it is believed that highly fertilized plants, growing
vigorously, suceumb more readily than those which have grown
on poorer soil." Orton also corroborated the work of Smith (99)
and Atkinson (792) by producing the disease in cotton plants
through infection experiments. 'This was accomplished by
inoculating the soil, in which cotton was growing, with pure
cultures of Fusarium vasinfectum. Не conducted experiments
with a number of fungicides, spraying these on the plants and

[Vor. 14
362 ANNALS OF THE MISSOURI BOTANICAL GARDEN

applying them to the soil in the hope of killing the fungus, but
none of the results obtained gave promise of combating the
disease successfully. Тһе fungicides used were Bordeaux mix-
ture, Bordeaux mixture and sulphur, copper carbonate, copper
acetate, lime, sulphur, lime-sulphur mixture, liver of sulphur, iron
sulphate, carbolic acid, caustic soda, formalin, and kainit. Fields
were selected which were uniformly infested, but the cotton died
in both the treated and untreated areas, and no difference trace-
able to the fungicides used could be observed.

At that time, Orton advocated such hygienic treatment as
rotation of crops and the removal of diseased plants, also the
avoidance of spread of the fungus by cattle, tools, or through
infested manure. Considerable stress was also placed upon devel-
oping wilt-resistant selections of cotton. Fulton (707), working
in Louisiana in 1907, advocated the planting of wilt-resistant
varieties of cotton and the employment of crop rotation as a
method of control.

In 1908, Orton published a popular account of cotton wilt in
which the geographical distribution of the disease, amount of loss,
symptoms of wilt, and the life history of the fungus, so far as
known at the time, were given. The influence of seasonal con-
ditions, the time of planting, soil conditions, and the effect of
fertilizers and fungicides upon wilt were noted. Attention was
also given to breeding cotton for wilt-resistance, and breeding
methods were described.

Lewis (711) noted the distribution of wilt in Georgia in 1911 and
estimated the annual loss in that state to be about three-quarters
of a million dollars. He studied the effect of fertilizers on the
disease and concluded that various combinations of acid phos-
phate, muriate of potash, and guano,' used at the rate of from
300 to 800 pounds per acre, were of no benefit in controlling it.
Тһе test plats were not replicated, however, and the size of the
plats in the experiments was not indicated.

Lewis (715) originated several wilt-resistant varieties of cotton
from 1905 to 1915, some of which are still widely planted in the
southeastern section of the cotton belt. He used both the
method of selection and of hybridization of desirable parent

1 Mixed fertilizer, 10-2-2.

1927]
NEAL—COTTON WILT 363

plants to obtain resistant strains. Doubtless his better-known
wilt-resistant cottons are Lewis 63, Dix-afifi, and Council Toole.

Rast (’22) conducted some fertilizer experiments in Arkansas
in 1920 on the control of cotton wilt, which apparently were
beneficial. A portion of a field was observed by him in 1920 in
both fertilized and unfertilized areas heavily infested with wilt.
The grower had used 500 pounds per acre of a mixture containing
10 per cent phosphoric acid, 3 per cent nitrogen, and no potash.
So many plants in this field on both the fertilized and unfertilized
areas died in 1920 that no record was kept of the yield of cotton.
The experiment had been planned to run on this area for 5 years,
and it was necessary to change radically the plan of the experi-
ment in order to justify the continuance of the work. It was
suggested that 500 pounds of the same fertilizer that was used in
1920 (containing 10 per cent phosphoric acid, 3 per cent nitrogen,
and no potash) be mixed with 500 pounds of kainit containing
12.5 per cent potash. This mixture contained, as shown by
analysis, 5 per cent available phosphoric acid, 1.5 per cent ni-
trogen, and 6.25 per cent potash. It was applied in the spring of
1921 at the rate of 1000 pounds per acre before planting. His
results are quoted: “Тһе plants on the unfertilized area began
to die long before they were mature and were evidently infected
with the wilt disease. By harvest time, no less than 95 per cent
of the plants on the unfertilized area were dead and not a dead
plant could be found in any part of the field where the fertilizer
was used. The fertilized area produced 1127 pounds of seed
cotton per acre; whereas the disease-infested part of the field to
which no fertilizer was applied produced only 225 pounds of seed
cotton per acre. Plants on an adjoining area to which 500
pounds per acre of fertilizer containing 10 per cent available
phosphoric acid, 3 per cent nitrogen, and no potash were applied
before planting and to which 500 pounds per acre of kainit were
used as an additional application after the plants were up and
growing, were equally resistant to the disease and just as prolific.”
Rast concluded that the above results were due to the potash in
the fertilizer, and Elliott (Rast, '22, р. 224) who examined the
plats, wrote as follows: ‘‘I do not attempt to make an explanation
of the facts, but to all appearances the treatment the kainit plat

[Уог. 14
364 ANNALS OF THE MISSOURI BOTANICAL GARDEN

received enabled the plants this year to very largely escape wilt
infection."
DESCRIPTION оғ Соттом WILT

А noticeable yellowing of the leaves of the cotton plant ассот-
panied by a stunted appearance somewhat early in the season is
usually a good indication of Fusarium wilt. The disease may be
noted at first in irregular spots over the field, and each succeeding
year these infested areas enlarge. Frequently the main stem of
plants infected with the disease remains short, while some of the
lower branches grow normally (pl. 31, fig. 1). It is, however,
by means of the internal appearance of affected cotton plants
that we obtain the most characteristic symptom of wilt. If the
stem of a freshly wilted plant is cut across near the ground and
a brown or black discoloration of the fibro-vascular tissues is
found, there is strong evidence of the disease (pl. 31, fig. 2). Тһе
tap-root of infected plants is shorter and the lateral roots less
abundant. If plants are examined late in the season at the time
they are approaching maturity, the wilt fungus may frequently
be observed not only in the lateral and tap-roots, but in the
vascular tissues of the main branches, the leaf petioles, and
pedicels.

MORPHOLOGICAL AND CULTURAL CHARACTERS OF THE FUNGUS

The fungus, Fusarium vasinfectum Atk., the pathogen respon-
sible for the occurrence of cotton wilt, has not been made the
object of extensive morphological or taxonomie study by the
writer. It seems desirable, therefore, to accept the criteria of
taxonomie differentiation of species as established through the
work of Wollenweber and associates (25). Тһе fungus is ac-
cordingly placed in section Elegans of the genus Fusarium.
Growth of the fungus is usually rapid on a variety of complete
nutrient media, and both micro- and macro-conidia are produced
readily in culture. Місго-сопійіа are present on the aerial
mycelium and are dominantly non-septate, ovoid to fusoid,
4-6 x 3-4.5 u. Macro-conidia are usually none to 3-septate,
attenuate at distal end, pedicellate, fusoid to sickle-shaped,
20-25 x 4-6. — Chlamydospores are produced in cultures 3
weeks old or less and may be either terminal or intercalary (pl. 32,

1927]
NEAL—COTTON WILT 365

figs. 3-4). Тһе color of the stroma is marked, and on sterilized
rice the acid modification is especially well developed, the rose-red
to wine-red color appearing in this medium in from 8 to 12 days
after the fungus is introduced. The aromatic odor of the fungus
when grown on rice is pronounced. Оп nutrient and bean agar
the color of the mycelium is white, but on starchy media, such as
potato-dextrose and corn-meal agar, the mycelium is tinged
noticeably purple.
SOURCES OF CULTURES

Several strains of Fusarium vasinfectum have been used during
the progress of this investigation. On September 15, 1925, the
writer isolated the fungus from the stem tissues of а Cook Tri-
umph hybrid cotton plant growing in a breeding plot of the Exper-
iment Station, A. & M. College, Mississippi, and later established
the pathogenicity of this culture by a series of infection experi-
ments. This isolation has been used very largely in the earlier
phases of the investigation and particularly in the laboratory
studies of the fungus. Оп July 15, 1926, the fungus was isolated
from а wilted plant of Trice cotton in the breeding plot at the
Experiment Station, A. & M. College. It was also isolated from
а wilted plant of Delfos 6102 cotton collected at Dunleith, Missis-
віррі, on August 16, 1926, and from Ттісе cotton growing at
Poplarville, Mississippi, on August 12, 1926. Strains of the
fungus supplied by Dr. V. H. Young and Dr. H. R. Rosen, of the
Arkansas Experiment Station, have been included in some of the
experiments. The pathogenicity of all cultures was later estab-
lished by infection experiments.

The method of isolation may be briefly described as follows:
Cotton plants showing unmistakable symptoms of wilt were col-
lected and free-hand sections made of the stem tissue and occa-
sionally the leaf petioles. Тһе sections with typical vascular
discoloration were treated from 3 to 5 minutes in a 1 : 1000
solution of mercuric chloride (holding with sterile forceps),
rinsed in sterile distilled water, and placed on nutrient, bean,
and potato-dextrose agar in Petri dishes. As a disinfecting agent
in the preparation of material, calcium hypochlorite has also
been used extensively in making isolations, and is equally satis-
factory for the purpose.

(Уот. 14
366 ANNALS OF THE MISSOURI BOTANICAL GARDEN

PHYSIOLOGICAL STUDIES OF THE FUNGUS
INFLUENCE OF TEMPERATURE

Temperature relations of many organisms causing plant diseases
are important in that the geographical distribution of these
diseases may be influenced very largely by their thermal relations.
For instance, it is a well-known fact that many of the Fusarium
diseases develop in destructive form only during periods of pro-
longed hot weather when soil temperatures are high, whereas
others may require moderately cool temperatures to attain the
maximum degree of development necessary for infection.

A very large amount of work has been reported by various
investigators on temperature relationships of many fungi causing
plant diseases, and particularly is this true of the Fusarium group
of diseases. It is impossible to go extensively into the literature
on this subject in the scope of this paper, but the work of a few
investigators may be noted:

Gilman (716) found that Fusarium conglutinans Wollenw., the
fungus causing cabbage yellows, has a high optimum temperature
and is very resistant to drying both in pure culture and in the
soil. The occurrence and development of the characteristic
symptoms of the disease required a temperature of about 17-
22° С. or higher. Lower temperatures (12-16% C.) under con-
trolled conditions prevented the occurrence of the trouble in the
greenhouse. Field observations made over a period of years
bore out this relation between the occurrence of the disease and
high temperature.

Edgerton (720) has worked on the temperature relations of
tomato wilt, and he and other investigators state that the tomato
wilt fungus, Fusarium Lycopersici Sace., is more commonly found
in the warmer regions throughout the world. He finds that this
fungus grows best at a temperature around 29? C. and that infec-
tion also takes place quicker and the disease develops in plants
more rapidly if the soil is kept around that temperature. Very
little development of the disease is noted at lower temperatures.

Tisdale (723) found that cabbage yellows is often most de-
structive in Wisconsin during midsummer when the soil is dry
and hot. On the other hand, there is little or no development
of the disease, even on soils heavily infested with the organism,

1927]
NEAL—COTTON WILT 307

during moist, cool weather. This author emphasizes the impor-
tance of the temperature relation of this fungus in the geograph-
ical distribution of the disease. For instance, in the southern
states, where cabbage is generally grown commercially as a
winter or early spring crop, the soil temperature is very probably
too low for the organism to gain а foothold on such plants.
However, when once introduced, the Fusarium establishes itself
on summer-grown cabbage and other related hosts and attacks
the crop whenever soil temperatures are favorable.

Clayton (23) found somewhat similar temperature relations
with the tomato-wilt fungus as did Edgerton (720), but noted that
the most favorable temperatures of soil and air, as determined in
tanks in the greenhouse, are 27? C. and 28? C. respectively, i.e.,
after the fungus was established in the stems of the plants.

Jones and Tisdale (22) found that in the case of flax wilt
caused by Fusarium Lini Bolley, the disease was aggravated by
hot seasons and that the midsummer temperature in the regions
where it occurred corresponded closely with the optimum for the
fungus.

In view of the rather extensive investigations which have been
made on the Fusarium wilts of other crops, it was interesting to
study some of the temperature relations of the cotton-wilt fungus,
Fusarium vasinfectum. Moreover, it has been noted by the
writer not only in Mississippi, but in Alabama, Louisiana, Arkan-
sas, and Texas, that wilt of cotton is usually more prevalent
during prolonged periods of dry, hot weather. Particularly was
this true in the Gulf Coastal States during the years of 1924-25,
two seasons which were notable because of extended periods of
dry weather and abnormally high temperature during the greater
part of the summer.

In studying the effects of temperature upon the development of
the fungus in pure culture, uniform Petri dishes containing a
measured amount (10 сс.) of potato-dextrose agar were used. In
order to obtain spores of the fungus free of mycelium, a suspension
of the spores was made in test-tubes containing 3 or 4 cc. of
sterile distilled water. Тһе spore suspension was made from a
7-day-old culture of the fungus growing on steamed rice. In one
Petri dish, a thin layer of nutrient agar was poured and then

[Vor. 14
368 ANNALS OF THE MISSOURI BOTANICAL GARDEN

several loopfuls of the spore suspension were placed at random
over the surface of the plate. As soon as the agar solidified, the
plates were examined by the low power of the microscope, and
single spores were located. These areas were immediately
marked by means of a blue wax pencil and the agar blocked out
by means of a needle and transferred to the center of the Petri
dishes. Each temperature series was run in triplicate and
measurements of the growth of the colonies of the mycelium were
made at the end of 2, 6, 10, and 15 days. Тһе results are given in
table r.

TABLE I
GROW'TH OF FUSARIUM VASINFECTUM AT VARIOUS ТЕ MPERATURES
Diameter of colony in em. at indicated intervals
Temperature 2 days 6 days | 10 days 15 days
Plate No. | Plate No. | Plate №. | Plate No.
11231123 | 1 | 2|з|11|213
6 0 0 0 0.2/0.2/0.2/1.2 |1.5 1.2 |1.5 |1.6 | 1.6
10 0.4/0.3/0.2/1.8/1.9/1.8|2.5 12.8 |2.9 13.8 |3.6 13.4
15 1.9|1.811.8|8.8|3.7|3.915.4 |5.2 |5.4 18.2 8.0 | 8.1
20 3.1|3.2|3.1|7.0|7.2|7.0/9.0*/9.0*]9.0*|9.0*]9.0*
24 3.8|]3.7|4.0/7.8|7.5|8.0/9.0*/9.0*9.0* — | — | —
28 4.1|4.0|4.5|7.9|8.0/|8.2/9.0*/9.0*/9.0*9.0*]9.0* | f
30 3.8|3.7|4.0|7.7|7.5|8.0/9.0*/9.0*/9.0*]9.0*]9.0*| —
35 1.8|1.4|1.6|3.2|3.3|3.6 3.5 3.0 3.5 | — | — | —
40 0.010.010.010.010.(

* Growth over plate.
1 Contaminated.

It will be noted from the above results that Fusarium vasin-
fectum grows very slowly at the lower temperatures, and that the
optimum is not far from 28? C. Тһе optimum temperature,
therefore, of the parasites of the very similar Fusarium diseases of
cotton, tomato, Нах, and cabbage is practically the same in pure
ишке, although the host plants have quite different temperature
relations. This fact is supplemented by the work of Edgerton
(20), Tisdale (728), and others. Тһе maximum temperature of
Fusarium vasinfectum is apparently around 38? C

The effect of temperature on the growth of Fusarium vasin-
fectum was also studied by means of liquid cultures. The dry
weight of the fungous mat was ascertained at various tempera-

1927]
NEAL—COTTON WILT 369

tures ranging from 6 to 40? C. at intervals of 5, 10, and 15 days.
Duggar's! solution was used for growing the fungus and the final
concentration of chemicals used in this solution was as follows:
M/4 dextrose, M/b KNO;, M/20 KH;PO,, M/100 MgSO,, and
a trace of FePO,. For obtaining these dilutions the following
stock solutions were prepared: M/2 dextrose, МЛ KNO,,
M/4 КН,РО,, M/10 MgSO,, апа M/1000 ЕеРО.. For 50 сс. of
medium the following amounts of these solutions were used:
25 се. dextrose, 10 се. КМО», 10 се. КН,РО,, 5 сс. MgSO,, and
6 drops ЕеРО.. Pyrex flasks were used for the cultures, and
these were run in triplicate at each temperature. А copper-
lined, differential incubator with compartments was used in these
experiments. It was heavily insulated with asbestos, and had an
ice chest at one end and on the opposite end a compartment for
water was heated by a small gas burner regulated by a thermostat.
With such radiation almost any temperature could be maintained
by simply increasing or diminishing the size of individual com-
partments opposite the ice chest and water-heated unit. Тһе
flasks were inoculated with a 7-day-old culture of the fungus by
means of a sterile needle in the usual manner, and care was taken
to introduce approximately the same amount of the inoculum
into each flask. At the intervals indicated, the mats were col-
lected оп а filter paper by means of a Buchner filter, dried to
constant weight in а Fries ovcn, and weighed.

Using the dry weight of the fungous mat as an index, it will be

TABLE II
GROWTH OF FUSARIUM VASINFECTUM AT VARIOUS TEMPERATURES
Temperature Dry weight of fungous mat in milligrams
degrees C 5 days 10 days 15 days

6 6.20 54.9 81.4
10 28. 84.5 97.1
15 35. 98.7 145.2
20 38. 149.4 251.5
25 49.1 112.3 455.2
28 226. ( 338.6 610.0
30 82. 498.7 725.2
35 86. 294.2 422.9
38 172.1 247.3

ж Contaminated.

1 A nutrient solution for fungi recommended by Dr. B. M. Duggar.

[Vor. 14
370 ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN

noted again that the optimum temperature for Fusarium vasin-
fectum is around 28 and 30? С. It grows slowly at temperatures
below 10? C.; the growth curve begins to fall sharply at about
30° C., while at 38? C. it falls off abruptly. The results are
graphically illustrated in fig. 1.

3
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15 2 25 BS 35 3

Fig. 1. Showing the effect of temperature on the growth of Fusarium vasinfectum.

RELATION TO HYDROGEN-ION CONCENTRATION

The effect of the hydrogen- and the hydroxyl-ion concentration
upon the germination of the spores and growth of various fungi
has been demonstrated by numerous investigators. That the
reaction of the substratum is a very important factor in influenc-
ing the growth of many disease-producing organisms is а well-
known fact. The literature bearing upon this subject is rather
extensive, and has been reviewed to advantage by Webb (719,721),
Lehman (723), and others. Webb (721) finds that with a certain
Fusarium sp., germination is equally as good, if not better, with
an alkaline reaction. However, with the majority of fungi he

"TEES | [РУХУ АЕ | eee

1927]
NEAL—COTTON WILT 371

employed the maximum for germination was between P43.0 and

Hopkins (721), using a synthetic medium in which the hydrogen-
ion concentration was adjusted by means of КН,РО,, K;HPO,,
H;PO,, and KOH, found that a Fusarium isolated from scabby
wheat, but not proved to be Gibberella, exhibited a similar depres-
sion in the growth-acidity curve as did Gibberella saubinettii, the
wheat-scab organism. In the case of the latter, the amount of
growth increased with decreasing acidity from Рһ2.5 toamaximum
of Pa 4.0-4.5. It then decreased to a minimum at Рн 5.0-5.5 and
rose again to a second maximum, but the highest point was not
determined.

MacInnes (22) observed that a strain of Fusarium isolated
from scabby wheat in Minnesota was capable of growing in
solutions varying through an unusually wide range of hydrogen-
ion concentrations. Тһе organism grew in nutrient media rang-
ing from Py 5.0 to Py 11.7. Хо determinations were recorded
for hydrogen-ion concentrations on either side of these values.

Kirby (22) found that the maximum hydrogen-ion concen-
tration for Fusarium moniliforme on corn was near 8.2, and the
range of growth was greater than Py 5.2-9.2.

Тһе results of the above-mentioned investigations and many
others clearly indicate that many fungi require varying degrees
of acidity in the substratum for optimum growth, whereas others
flourish to better advantage in an alkaline medium or in one ap-
proaching the neutral point. In the case of the organism causing
the ‘‘take-all disease" of wheat, Kirby (722) states that the fungus
requires alkalinity for optimum growth, and that may explain
why additions of alkaline substances to the soil have been re-
ported as favoring the fungus, while additions of acid-forming
substances decrease the amount of infection. From the work
of numerous investigators, notably Martin (720, '21), Lint (714,
215,716), and others, it is known that there is a correlation between
increase in soil acidity and a decrease in the prevalence of the
potato-scab organism, Actinomyces scabies. According to Chupp
(25) and others, a neutral soil with Py value of 7 will almost
invariably give а crop 100 per cent scabby, while an acid one
with а Ри value of 5.6 or greater will produce an almost clean
crop.

[Vor. 14
372 ANNALS OF THE MISSOURI BOTANICAL GARDEN

In order to gain an idea of the limiting hydrogen-ion concen-
tration for the growth of Fusarium vasinfectum some growth tests
were made. Duggar’s solution previously referred to was used
and contained inorganie salts and dextrose in the following volume
molecular concentration: M/4 dextrose, M/5 KNO;, М/20
KH;PO,, and М/100 MgSO.. Six drops of M/1000 FePO, were
used for each 50 се. of solution, and M/5 Са(ХО:); was substi-
tuted for the М КМО.. The inorganic salts were Merck’s
“blue label” reagent grade, and the dextrose was the grade desig-
nated as “со standardized.” Тһе nutrients were prepared in
900-cc. Pyrex flasks in quantities of 475 сс. for each series, and
this quantity of solution was adjusted to the desired Py value
by the addition of previously determined amounts of 0.1647 N
H;PO, or 0.1613 N KOH. The range of initial Рн values was
from 2.5 to 9.0 After Py adjustments were made, the 400 сс.
of nutrient solution were divided as follows: 50 cc. of the solution
were pipetted into each of eight 125-cc. Pyrex flasks so as to have
each series in duplicate for examination after certain growth
periods. The remaining 75 сс. of solution of each initial Рн
value were put into Jena test-tubes and sterilized for 30 minutes
at 15 pounds pressure in order to note the effect of sterilization
upon P, and to serve as controls for final Py reading at the end of
designated growth periods. It was found that autoclaving did
not change the Рн appreciably in the tubes, but to guard against
enolization of sugars in the alkaline ranges, it was deemed best
to sterilize the dextrose portion separately for each flask. This
was added later to the solution of salts by means of a sterile
pipette.

The cultures were inoculated with a spore suspension of
Fusarium vasinfectum in sterile distilled water, one loopful being
added to each flask, and placed in the incubator at а temperature
of 29° C. Тһе cultures were also wrapped with carbon paper to
exclude the light from the bulbs. Two flask cultures from each
series were examined at the end of 8, 12, 16, and 28 days, and the
dry weight of mat, hydrogen-ion concentration, and titrable
acidity were determined at each examination. Тһе Рн values
were determined colorimetrically by the method of Clark and
Lubs (717).

1927]
NEAL—COTTON WILT 373

By means of a Buchner suction filter, the mats were collected
on filter-paper, the dry weight of which had been previously de-
termined. The mats were dried for 3 days in а Fries oven at
100? C., eooled in a calcium chloride desiccator for а uniform
period, and weighed. Тһе filtrate from each mat was diluted
to 100 сс. with distilled water and the total acidity was deter-
mined by titrating a 10-cc. aliquot of the filtrate with N/50 KOH,
using phenolphthalein as an indicator. Тһе amount of unused
sugar present at each interval was also determined for series 2,
7, and 12, and for these determinations the method of Shaffer
and Hartman (’21) wasemployed. The data obtained are shown
in table 111.

TABLE III
GROWTH OF FUSARIUM VASINFECTUM IN NUTRIENT SOLUTION*
Growth | Dry wt. Cc. N/50 KOH AN of min p
Series | Initial | period mat Final Рн | to neutralize . of so
No. Рн ays mgms. 10 cc. filtrate
8 9.0 2.9 19.45
1 2.5 12 121.3 2.9 18.07
16 387.4 3.0 19.00
28 348.4 6.6 11.2
8 49.6 3.2 19.9 43.2
2 2.8 12 145.0 3.4 17.6 24.0
16 385.0 3.4 16.5 2.24
28 397.0 6.2 11.9 0
8 51.2 3.5 16.3
3 3.0 12 301.0 5.5 13.4
16 414.0 5.8 13.1
28 756.3 6.4 5.4
8 112.2 4.6 16.6
4 3.5 12 343.5 o. 14.2
16 417.1 5.8 13.6
28 761.0 6.9 1.2
8 116.0 4.5 16.8
5 4.0 12 345.2 5.8 13.4
16 419.0 5.8 14.2
28 758.3 7.8 .8
8 122.3 5.3 14.9
6 4.5 12 357.2 6.6 10.7
16 418.2 6.9 3.9
28 747.0 7.4 1.9
8 133 5.3 15.2 28.08
7 5.5 12 362 6.6 9.8 4.8
16 413.0 6.8 у! 2.88
28 782 7.2 2.3 0

[Vor. 14

874 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE III (Continued)

Growth | Dry wt. Ce. N/50 KOH | Wt. of sugar per
Series | Initial | period ma Final Рн | to neutralize cc. of solution
No Рн а mgms 10 сс. filtrate
8 195.0 5.6 14.7
8 6.0 12 380.0 6.4 5.9
16 441.6 6.4 6.1
28 577.8 7.3 3.7
8 199.0 6.2 10.6
9 6.5 12 353.0 6.6 9.9
16 434.1 7.2 8.9
28 452.0 7.4 1.3
8 189 6.4 4.3
10 7.0 12 318 6.4 4.35
16 447 6.2 8.6
28 512 7.2 1.75
8 144 6.4 3.4
11 7.5 12 358 6.2 9.6
16 455 6.3 9.4
28 502 7.4 0.30
8 109 6.9 3.8 30.2
12 8.5 12 299 7.0 .4 22.2
16 421 7.6 .551 9.8
28 529 8.4 1.31 0
8 6.7 5.2
13 8.8 12 257 6.9 2.7
16 405 t— ——
28 487 8.5 1.21
8 96 8.4 .95}
14 9.0 12 251 7.9 701
161 --- --- س‎
28 427 7.8 ‚851

* The values given аге the averages of 2 cultures.

1 Contaminated.

1.2 N HCl

The relations of time to growth in the series are shown in the
curves in fig. 2. At the end of the 28-day period, the best growth,
as indicated by the dry weight of mat, occurred in series 3-8 in-
clusive, represented by Py values of 3.0, 4.0, 4.5, and 5.5. Тһе
maximum growth occurred in the cultures started at Py 3.5.
The growth of the fungus was slow in series 1, 2, and 3 for the
first 8 days, but later the growth rose rapidly and reached a
maximum at the end of 28 days as high as other series started
with higher Py exponents. Judging from the behavior of the
cultures in series 1 and 2, the fungus tolerates a culture solution

1927]
NEAL—COTTON WILT 375

Dry Weight Of Mat Milligrams

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Мұз Бе s Initial Py
6028 В bs À als P Ld ake 5 s

Fig. 2. Showing the relation of time to the growth of Fusarium vasinfectum.

strongly acid. However, the maximum growth attained at the
close of the experiment was notably less with the more acid
cultures. АП series initially acid became less so, while all
initially alkaline became slightly acid excepting a few started on
the extreme alkaline side of Py 7, which, although becoming acid
at first, reversed the reaction later and were distinctly alkaline at
the close of the growth period.

[Vor. 14
876 ANNALS OF THE MISSOURI BOTANICAL GARDEN

In the majority of the series changes in total acidity were
closely correlated with changes in hydrogen-ion concentration.
Decrease in the amount of sugar in the culture was also cor-
related with an increase in dry weight of mat. At the close
of the experiment (28 days) sugar had entirely disappeared from
the culture solution.

FACTORS INVOLVED IN PRODUCING WILT

The wilting in the case of many Fusarium diseases of plants has
been explained by numerous plant pathologists and botanists as
being due to mechanical plugging up of the vascular system by
the vegetative growth of the fungus. Smith (799) suggested
this theory in the case of cotton, cowpea, and watermelon plants,
which were infected with closely related fungi of the genus
Fusarium. Не found considerable mycelium within the water-
conducting tubes, and assumed that the amount present was suf-
ficient to act as an obstruction of liquid upward, thus causing the
condition known as wilt. Among others who have held similar
views in the past in connection with cotton wilt are Duggar (709),
Stevens (721), and Orton (700, '08, 19). It may be stated in
fairness to the above investigators that in the case of the majority
of the Fusarium diseases, this idea was generally held by botanists
and plant pathologists until a few years ago; the writer is fully
aware, too, that some of the investigators mentioned above,
perhaps all of them, no longer hold such views.

The theory that the wilting may be due to simply mechanical
plugging of the water-conducting vessels of the plant, however,
has recently been questioned by a number of investigators, and
the numerous experiments along this line have been concerned
with а number of Fusarium wilt diseases of plants of very diverse
habitat. Because of the fact that the writer has frequently
observed considerable fluctuation in the prevalence of cotton
wilt in fields which were known to be infested, and the fact that
the disease has appeared to be more severe during dry seasons,
especially in recent years, it was of interest to make some studies
regarding the phenomena concerned in wilting by Fusarium
vasinfectum.

If the cause of wilting was simply a response to mechanical

bai cur os co ока

1927]
NEAL—COTTON WILT 377

obstruction, it is conceivable that with ample moisture, there
would perhaps be a greater vegetative growth of the fungus within
the tissues and a corresponding increase in wilt because of greater
obstruetion by the mycelium. In this connection, observations
made in Mississippi by the writer are somewhat in disagreement
with the statement by Elliott (21). He remarks: “It is a well
observed faet that cotton wilt is more severe in а wet season
than in a drier one. The reason for this is because the weather
conditions which favor a rapid ‘sappy’ growth of the cotton
plant also favor the development of the wilt fungus, and if the
water supply becomes at all short, such plants suffer quickly and
severely." Observations made in Mississippi and other states
lead the writer to believe that the reverse may be true; i. e., the
disease is certainly more serious during prolonged panda of
abnormally dry weather, but may frequently develop suddenly
and in destructive form as soon as such periods are broken by
ample rainfall. Beal (726) has also noted a greater prevalence
of the disease during periods of dry weather. Therefore, since
а study of the relation of moisture conditions to the occurrence
of wilt did not strengthen the theory that the wilt may develop
as the result of mechanical plugging of the vessels owing to
increased growth of the fungus, it was desirable to study other
factors which might be involved. "This appeared essential, too,
since mieroscopie examination of the vascular system of many
cotton plants with typical wilt symptoms failed in many instances
to reveal the presence of hyphal filaments in the vessels, and in
others, when the fungus was detected, it was only in very meager
amounts.

Some of the experiments of other investigators on the factors
involved in wilting of plants by other Fusaria may be mentioned.
Brandes (719) found that cultures of Fusarium cubense, grown in
Richards’ solution, when filtered free from the fungus, produced
а marked wilting in buckwheat and bean plants, as well as banana
leaves. He obtained the same results using Uschinsky's solution
аз а medium, and concluded that in the disease known as banana
wilt, wilting is not due to plugging of the vessels by mycelium,
but is probably the result of toxic excretions by the fungus.

Bisby (719) reports the results of experiments in which leaves

[Vor. 14
378 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of different plants were placed in filtrates from Fusarium оху-
sporum, F. discolor sulphureum, Fusarium from pea, and other
fungi. Wilting occurred within a few hours with some extracts,
& fact which he interprets as not to be explained by changes in
acidity. Excised leaves placed in uninoculated media and іп
water did not wilt until considerable time had elapsed. Тһе
injurious effect was also noted after boiling the filtrates and after
diluting them considerably. He concludes that the wilting is
not due to some poison which is specifically active against certain
plants, since potato leaves wilt as readily in old Rhizopus solu-
tions as in solutions in which F. oxysporum had grown.

Haskell (19) states that from numerous microscopic observa-
tions of the stems of potato plants infected with Fusarium оху-
sporum, he found no instance of a stoppage of the tracheae suf-
ficient to shut off the passage of sap.

In 1920, Young and Bennett (721) made an effort to determine
the method by which Fusarium oxysporum induces wilting of
potato plants. They found that filtrates of this fungus growing
upon Richards’ solution were increasingly toxic to potato plants
as the age of the cultures increased up to the fortieth day.
Furthermore, with this increase in toxicity there was noted,
after a 10-day period, a gradual decrease in hydrogen-ion con-
centration, from an initial reaction of Py 5.0 to Py 7.4 on the
fortieth day. But when the control (uninoculated) solution was
adjusted to Pg 7.4, there was no toxicity noted when cut stems
were placed in it. From this it is concluded that the change
in reaction, while not the primary cause of wilting, indi-
cated the presence of a compound with a slightly alkaline re-
action. Autoclaving or boiling did not alter the toxicity of this
substance.

Іп 1921, Ajrekar and Bal (721), of India, working on cotton
wilt, were unable to obtain wilting of cotton plants by means of
the alcoholic extract or by the method described by Hutchinson
(13). They did not, however, conclude from their very meager
tests (only 2 plants were used) that no toxins were produced by
the fungus; on the contrary, when they sectioned diseased plants
and examined them microscopically the number of vessels which
were not occupied by the fungus was so great in comparison with

1927]
NEAL—COTTON WILT 379

the number occupied that they suspected the action of a toxic
substance rather than a mechanical plugging up by fungus
mycelium.

Other investigators who have attempted to determine the
cause of wilting by various Fusaria or the nature of substances
excreted by such parasitic fungi are Picado (723), Fahmy (723),
Goss (24), and others.

In the biennial report of the Director of the Kansas Agricul-
tural Experiment Station (page 75) which appeared in 1924, it is
recorded that Fusarium Lycopersici has been found to secrete an
enzyme which, when precipitated, dried and re-dissolved in dis-
tilled water, will cause young plants to wilt rapidly when the cut
stems are immersed in the solution. Тһе toxicity of the enzyme
is destroyed by boiling. Both residue and the dialysate of
nutrient solutions upon which the fungus was grown produced
wilting of young plants. From this it is concluded that two toxic
substances exist, one a colloid and the other a crystalloid.

Quite recently, Rosen (726) has shown that filtrates represent-
ing cultures growing on Richards’ solution are quite toxic to
cotton plants, while filtrates of cultures growing on media
containing organic nitrogen, such as Uschinsky’s asparagine
solution or peptone-beef broth, are non-toxic. He raises the
question if this might not explain why cotton wilt is more preva-
lent in light sandy soils devoid of appreciable quantities of organic
nitrogen than in richer types of soils. By subjecting filtrates of
the fungus grown on Richards’ solution to distillation in vacuo,
as well as to ordinary boiling and testing, the distillates and also
the residues were found to possess toxic properties, the residue
being considerably more toxic. Тһе filtrates of 2- or 3-weeks-
old cultures on Richards' solution gave positive tests for nitrites.
These were found in quantities ranging from 0.012 mgm. to 0.04
mgm. of nitrogen for each cc. of solution. Using chemically pure
sodium nitrite solution comparable to the concentration found in
the filtrates, it was determined that this is markedly toxic to
cotton plants. Microscopic observations of the vascular ele-
ments of wilted cotton plants by Rosen (’26) substantiate the
findings of the writer and others that wilting is not due to
mechanical plugging up of the vessels by the fungus. Rosen

[Vor. 14
380 ANNALS OF THE MISSOURI BOTANICAL GARDEN

records that in badly infested fields wilted cotton plants are found
occasionally in whose vascular tissues the fungus may be appar-
ently absent, and he suggests that the wilting and interior dis-
coloration of the xylem in these cases is due to the formation of
toxic substances by the fungus in the soil. "This author concludes
that the wilting of cotton infected with Fusarium vasinfectum is
considered to be due to poisonous chemical substances formed
by the fungus, and that filtrates of Fusarium vasinfectum growing
on Richards’ solution possess at least two substances poisonous
to cotton. One is a volatile compound with an alkaline reaction
and the other is an inorganic salt in the form of nitrite. It is
evident from the above references and others not enumerated in
this paper that in many of the Fusarium diseases, including those
of banana, cotton, potato, and tomato, the theory of mechanieal
plugging of the fungus as being the factor responsible for wilting
is no longer accepted. It is now suggested that toxic substances
are probably responsible for wilt symptoms and related patho-
logical phenomena.

EXPERIMENTS RELATING TO TOXIC ACTION BY THE FUNGUS

In order to obtain some information on the probable toxic
properties of substances given off by Fusarium vasinfectum, the
following experiment was performed. Тһе fungus was grown at
room temperature for 18 days іп 150-cc. Erlenmeyer flasks on
50 cc. of Duggar's nutrient solution previously described, and
treated as follows: In a series of 4 flasks, the mats were collected
on filter paper by means of a Buchner suction filter, and the
filtrates boiled for 10 minutes. Another series of 4 flasks was
treated similarly but not boiled. In another series of 4 flasks,
the mycelial mats were collected on filter paper as described above,
ground with 20 grams of fine quartz sand for 15 minutes in &
mortar, and the mixture extracted with 75 cc. of sterile distilled
water. Another was prepared in the same way and the extract
boiled for 10 minutes. А final series of filtrates was also prepared
as described above, and the solution afterward diluted with sterile
distilled water at the rate of 1 part filtrate to 3 parts of water.
After cutting away the roots, cotton seedlings which had been
sprouted in washed sand and which averaged 6 to 8 inches in

1927]
NEAL—COTTON WILT 381

height were immersed in 2 of the flasks in each series. In the re-
maining 2 flasks of each series, the roots of the plants were left
intact. Control flasks contained Duggar’s nutrient solution and
sterile distilled water. All series were placed in the greenhouse
in bright light at a temperature of approximately 75-80°

and examined at intervals of 3, 6144, and 36 hours. Тһе daia
obtained are presented in table ту.

TABLE IV

ЕЕЕЕСТ OF FILTRATES AND MYCELIAL EXTRACTS OF 18-DAY CULTURES
OF FUSARIUM VASINFECTUM ON COTTON SEEDLINGS

| Wilt symptoms present after

"e Treatment
| зы. | бум. 36 hr.
Filtrate
1 Boiled None Positive Very pronounced
la Unboiled None Positive Very pronounced
rps Boiled Slight Positive Very pronounced
1 с* Unboiled one None None
Extract of mycelial mat
2 Boiled None Positive Very pronounced
2а Unboiled None None ositive
2 b* Boiled None Slight Pronounced
25* Unboiled None Slight Very pronounced
Filtratet
3 Boiled None None Slight
3a Unboiled None Slight Positive
3 b* Boiled None None None
3 c* Unboiled None Slight Positive
Control |
4} Boiled None None None
4 at Unboiled None None None
5$ Boiled None None one
5a$ Unboiled None None None

* Root removed at Pan of plants.
1 Filtrate diluted 1

1 Duggar's nutrient ux.

$ Sterile distilled water.

From the data recorded, it is observed that both filtrates from
the 18-day-old culture of the fungus and an extract made of
the mycelial mat of the same age possess toxic properties. In
the case of the filtrates, slight evidence of wilting was noticeable

[Vor. 14
382 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

at the end of 3 hours and became quite definite at the end of
6/2 and 36 hours. In uninoculated controls of both Duggar's
nutrient solution and sterile distilled water the seedlings remained
healthy and turgid, even after 36 hours (pl. 32, figs. 2-3). The
extracts made from the mycelial mats appear also to possess toxic
properties in about the same degree as the filtrates (pl. 32, fig. 2).
The seedlings whose roots had been cut away prior to immersing
did not reveal wilt symptoms any more pronounced than those
in which the roots were left intact. Diluting the filtrates in the
ratio of 1 part filtrate to 3 parts of sterile distilled water appears
to have lowered the toxicity to some extent, and in these series
wilt symptoms became definite only after the maximum period
of 36 hours. Тһе toxic substance is approximately thermostable
in both the filtrate and mycelial extract used in these experi-
ments. Тһе Py of the nutrient solution prior to inoculating was
5.6, and at the end of the 18 days the filtrate was 6.8. Additional
experiments are now in progress to ascertain if Rosen's (26)
results are correct concerning the presence of nitrites in filtrates
of Fusarium vasinfectum.

PATHOGENICITY OF FUSARIUM VASINFECTUM АТК.

During the last few years different investigators have raised
the question as to the real parasitic nature of the fungus Fusariwm
vasinfectum in connection with wilt of cotton. For instance,
Ajrekar and Ва (21), in their inoculation experiments, do not
appear to have secured uniformly successful infection, and they
suggested that such negative results perhaps may have been due
to the fact that the season was far advanced when their experi-
ments were carried out. They concluded from the result of their
inoculation experiments that the Indian species of cotton wilt
was not a virulent parasite.

In 1924 Dastur advanced the theory that cotton wilt is not due
to the parasitic action of a species of Fusarium, but that it may
be caused by the excessive accumulation of aluminium and iron
salts within the tissues of the plants. Не arrived at this con-
clusion because there was a constant accumulation of iron and
aluminium compounds in the tissues of wilting plants and a
constant absence of these compounds in the tissues of healthy

1927]
NEAL—COTTON WILT 383

plants. Не used qualitative microscopical tests for these metals.
Tissues of plants which were infected with Rhizoctonia Solani
Kühn were also examined for aluminium and iron compounds
with negative results. Dastur also failed to isolate a parasitic
organism from the wilting plants, and suggested that the ассит-
ulation of the above compounds may have some correlation with
the wilt and that the species of Fusarium which had been isolated
from wilting plants in different cotton tracts may be only a con-
tributory factor in hastening the death of the plant and merely
follows in the wake of the accumulations of these compounds in
the tissues.

Dastur worked on varieties and selections of cotton listed as
AK, AK, and Roseum. These were inoculated with different
strains of Fusarium which had been isolated from wilted cotton
plants. He used both virgin soil and sand in pots, and inocu-
lations were made after scraping away the soil an inch or two
deep around the plants; and the entire contents of agar tubes, in
which the fungus was growing luxuriantly, were placed around
the roots. The soil thus scraped away was then returned. Іп
some pots the upper lateral roots were cut before introducing the
fungus, but the tap roots were uninjured. Ніз infection experi-
ments were uniformly negative; the controls and inoculated
plants were equally healthy and developed good flowers and
bolls.

During the progress of this investigation, the writer has en-
deavored to substantiate the findings of Dastur with reference
to the presence of iron and aluminium compounds in the tissues
of cotton plants, particularly with plants showing typical wilt
symptoms, such as stunted growth, yellowing of leaves, and pro-
nounced vascular discoloration. On August 28, 1926, 3 Trice
cotton plants with typical wilt symptoms and 3 healthy plants
of the same variety were collected in a field of the Mississippi
Experiment Station. These were referred to Dr. W. F. Hand,
State Chemist, with a request that quantitative analyses be
made for aluminium and iron oxides in the leaves, stems, and
roots. By using official quantitative methods, the per cent of
iron oxide was determined in the above parts of both healthy and
diseased plants, but the results for aluminium were discarded

[Vor. 14
384 ANNALS OF THE MISSOURI BOTANICAL GARDEN

because of the probable inaccuracy of the present method for
quantitative determinations of these compounds in the amounts
occurring in cotton plant tissues. "The per cent of iron oxide and
total ash of the leaves, stems, and roots of healthy and diseased
Trice cotton plants are given in table v.

TABLE V
QUANTITIES OF ҒеОз FOUND IN TRICE COTTON PLANTS*
Healthy Per cent Diseased Per cent
'Total ash in leaves 14.32 'Total ash in leaves 18.54
Iron oxide in leaves 0.028 Iron oxide in leaves 0.04
otal ash in 8 .75 Total ash in ste 4.11
Iron oxide in s 0.006 Iron oxide in 0.008
ash in roots 3.45 ''otal ash in roots .70
Iron oxide in roots 0.016 Iron oxide in roots 0.014
Total ash in plant 21.32 Total ash in plant 26.35
Iron oxide in plant 0.050 Iron oxide in plant 0.062

* Average of 3 plants

Additional Ттісе cotton plants in а more advanced stage of
maturity were collected at the South Mississippi Experiment
Station on September 8, and quantitative analyses made for iron
oxide in the stems and roots. Тһе results, however, were ргас-
tically the same as those embodied in table v; i.e., the amount of
iron oxide found in both healthy and wilt-infected plants was
about the same. The above results do not indicate any close
correlation between iron accumulations in the tissues of the
plants and wilt infection. Тһе writer has also made numerous
qualitative tests for iron with healthy and wilt-infected cotton
plants of approximately mature ages. For these tests, potassium
acid thiocyanate solution was used, but no appreciable difference
could be observed in the stems and roots of healthy and diseased
tissues.

In order to test further Dastur's (724) theory of aluminium and
iron accumulations in the tissues of plants as a primary factor
involved in producing wilt, another experiment bearing upon this
question was planned. In the summer of 1926, a field was visited
at the South Mississippi Station which was badly infested with
wilt and which had been so for years previous. А quantity of
soil from this field was collected and shipped to the laboratory for

1927]
NEAL—COTTON WILT 385

some pot experiments. Eighteen pots were filled with the soil
and half of them sterilized in the autoclave at 15 pounds pressure
for 5 hours. Тһе pots containing the sterile and unsterile soil
were then planted with delinted Delfos 6102 cotton seed and
placed in benches in the greenhouse, the date being September 15.
It was considered that if the disease could be induced in the
sterile pots in the absence of the fungus, some significance could
be given the probable accumulative effect of certain compounds
in the tissues of the plants. The plants were allowed to grow until
December 3. Approximate readings were made on November 1,
and more accurate readings made by cutting the stems of the
plants for evidence of vascular discoloration, on December 3.
The results obtained are shown in table ут.

TABLE VI
EXPERIMENTS TO DETERMINE THE PATHOGENICITY OF COTTON WILT,
FUSARIUM VASINFECTUM ATK., IN THE GREENHOUSE,
SEASON 1926—VARIETY DELFOS 6102

Number infected plants
Pot No. Soil treatment
Apparent wilt Nov. 1 | Actual wilt Dec. 3

1 Unsterile 2 3
1A Sterilized 0 0
2 Unsterile 1
2A Sterilized 0* 0
3 Unsterile 1 2
ЗА Sterilized 01 0
4 Unsterile 2 6
4А Sterilized 0

Unsterile 41 5
5А Sterilized 4 0

Unsterile 0 1
6A Sterilized 0 05
7 Unsterile 8 8
ТА. Sterilized 0 0
8 Unsterile 0 8
8A Sterilized 0 0
9 Unsterile 0 2
9A Sterilized 0 0

* 2 dead with Rhizoctonia.

t1 dead with Rhizoctonia.

13 killed by Fusarium vasinfectum.
$ All infected with Rhizoctonia.

The results obtained in the above experiment indicate very
clearly that the strain of Fusarium vasinfectum present in the
soil obtained from the infested field at the South Mississippi

[Vor. 14
386 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Station was pathogenic and quite virulent, since infection occurred
in all of the pots which were not sterilized, and in some cases all
of the plants (8 in number) were infected when the plants were
taken down and the experiment terminated (pl. 32, fig. 1).

It is quite likely that the failure of Dastur (24) to secure in-
fection as the result of his inoculations was due to the method he
employed. Тһе writer has also frequently found it difficult to
infect plants under greenhouse conditions in sand cultures par-
ticularly, even after an interval of 90 days, if the fungus is placed
merely around the roots of the seedlings. Such a method is not
as reliable as that of mixing the fungus thoroughly with the soil
or sand prior to sowing the seed. From these and other infection
experiments it would seem that cotton wilt cannot be attributed
to the accumulation of certain compounds within the tissues, such
as aluminium and iron, but that the fungus is responsible for the
disease and the attendant symptoms. Тһе recent work of
Butler (726) substantiates this view.

Although the above experiments do not indicate any relation
between cotton wilt and the accumulation of iron or aluminium
within the tissues of the plants, it is interesting to note the results
which Hoffer (723) has obtained bearing upon this question as it
relates to corn root-rot. Не has found that in the presence of
soluble iron and aluminium salts in the soil or in the presence of
other growth-inhibiting factors corn exhibits an abnormal ac-
cumulation of these elements in the vascular plate tissues in the
nodes of the stalk. There is also some concentration in the leaf
tissues. Conditions result which retard the translocation of food
to the roots. The roots later cease to function and accordingly
their vitality is greatly impaired. He has contributed some very
definite data on this point. Тһе most severe cases of root-rot
with which he has come in contact were found in soils notable
for their deficiencies of lime and phosphate. Potash deficiencies
were also often noted. His experiments indicate that root-rot
is not due to fungi (as the primary factor), but that these organ-
isms are usually associated with and involved in the crop failure.

In some later experiments (23), Hoffer and Trost have added
materially to the above. А set of acid soils was treated in a
manner to increase or decrease the aluminium absorbed by the

1927]
NEAL—COTTON WILT 387

plant and to determine the intensity with which soil organisms
would attack plants under the conditions. These experiments
showed that both lime and phosphate lowered the assimilation
of aluminium. Тһе roots of all plants showed rot lesions caused
by Fusarium moniliforme, but the severity of the damage was
modified by the soil treatment and was serious only in the presence
of aluminium salts and deficiencies of lime and phosphates.
Hoffer also determined the relation of aluminium salts to the
susceptibility to root-rot or the invasion by one of the root-rot
fungi. In these experiments he used Gibberella saubinetti. The
results showed that in the presence of aluminium salts there was
a predisposition of the plant to invasion by the fungus. А
potash reserve in the plant increases the resistance by lessening
markedly the accumulations of iron and aluminium in the nodal
joints.

In this connection, McGeorge (725) has also recently found that
aluminium is a factor directly associated with the retarded growth
of sugar-cane on the acid mauka lands in Hawaii, and further he
has found abnormal accumulations of aluminium and iron at the
nodes of certain varieties of cane. In the case of these highly
acid soils, he considers the retarded growth of the sugar-cane as а
direct toxic action of aluminium and not a phosphate deficiency.

ADDITIONAL EXPERIMENTS BEARING UPON ALUMINIUM TOXICITY
OF COTTON AND COTTON WILT
Because of the interest manifested regarding the probable
absorption of soluble aluminium salts in certain soils and predis-
posing such plants as согп and perhaps sugar-cane to invasion
by soil-inhibiting organisms, it was of interest to study further
the róle that such compounds might play in predisposing cotton
plants to infection by Fusarium vasinfectum. Accordingly, 2
series of water cultures in battery jars containing aluminium
salts in varying concentrations were prepared. Shive’s (716)
В.С, nutrient solution was used for this purpose and the alumin-
ium was supplied in the form of the chloride and the sulphate,
and added to the nutrient solution to give the following normal-
ities. All treatments were made in duplicate:

[Vor. 14

388 ANNALS OF THE MISSOURI BOTANICAL GARDEN
SERIES I
p.p.mil. А1
1. Со, Бато ИМЕН... on rl None
2. Basic nutrient plus aluminium sulphate to make N/2500.. 3.6
3. Basie nutrient plus aluminium sulphate to make N/1000.. 9
4. Basic nutrient plus aluminium sulphate to make N/500. . .18
5. Basic nutrient plus aluminium sulphate to make N/250. . .36
SERIES II
p.p. mil. AI.
Control, basic nutrient опіу......................... None

Basic nutrient plus aluminium chloride to make N/2500.. 3.6
Basic nutrient plus aluminium chloride to make N/1000.. 9
Basic nutrient plus aluminium chloride to make N/500. . .18
Basic nutrient plus aluminium chloride to make N/250. . .36

oo уи

Delfos 6102 cotton seedlings, the seed of which had been
previously delinted with sulphuric acid and sprouted in washed
sand, were used in these cultures. Twelve seedlings, supported
by galvanized wire of a suitable mesh, were placed in each jar
when the radicles were about 4 inches long and allowed to grow
in the nutrient solutions for 29 days. The solutions were changed
in the jars regularly once a week and precaution taken to prevent
any contamination with algal forms, etc.

On comparison of the cultures from time to time, toxicity was
scarcely apparent in either the series containing the aluminium
sulphate or aluminium chloride until a concentration of N/250
(36 p.p. mil. aluminium) was reached. In these cultures, repre-
sented in each series by number 5, some toxicity was apparent,
not only as related to height of the seedlings, but as to the root
development which was also considerably retarded. Тһе effect
of these salts when added to the nutrient solution is best indicated
by the photographs in pl. 33, figs. 1-2. At the end of the 29-day
growth period, the lengths of the tops of the plants were measured
and notes made on the condition of the roots. These data are
summarized in table уп.

Of the two salts used, the aluminium chloride appears to be
slightly more toxic to cotton under the conditions of the above
experiment, especially the N/500 and N/250 concentrations.

1927]
NEAL—COTTON WILT 389

TABLE VII

RELATIVE LENGTH OF COTTON SEEDLINGS GROWN IN VARYING AMOUNTS
OF ALUMINIUM SULPHATE AND ALUMINIUM CHLORIDE FOR 29 DAYS

Length of tops Condition of r
oots

Culture Inches

Sulphate | Chloride Sulphate Chloride
Control 11.5 12 Good—normal Good—normal
N/2500 12.0 10 Good—normal Good—normal
N/1000 10 9.5 Good—normal Good—normal
N/500 12 10.5 Good—normal Fair—few laterals
N/250 8.4 7.8 Fair—few laterals PET few

aterals

After the length of plants in the above experiment had been
determined they were put back into the battery jars and to each
was added freshly prepared Shive’s Е.С; nutrient solution.
Each culture was then immediately inoculated with a spore
suspension of Fusarium vasinfectum, in order to ascertain whether
the plants which had received varying amounts of aluminium
sulphate and chloride would reveal any decided differences to
invasion of the fungus as compared with the control plants or the
ones which had received the basic nutrient solutions only. At
the end of one week after inoculation, all cultures were infected
with the fungus (pl. 33, fig. 3) and so far as could be observed, the
control plants were equally susceptible to attack by the fungus.

Judging from the above experiment, it seems that aluminium
sulphate and aluminium chloride are not appreciably toxic to
cotton plants in concentrations lower than 36 parts per million.
Moreover, it is believed by the writer, too, that such a concen-
tration of soluble salts of aluminium does not occur in soils of
cotton tracts generally in the southern states, and that this
factor is perhaps of little consequence in predisposing plants to
infection by Fusarium vasinfectum as has been supposed by
Dastur (724).

EFFECT OF NUTRITION UPON THE DEVELOPMENT OF COTTON WILT

Greenhouse experiments.—In studying the effects of varying
conditions of nutrition upon the development of wilt of cotton,
a nutrient solution recommended by Hartwell апа Pember (715)
was used as a basic nutrient. This nutrient had the following
composition:

[Vor. 14

390 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ЖАО ВИ, 15 сс. per litre
EN TGLNUM TS В 10 ec. per litre
A A 55 PM FNERIT EAS 8 cc. per litre
AN MISOLTHEIOL. ар 8 cc. per litre
8.3 grams per litre СаН(РО,.),........ 1 cc. per litre
о асыусы Trace

Stock solutions of the salts in the above concentrations were
prepared in 3-litre bottles and 12 similar bottles were used for
the desired dilutions for the different series of treatments.

The series of treatments used were as follows:

Series I—Basic nutrient only.

Series II—Same as I but with phosphorus omitted.

Series III—Same as I but with phosphorus increased 20 times.

Series IV—Same as I but with potassium omitted.

Series V—Same as I but with potassium increased 10 times.

Series VI—Same as I but with potassium increased 20 times.

Series VII—Same as I but containing 1/10 less nitrogen.

Series VIII—Same as I but with nitrogen increased 2 times.

Series IX—Same as I but with calcium omitted.

Series X—Same аз I plus 24 p.p.m. iron from ferric ammonium
sulphate.

Series XI—Same as I plus 54 p.p.m. iron from ferric ammonium
sulphate.

Series XII—Same as I plus 12 p.p.m. aluminium from potas-
sium aluminium sulphate.

Equivalent concentrations of NaNO, Na;HPO, were used
in the above Hartwell and Pember nutrient solution when cal-
cium was omitted from the treatment, and likewise an equivalent
of NaCl was employed in the treatments which did not receive
potassium.

Three-gallon glazed pots containing washed sand were used
for growing the plants, and each series of treatments contained
4 pots. Тһе experiment was begun on August 9, 1926, and on
this date each pot of the different series of treatments was
planted with delinted seed of Delfos 6102, a staple cotton which
is quite susceptible to wilt. After the seedlings were a few days
old, they were thinned to 6 plants in each pot. During the first

1927]
NEAL—COTTON WILT 391

6 weeks of growth, each series of cultures was given 300 cc.
per pot of the nutrient once а week, and after 3 such weekly
applieations each pot was washed out thoroughly with one
gallon of distilled water to prevent the possibility of excess salt
accumulation. After 6 weeks had elapsed and the plants became
larger they were given 300 cc. of the nutrients per pot every 10
days, and after each pot had received 3 such consecutive 10-day
treatments they were washed out with the distilled water as
indicated above. On September 9, when the plants were &
month old, all series were inoculated with Fusarium vasinfectum
by applying the entire contents of a bean-agar test-tube culture
of the fungus to the roots of the plants. On Oetober 5, all
plants in each series were again inoculated by applying to the
roots the mat of the fungus which had grown on 50 cc. of Duggar's
solution for 10 days. А final inoculation was made on October
20, bits of diseased stems of mature Delfos 6102 cotton plants from
а wilt-infested field being placed around the roots of the plants.

The schedule of treatments given above was followed as closely as
possible, but no attempt was made to keep the pots at a constant
soil moisture. Moisture determinations were made, however, with
untreated control pots until the plants died of starvation, and
these results indicated that the moisture content in the treated
pots was approximately 25 per cent of the water-holding capacity
of the sand.

Very few of the plants in any of the series of treatments were
killed by the wilt, but on March 5, after an interval of almost 7
months, the different series of treatments were terminated and
examinations of the plants made as to their green weight, dry
weight, vascular discoloration, lateral root discoloration, and the
number of plants showing positive infection. Evidence of
positively wilt-infected plants was recorded only when such a
symptom as pronounced vascular discoloration was obtained by
splitting the plants with a stainless steel knife. Тһе numbers
with lateral root discoloration were recorded, and undoubtedly,
if these plants had been cultured, Fusarium vasinfectum could
probably have been isolated, though time would not permit such
cultural tests for evidence of the fungus. Тһе results obtained
in these nutrition experiments are presented in table уш.

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

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1927]

398

NEAL—COTTON WILT

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[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

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1927]
NEAL—COTTON WILT 395

The plants in the pots of each series were photographed on the day
the experiment was completed, and they are shown in pls. 34-35.

An examination of the data recorded in table VIII reveals some
interesting facts with reference to the nutrition of cotton and the
probable réle it may play in increasing or lowering the resistance
of the plant to invasion by Fusarium vasinfectum. So far as the
writer is aware, no previous attempt has been made to study these
questions with respect to cotton and their relation to the develop-
ment of wilt under greenhouse conditions. It was observed that
the disease developed in all of the series of treatments. It was
less in series I, which received the basic nutrient without any
variation of its nitrogen, potassium, phosphorus, or iron content.
As evidenced by the vascular discoloration of stems and tap roots,
the infection was highest in series X and XI, containing respec-
tively 24 and 54 parts per million iron from ferric ammonium
sulphate. Next in number of wilt-infected plants was series
VII, containing 1/10 less nitrogen, series XII, with 12 parts per
million aluminium from potassium aluminium sulphate, and series
IV, in which potassium was omitted. The pots in which the potas-
sium content was increased were notably free of the disease, as
indicated by the lack of vascular discoloration in both stems and
roots and the absence of lateral root discoloration. The same is
decidedly true of series ІХ, in which calcium was omitted. In
the pots which received marked increases of phosphorus and
nitrogen the number of plants with vascular discoloration was
low, but the lateral root discoloration was somewhat high.

As revealed by the dry and green weights, the plants in the
pots from which calcium was omitted made the most vigorous
growth. This was very noticeable as compared with the other
series. No toxicity developed in series XII, which received the
basic nutrient plus 12 parts per million aluminium salt, and the
growth of these plants was good despite the fact that the examin-
ation of the roots revealed 7 infected plants and several with
lateral root discoloration.

The data as presented do not show any consistent diminution
or increase of infections by the various treatments with the
exception of the ones receiving increased potassium, the ones from
which calcium was omitted, and those in which the iron content

[Vor. 14
396 ANNALS OF THE MISSOURI BOTANICAL GARDEN

was high. Тһе infection as evidenced by vascular discoloration
of stems and roots was low in the pots receiving twice the amount
of nitrogen, but, on the other hand, the lateral root discoloration
in this series was high, and the fact that the green and dry
weight was only a little better than series I (control pots) does
not indicate any marked effect as the result of increasing nitrates
in the nutrient. Тһе same may be said of inereased phosphate
in these treatments. Increased potassium content appears to
have increased resistance to the fungus, but these plants did not
make any better growth, as judged by the dry weight, than did
the plants in series IV, in which potassium was omitted. Тһе
specific rôle that potash may play in increasing resistance in
cotton to invasion by a fungus like that causing wilt is unknown.
We know that it is important in the translocation of food materials
within plants, and, as Hoffer (723) suggests in the case of corn,
it may also be of importance in the maintenance and normal
functioning of the vascular elements of cotton.

On studying the behavior of the plants in the series with high
iron content, we find the infection is somewhat high, and this
would lend support to Dastur's (724) theory of iron accumulation
as being а predisposing factor to invasion by the fungus. How-
ever, this fact is not borne out by quantitative analyses of the
tissues of diseased and healthy plants nor by inoculation experi-
ments, as referred to previously in this paper. Because of the
pressure of other work, analyses of the ash of the plants in these
experiments were not made, but additional nutrition experi-
ments are in progress and this point will receive attention.

The effect of commercial fertilizers upon cotton-wilt infection
has also been studied to some extent under greenhouse con-
ditions. On February 3, 1926, duplicate sand cultures were
prepared in 10-quart galvanized zine buckets, each of which
had been equipped with special aerating equipment in the form
of 2 galvanized zinc tubes attached to the opposite ends of an
inverted trough. The sides of the trough supporting the tubes
were cut ‘‘tooth-like” to facilitate percolation of water when
introduced into the buckets. Тһе tubes were also, of course,
hollow and when placed in the buckets prior to filling the latter
with sand, permitted the plants to be sub-irrigated; at the same

1927]
NEAL—COTTON WILT 397

time this facilitated evaporation from the bottom. Approxi-
mately 8 kilograms of sand were used in each bucket and the
fertilizer was used at the rate of about 800 pounds per acre, re-
garding 2,000,000 pounds of soil six inches deep as representing
approximately one acre. This is in accordance with the usual
fertilizer practice in pots when computing on an acre basis.! In
these experiments, Mississippi Station Trice cotton was used,
and the seed delinted with sulphuric acid before planting.

The sand in one of the buckets of each treatment was mixed
thoroughly with a bean-agar culture of Fusarium vasinfectum and
the other left uninoculated аз a control. The plants were allowed
to grow for 60 days, and then data recorded as to relative height.
of plants, vigor, and the number infected as evidenced by yellow-
ing of leaves, vascular discoloration, etc. The results are shown
in table rx.

TABLE IX
EFFECT OF COMMERCIAL ym ee UPON THE DEVELOPMENT OF
COTTON Td IN THE HOUSE. RATE OF APPLICATION
800 PU MEE PEL yv SEASON 1926
Fertilizer | Relative height of Vigor No. Infected plants after 60 days
plants, inches plants Number Per cent

-4-0 check 9.5 Good 0 0

-4-0 10.0 Good 8 37.5
8-4-4 check 11.5 Good Ч 0 0

—4-4 10.5 Good 0 0

-0-4 check 9 Good Ч 0 0

-0-4 8.5 Good 5 62.5

-4-4 check 8.5 Fair 0 0

0—4—4 8 Fair ; 4 50
1 check 12 Good ; 0 0
12-4-4 11 Good ; 0 0

From the above results, it appears that potash, when used at
the above rate, may have increased the resistance of the plants
to infection by the fungus. Тһе results obtained are best indi-
cated in pl. 36, fig. 1.

In other experiments of this nature, the potash in the formula
was increased to 6 per cent and the mixture applied at the rate
of 600 pounds per acre. However, at the end of 60 days, when
the experiment was terminated, no appreciable difference in the

! From information жол the writer by Dr. Oswald Schreiner, of the Bureau
of Plant Industry, U. S. Dept. Agr.

[Vor. 14
398 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN

amount of infection was noted between plants receiving potash
and those to which none was added.

CONTROL

Field experiments.—Field experiments to determine the effects
of commercial fertilizers upon the control of wilt were conducted
during the season of 1926 at 2 localities in Mississippi. One set
of experiments was located at the Central Experiment Station,
А. & M. College, and another at the Poplarville Branch Experi-
ment Station. Тһе soil in the experimental plots at the Central
Station was artificially inoculated, whereas the field used for
recording infection data at the Poplarville Branch Station was
already infested with wilt and the plots had been fertilized in the
same manner for a period of 5 consecutive years. The size of
the plots in the fertilizer treatments at the Central Station was
approximately 1/40 of an acre, and at Poplarville approximately
1/12 of an acre.

TABLE X
COTTON WILT OCCURRING IN LONE STAR 65 VARIETY. FERTILIZER
TREATMENTS AT CENTRAL STATION. SEASON 1926
Yield Total number plants % infection

F n Miei Pers in area* | in areaf | inarea | in area

inocu- inocu- inocu- inocu-
formula bs. ж” Бей lat lat ма Тоба!

- опсе twice once twice
Chec 140 219 42 3.50 5.56 9.06
34-8 600 712 288 63 2.20 3.17 5.37
-4-4 600 572 267 59 8.87 7.35 11.22
600 260 264 61 4.16 5.47 ).63
Check س‎ 128 258 55 3.49 6.67 10.16
8-4 600 684 268 65 3.61 4.30 7.91
-6-4 600 512 251 54 4,25 6.18 10.43
-4 600 492 227 52 5.00 7.06 12.06
Chec = 140 261 51 3.96 1.98 .94
6-4-4 600 536 279 59 4.66 8.49 13.15
4-4-4 60 496 296 64 4.28 7.81 12.09
10-5-5 600 708 267 56 8.74 4.80 . 54
Chec ----- 208 257 55 4.41 4.85 9.26
—4—4{ 60 652 228 42 4.50 6.76 11.26
-4-4 900 724 273 53 4.03 7.55 11.58
-4-4 1200 804 254 54 2.37 2.48 4.85
Check 152 226 47 9.70 5.00 8.70

* Soil inoculated May 4 by plowing under re-inoculated sterilized oats.

T Soil inoeulated again August 9 by pouring 100 cc. of spore suspension of Fusarium
vasinfectum around roots of

1 Lime applied at rate of 4500 pounds per acre, to approximately neutralize soil.

1927]
NEAL—COTTON WILT 399

Тһе plots at the Central Station were seeded to Lone Star 65
eotton early in May, and in due time the total number of plants
and the percentage of infections were determined. Yield records
were obtained in October. "These records are given in table x,
and represent the average of 3 replications of the one-fortieth-acre
plots. The results on infection counts are given іп 2 areas in
each treatment, one being inoculated just before planting time,
the other being a section of the row of each plot which was re-
inoculated with 100 сс. of а spore suspension of the fungus in
August. Тһе results obtained in a single season at the Central
Station on artificially inoculated soil are decidedly inconsistent,
and it was only where very heavy rates of fertilizer (1200 pounds
per acre) were used that the infection was materially reduced.
Тһе addition of lime at the rate of 4500 pounds per acre had по
effect in reducing wilt.

Тһе arrangement of plots and the fertilizers used at the Poplar-
ville Branch Station are indicated in table xr.

TABLE XI

ARRANGEMENT OF PLOTS AND AMOUNTS OF FERTILIZERS USED ON
COTTON SOIL INFESTED; WITH THE WILT FUNGUS, FUSARIUM
VASINFECTUM. POPLARVILLE BRANCH STATION. PERIOD OF

REATMENTS 5 YEARS

Plot 3 Plot 6 Plot 9 Plot 12
400 Ibs. acid 400 Ibs. acid 400 lbs. acid
phosphate phosphate phosphate
150 Ibs. ammonium} 150 lbs. ammonium} 150 lbs. ammonium} 300 lbs. ammonium
sulphate ulphate

sulphate sulp р
50 lbs. potassium | 100 lbs. potassium | 100 lbs. potassium | 250 lbs. potassium
sulphate sulphate sulphate sulphate

Plot 2 Plot 5 Plot 8 Plot 11
Check Check Check Check

Plot 1 Plot 4 Plot ? Plot 10
400 Ibs. acid 400 Ibs. acid 200 Ibs. acid 400 lbs. “ere
phosphate phosphate phosphate phos
150 lbs. ammonium} 150 lbs. ammonium] 150 lbs. ammonium] 100 lbs. Enn
sulphate sulphate

sulphate sulpha 4
100 Ibs. potassium | 200 Ibs. potassium
sulphate sulphate

Тһе plots at the Poplarville Branch Station were planted about
the middle of April to Mississippi Station Trice cotton, and on
September 2 wilt infection counts made in each plot. Тһе data
obtained are presented in table xir.

[Vor. 14
400 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE XII

ИТГ E COTTON WILT IN FERTILIZER PLOTS ас зз ривая
АМОН EXPERIMENT STATION. SEASON

Plot Seed cotton | Total number Number Per cent
No. per acre, lbs. plants wilted wilted
1 666 493 226 45.84
2 468 622 138 22.19
8 1206 691 102 14.76
4 1278 773 8 1.04
5 810 18 D. 02
6 1080 762 .92
7 1080 660 1/21
8 1 612 17 2.78
9 630 665 1 .15

10 486 613 0 0
11 4 512 1 .19
12 990 590 0 0

The results with the above fertilizer treatments indicate that
applications of potassium may have been beneficial in reducing
the amount of wilt infection. In plot 1, which received adequate
phosphate and nitrogen as ammonium sulphate, but no potas-
sium, the infection was even considerably higher than the un-
treated plots. This is probably explained by the fact that these
plants made a very vigorous growth last season owing to the
liberal amounts of phosphate and nitrogen used, and such appli-
cations may have exhausted the potash reserve in this plot,
resulting in a greater prevalence of the disease. At any rate, the
writer observed the plants in plot 1 early in July and they were
at the time quite large, with wilt beginning to appear over the
entire plot. Photographs of the condition of the plants in plots
1, 4, 5, 6, and 12 were made on September 2, also on October 3,
when they were maturing. These are shown in pls. 37-39. Тһе
spotted areas in plot 1 are especially noticeable, these being due
to plants killed early in the season by the wilt organism. It is
regrettable that there were not available more plots which had
not received potash for a similar period; and until a larger number
of such treatments have been made it cannot be definitely stated
that the disease may be kept in check by such field fertilizer
treatments. А field of Delfos 6102 cotton as shown was photo-
graphed by the writer on August 11, 1926 (pl. 38, fig. 2). This
field had been planted to cotton for several years and had devel-
oped considerable wilt. At the suggestion of the County Agent,

1927]
NEAL—COTTON WILT 401

the grower fertilized the crop with 700 pounds per acre of an
8-4-4 fertilizer and left а small portion of the field untreated as
а check. Wilt infection counts were made in this field in August
and the fertilized portion developed 4.5 per cent wilt, whereas
the per cent of infected plants in the unfertilized portion was
26.8 per cent.

It is hoped that the enlarged program of field experimentation
which has been inaugurated during the present season of 1927 in
Mississippi with reference to the value of fertilizer applications
in reducing wilt damage will yield further results.

Етето Тизтз оғ WILT RESISTANCE

Tests of wilt resistance have been carried forward for a period
of years in cooperation with the Department of Plant Breeding
of the Mississippi Experiment Station. These experiments
were made on soil which had been artificially inoculated with
Fusarium vasinfectum. АП of the varieties received 600 pounds
per acre of an 8-6-4 fertilizer. Тһе total money value per acre of
each variety is estimated at twenty-four dollars for cotton seed
and average lint prices for middling grade of the indicated length.
These lint prices were obtained by averaging quotations of
November 1, received from 5 southern market centers. The
data obtained in these tests on 19 varieties of cotton for the
season of 1926 are summarized in table хиг and were prepared by
the Department of Plant Breeding of the Mississippi Experiment
Station.

The results indicate that the most promising wilt-resistant
staple cottons are Lightning Express, Watson, and Super Seven.
Salsbury, another staple cotton, although not included in the ex-
periment in 1926, is somewhat resistant as indicated by experi-
ments made in former years. Rhyne’s Cook, Dixie Triumph,
Cleveland 54, and Solomon and Oates’ Big Boll are promising
wilt-tolerant, short-staple varieties. It will be observed that
Trice ranked fifth in money value in this experiment. This
variety matures its fruit early and rapidly. Frequently, such
cottons, even though highly susceptible to wilt, are able to
mature а good сгор of bolls, especially when the plants are
heavily fertilized and the season for growth is favorable.

[Vor. 14
402 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE XIII
TESTS OF WILT-RESISTANCE AT MISSISSIPPI EXPERIMENT STATION.
SON 1926

Pounds per acre Lint data Tot d Та
Variety ый Pe Los ollars in
. Length | Cents
cotton Lint cent | inches | per Ip. | Ре acre value
Trice Mississippi Sta. 891 | 272.6 .6| 1% 13.37 | 43.87 5
Half & Half, Mahon 534 | 205.6 .5 7$ .48 | 27.54 19
Cook, Rhyne 762 | 287. .5 % 2.03 | 39.27 9
Cook, 307-6 654 | 228.9 4 7% f .67 | 31.: 16
Toole, Mathis 741 | 263 А Bg f | 12.37 | 38.28 11
oole, Per 753 | 255.3 E 5% 2.03 | 36.68 13
(итеш, Plo. 903 | 293.5 d Bg f | 12.37 | 43.02 6
leveland, Humco 20 657 | 231.9 К 56 2.03 | 33.00 15
Cleveland 54 278.2 .( | 2.70 | 41.‹ 7
uper Seven 888 | 291 S [78 49.4 2
Dixie Triumph 888 | 293.9 к | [ 44. 4
Kelly Big Boll 660 | 236. E f 35. 14
Sol. & Oates Big Boll 531 | 188.0 E! lý f 30. 17
Шег 528 | 178. .8 lý 28. 18
D. & P. L. №. 4 669 | 244. .6 1% | 37.3 12
D. & P. L. №. 5 600 192. M 3% f 3 88. 10
Watson 822 259. a 3% 6.53 49.4 1
Lightning Express 846 | 256.: Ж % | 16.53 | 49.4 3
Delfos 6102 771 | 241.3 | 81. 1% | 14.53 | 41.4 8
SUMMARY

1. Тһе disease of cotton known as wilt has been further studied
and described. An attempt has been made to present a complete
discussion of the literature pertaining to the subject.

2. The disease is widely distributed in the Southern States, and
the annual loss to the cotton crop in the United States is in excess
of 350,000 bales. Such loss varies from year to year, and the
amount of it depends somewhat on environmental factors.

3. Тһе disease causes a yellowing of the leaves of the cotton
plant accompanied by a stunted appearance of the main stems,
usually rather early in the season. Under certain conditions
new outbreaks occur at intervals throughout the growing season.
The characteristic internal symptom, as in the case of other wilt
diseases, is the discoloration of the fibro-vascular tissues of roots,
stems, and occasionally leaf petioles. The fungus responsible
for the occurrence of the disease is Fusarium vasinfectum Atk.
Some of the morphological and cultural characters of the fungus
are given. Тһе fungus is placed in the recognized generic sub-
division “Elegans.” This species produces readily (in culture)

1927]
NEAL—COTTON WILT 403

miero- and macro-conidia and chlamydospores. The latter are
formed in corn-meal or potato-dextrose agar cultures in less than
3 weeks. Тһе color of the stroma is marked and the acid effect
well emphasized. Тһе aromatic odor of the fungus when grown
on rice is pronounced.

5. Physiological studies of the fungus have been made. "These
include growth-temperature relations of the fungus in pure
cultures, hydrogen-ion relations, and a study of the factors in-
volved in producing wilt. Тһе fungus grows slowly at temper-
atures below 10? C., the optimum temperatures being 28-309 C.,
and the maximum temperature about 38? C. It grows well over
а rather wide range of hydrogen-ion concentrations. Тһе best
growth occurred in cultures represented by Py values of 3.0,
4.0, 4.5, and 5.5. Тһе maximum growth occurred in cultures
started at Рн 3.5. The fungus tolerates a culture solution strongly
acid. Тһе growth in the more acid cultures was slow in the
beginning, but later more rapid, reaching a maximum growth
rate almost as high as in the other series initially less acid.
Change in reaction of the culture solution occurred during growth.
Cultures originally acid became less so, while those originally
alkaline became slightly acid, except the ones started on the
extreme alkaline side. As the weight of mat increased, sugar
disappeared from the culture solution.

6. The literature on the factors involved in producing wilt in
the ease of several Fusarium diseases, including cotton wilt, is
reviewed, and experiments relating to toxic action induced by the
cotton wilt fungus, Fusarium vasinfectum, have been performed.
Both filtrates and extracts of the mycelial mat of the fungus
possess toxie properties. 'These toxic substances are not de-
stroyed by boiling, but diluting the filtrate lowers its toxicity.

7. Quantitative analyses for iron oxide, in the tissues of mature,
healthy, and diseased cotton plants, exhibit no correlation
between iron accumulation in the tissues and wilt infection. Тһе
pathogenicity of the fungus has been established under certain
conditions in the greenhouse.

8. The toxicity of aluminium salts to cotton seedlings is not
pronounced, and the susceptibility to invasion by Fusariwm vasin-
fectum was no more evident in plants subjected to different

[Vor. 14
404 ANNALS OF THE MISSOURI BOTANICAL GARDEN

strengths of these salts in a nutrient solution for a certain period
than in those grown in the absence of these salts.

9. Тһе effect of nutrition upon the development of the disease
in the greenhouse does not indicate any consistent lowering or
increase of infection by the various treatments used. Excep-
tions are noted in the case of certain pot experiments with
potassium, calcium, and those receiving additional iron. The
most vigorous plants, as evidenced by their green weight, were
those in which no additional calcium was added. Increasing
the nitrate and phosphate content did not decrease the tendency
of these plants to become infected, as evidenced by lateral root
discoloration, although the vascular discoloration of the stem
revealed by these plants was low. Infection was highest in the
series of pots with added iron, but, as previously indicated,
quantitative analyses of diseased and healthy tissues do not
indicate a correlation between iron oxide content of the tissues
and a tendency to become infected when inoculated.

10. Both greenhouse and field experiments have yielded results
indicating that potassium salts may be beneficial in reducing wilt
damage. Such fertilization methods appear to bring about
delayed infection and the plant is able to mature a portion of its
bolls, especially if such a system is combined with consistent
rotation of crops so as to maintain the organic content of the soil.

11. A study of susceptible and wilt-resistant varieties of cotton
has been made, and the behavior of these varieties on soil arti-
ficially inoculated with the fungus is described. It is suggested
that preference be given to wilt-resistant cottons in tracts where
the soil is heavily infested with the fungus. As a further measure
of control of the disease, liberal fertilization of the crop is impor-
tant and preference should be given to fertilizer formulas con-
taining from 8 to 10 per cent phosphoric acid, 4 to 5 per cent
potash, and from 4 to 6 per cent nitrogen.

ACKNOWLEDGMENTS

In conclusion, the writer wishes to express his grateful appreci-
ation to Dr. B. M. Duggar, under whose guidance the laboratory
and greenhouse aspects of the problem have proceeded, and to
thank him for the many suggestions and helpful criticisms which

1927]
NEAL—COTTON WILT 405

he has given. He is also indebted to Dr. W. F. Hand, of the
Department of Chemistry of the Mississippi A. & M. College,
for the services he has rendered on many occasions as to certain
chemical phases of the work. Acknowledgments are also due
the National Fertilizer Association, who, through their appro-
priation of funds, made this investigation possible; to Mr. H. H.
Wedgworth, of the Mississippi State Plant Board, who kindly
made certain of the photographs and recorded wilt data at the
Poplarville Branch Station; and to Dr. George T. Moore, for the
privileges and facilities of the Missouri Botanical Garden.

BIBLIOGRAPHY

Ajrekar, S. L., and Bal, D. V. (21). Observations on the wilt of cotton in the
ntral Provinces. Agr. Jour. India 16: 598-617. 1921.
Atkinson, С. Е. (92). Some diseases of cotton. Ala. Agr. Exp. Sta., Bul. 41:
9-29.

Beal, J. М. (26). Plant disease reporter. U.S. Dept. Agr. Bur. РІ. Ind. Suppl.

5: 52. I9

Bisby, С. В. (719). Studies on Fusarium diseases of potatoes я truck crops іп
Minnesota. Minn. Agr. Exp. Sta., Bul. 181: 5-47. f. 1-30. 1919.

pre Е. W. (19). Banana wilt. Phytopath. 9: 339-389. pl. 21-34, f. 1-5.

W. Wilt of the pigeon pea. India Dept. Agr. Mem. Bot. Ser. 2:‏ ا
.1-60

——— Fd The wilt jagen of cotton and sesamum in India. Арт. Jour.
e 208-273. f.1 1926.

Chupp, C. (25). Manual of КЕСТІ garden diseases. 647 pp. Мем York, 1925.

Clark, W. M., and Lubs, Н. А. (17). Тһе colorimetric determination of че сам
ion concentration and its applications in bacteriology. Jour. Bact. 2: 1-34,
109-136, 191-236. f. 1-8. 1917.

Nee E. E. (23). The relation of temperature to the Fusarium wilt of the

ato. Am. Jour. Bot. 10: 71-88. pl. 8-11. 1923.

us J. F. (24). A preliminary account of ihe investigation of cotton wilt
in Central Provinces and Berar. Agr. Jour. India 19: 251-260. 1924.

Duggar, B. M. (09). Fungous diseases of plants. 508 рр. New York, 1909.

Edgerton, C. W. (20). Tomato wilt. La. Agr. Exp. Sta., Bul. 174: 3-55. f. 1-19.

1920.

Elliott, J. >; г Arkansas cotton diseases. Ark. Agr. Exp. Sta., Bul. 173: 3-26.
pl. 1

Fahmy, T. Perg D production by Fusarium Solani of a toxic excretory substance

capable of causing wilting in plants. Phytopath. 13: 543-550. 1923.

Fulton. Н. В. (07). Cotton wilt. La. Agr. Exp. Sta., Bul. 96: 5-15. f. 2. 1907.

Gilman, J. C. (16). Cabbage yellows and the ее of temperature to its occur-
rence. Ann. Mo. Bot. Gard. 3: 25-81. pl. 1-2, f. 1-16. 1916

Goss, R. W. (24). Potato wilt and stem-end rot caused by Fusarium eumartii.
Nebr. Agr. Exp. Sta., Bul. 27: 1-83. рі. 1-9, f. 1-4. 1924

[Vor. 14
406 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Hartwell, B. L., and Pember, Е. В. (18). Тһе presence of aluminium as а reason
for is фен іп the effect of so-called acid soil on barley and “ш боп
Sci. 6: 259-279. f. 1-2. 1918.
Haskell, К. J. (19). Fusarium wilt of potato in the Hudson River Valley, New
ork. Phytopath. 9: 225-260. рі. 13-15. 1919.
Higgins В. В. (09). Is Neocosmospora vasinfectum (Atk.) Smith, the perithecial
age of the Fusarium which causes cowpea wilt? N. C. Agr. Exp. Sta., Ann.
Rept, 32: 100-116. /. 1-16. 1909.
Hoffer, С. N., and Carr, В. Н. (23). Accumulation of aluminium and iron com-
pounds in corn plants and its probable relation to rootrots. Jour. Agr. Res.
23: 601-823. рі. 1-21. 1923. р
, апа Trost, Ј. Е. (723). Тһе accumulation of iron and aluminium com-
pounds in corn plants and its probable relation to root rots. II. Am. Soc.
Agron., Jour. 15: 323-331. я
Hopkins, Е. Е. (21). Growth and germination of Gibberella saubinetti at varying
hydrogen-ion concentrations. Phytopath. 11: 36. 1921
М2 С. M. (13). Rangpur tobacco wilt. India Debt. Agr., Mem. Bact.
: 67-83. 1913.
Rea L R., and Tisdale, W. B. (722). Тһе influence of soil — upon the
бетоны of Вах wilt. Phytopath. 12: 409-413. f. 1. 19
Kirby, R. S. (22). Тһе take-all disease of cereals and other i gaans. Ibid. 68-88.
l. 2-4, f. 1-3. 1922.
Lehman, S. G. (723). Pod and stem blight of soy bean. Ann. Mo. Bot. Gard. 10:
111-178. pl. 9-18, f. 1-13. 1923.
Lewis, А. C. (11). Wilt disease of cotton in Georgia and its control. Ga. State
Bd. Ent., Bul. 34: 1-31. 1911.
Lint, Н. С. (14). Report of potato scab experiments. М. J. Agr. Exp. Sta.,
Ann. Rept. 35: 477-488. А
‚ C15). Ibid. 36: 618-625. 1915.
—— — — ——, (16). Ibid. 37: 615-625. 1916.
MacInnes, J. (22). The growth of the hee scab organism in relation to hydrogen-
ion concentration. Phytopath. 12: .1. 1922.
Martin, W. H. (20). Тһе relation of pe to soil acidity and to the control of
potato scab. Soil Sci. 9: 393-408. /. 1-2. 1920.
a | A comparison of inoculated and pecu — for the

кайне У. T. (25). Тһе influence of aluminium, ias ese and iro
pon the growth of sugar cane, and their relations to the infertility of acid island
Hawaiian Sugar Planters’ Assoc. Bul. 49: 1-95. f. 1-33. 1925.
Pg W. A. (00). The wilt disease of cotton. d S. Dept. Agr., Div. Veg.
Physiol. and Path. Bul. 27: 1-16. pl. 1-4, f. 1
— ————, (08). Cotton wilt. U.S. Dept. Agr., . Bul. 333:1-24. f.1-
I 1908.
, and Gilbert, W. W. (712). The control of cotton wilt and root knot.
8. е Agr., Bur. РІ. Ind. Сис. 92: 3-19. f. 1-12. 1912.
Picado, C. (23). Sur l'action à distance des champignone phytopathogenes.
Cong. Path. Veg. Strasbourg, pp. 28-34. 1923.
Rast, L. E. (722). Control of cotton wilt by the use of potash fertilizer. Am. Soc.
Agron., Jour. 14: 222-224. f. 1-3. 1922.

1927]
NEAL—COTTON WILT 407

Rosen, H. R. (26). Efforts to determine the means by which the cotton wilt fungus,
Fusarium vasinfectum, induces wilting. Jour. Agr. Res. 33: 1143-1162. f. 1-3.
1926.

Shaffer, P. A., and Hartman, A. F. (21). ew iodometric determination of copper
1,

and its use in sugar analysis. Jour. Biol. Chem. 45: 349-390.
Shive, J. W. (16). А study m vltesiolegeul balance in nutrient media. Physiol.
es. 1: 327-397.

ut E. F. (99). Wilt divin of cotton, watermelon, and cowpea. U.S. Dept.
„ Div. Veg. Physiol. and Path. Bul. 17: 1-53. ot 1-10. 1899.
Р F. Г. (21). Diseases of economic plants. 507 pp. Rev: ed. New York,
1921

"Tisdale, W. В. (23). The influence of soil temperature and soil moisture upon the
Fusarium disease in cabbage seedlings. Jour. Agr. Res. 24: 55-86. pl. 1-2, f.
1-10. 1923.

Webb, В. W. (19) Studies in the physiology of the fungi. X. Germination of the
spores of certain fungi in relation to hydrogen ion concentration. Ann. Mo.
Bot. Gard. 6: 201-222. 9.

——— — ——, (21). Ibid. XV. Ibid. 8: 283-341. f. 1-89. 1921.

Wollenweber, Н. W., et al. (25). Fundamentals for taxonomic studies of Fusarium.
Jour. Agr. Res. 30: 833-843. f.1. 1925.

Young, H. C., and Bennett, C. W. (21). Studies in parasitism. 1.--Тохіс substances
оды by fungi. Mich. Acad. Sci., Ann. Кері. 22: 205-208. 1921.

[Vor. 14, 1927]
408 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 31

Fig. 1. A cotton plant infected with the wilt fungus, Fusarium vasinfectum.
Note that the main stem is short and stunted and the lower branch has grown
normally.

Fig. 2. Stems of mature cotton plants sectioned to show the symptoms of the
disease in the vascular elements. At left, three sections of stems from a diseased

plant. Note the brown to black irit of the fibro-vascular tissues. At
right, three sections from a healthy p

ig Photomicrograph of Fusari rium vasinfectum taken from a 12-day-old
culture of the fungus growing on potato-dextrose agar showing terminal and inter-
calary formation of chlamydospores. X 1

Fig. Photomicrograph of Fusarium vasinfochum taken from a 10-day-old
culture of the fungus growing on potato-dextrose agar showing micro- and macro-
conidia. Х 100.

DTM NOLLOO— IVAN

№ ка диг е а = nd ч р rote) PHIL

ANN. Mo. Bor. Ganp., Vor. 14, 1927 Prate 31

(Уот. 14, 19:7.
410 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 32

Fig. 1. Experiments to determine the pathogenicity of cotton wilt, Fusarium
vasinfectum Atk., in the greenhouse. Season

The soil in the above pots was heavily infested with the cotton wilt fungus, and
was obtained from plot 1 in the cotton fertilizer experiments at the South Mississippi
Experiment Station. The soil in the 2 pots at left was sterilized in autoclave for

5 hours prior to sowing seed. The soil in the 2 pots at right was unsterilized. The
seed was planted September 15, 1926, and photographed December 3, 1926. Note
the wilt in the unsterilized pots. Variety Delfos 6102.

Fig. 2. At left, uninoculated control flasks containing Duggar's solution and cot-
ton seedlings. At right, cotton seedlings in flasks containing filtrates from 18-day
cultures of Fusarium vasinfectum grown on pend s solution. Note toxic effect.
АП flasks photographed after treatment of 36 hou

Fig. 3. At left, uninoculated control flasks containing s sterile distilled water and
cotton seedlings. At right, cotton seedlings in IM con A extracts of mycelial
mats from 18-day cultures of Fusarium vasinfectum grown on Duggar’s solution.
Note toxic effect. All flasks photographed after aid "of 36 hours.

E
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PLATE

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NEAL E

-COTTON WILT

[Vor. 14, 1927]
412 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 33

Fig. 1. Delfos 6102 cotton seedlings grown in Shive’s RC; nutrient solution
alone and in this nutrient plus 3.6, 9, 18, and 36 parts per million, respectively, of
aluminium, from aluminium sulphate. Slight toxicity is observed in culture 5.
Growing period 29 days.

Fig. 2. Delfos 6102 cotton seedlings grown іп Shive’s RC; nutrient solution
alone and in this nutrient plus 3.6, 9, 18, and 36 parts per million, respectively, of
aluminium from aluminium chloride. Note toxicity in culture 5 and retarding effect
upon lateral root development. Growing period 29 days

Fig. Delfos 6102 cotton seedlings grown in Shivo’ В.С; nutrient solution
alone and in this nutrient plus 3.6, 9, 18, and 36 parts per million, respectively, of
aluminium, from aluminium chloride for a period of 29 days. All cultures transferred
to Shive’s К.С; nutrient solution on thirtieth growing day. Seedlings in jars at
right of each culture inoculated with Fusarium vasinfectum and photographed one
week after inoculation. Note that seedlings in all inocula ted jars are infected.

ANN. Mo. Вот. Ganp., Vor. 14, 1927 PLATE 33

та,
| МХ
қ ЛА B

my mero
3 $

NEAL—COTTON WILT

ГУ оз. 14. 1927]
414 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 34

Variety of cotton in each series, Delfos 6102. All plants photographed after
growing in inoculated sand cultures and receiving the nutrients indicated for approxi-
mately 7 months.

Fig. 1. Series I.

Fig. 2. Series IT.

Fig. 3. Series III.

Fig. 4. Series IV.

Fig. 5. Series V.

Fig. 6. Series VI.

ANN. Mo. Вот. GARD., Vor. 14, 1927 PLATE 34

NEAL—COTTON WILT

[Vor. 14, 1927)
416 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE
PLATE 35

Variety of cotton in each series, Delfos 6102. АП plants photographed after
growing in inoculated sand cultures and receiving nutrients indicated for approxi-
mately 7 months.

ig. Series VIII.

Fig. 2. Series IX.

Fig. 3. Series X.

Fig. 4. Series XI.

Fig. 5. Series XII.

ANN. Mo. Вот. Garb., Vor.

14,

1927

NEAL—COTTON WILT

]Vor. 14, 1927]
418 ANNALS OF THE MISSOURI BOTANICAL GARDE

EXPLANATION OF PLATE

PLATE 36

Fig Effect of commercial fertilizer upon the development of cotton wilt,
Fusarium vasinfectum, in the greenhouse. Rate of application 800 pounds per acre.
At top, plants uninoculated. At bottom, plants inoculated. Variety, Mississippi
Station Trice. Season 192

Fig. 2. Cotton plants growing in a field in which fertilization has not been fol-
lowed. Note the high per cent of wilt. Photographed September 8, at Mayhew,
Mississippi. Season 1926.

ANN. Mo. Вот. Ganp., Vor. 14, 1997 PLATE 36

NEAL—COTTON WILT

[Vor. 14, 1927]
420 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 37

Fig. 1. Cotton fertilizer experiments at South Mississippi Experiment Station,
Poplarville, Plot 1, in the foreground, fertilized with 400 pounds acid phosphate
апа 150 pounds ammonium sulphate. Plot 4, in the background, fertilized with 400
pounds acid phosphate, 150 pounds ammonium sulphate, and 100 pounds potassium
sulphate. Note the high per cent of wilt in plot 1. Season 1926.

"ig. 2, Cotton fertilizer experiments at South Mississippi Experiment Station,
Poplarville. At right, plot 5, check. At left, plot 6, 400 pounds acid phosphate,
150 pounds ammonium sulphate, and 150 pounds potassium sulphate. Season 1926.

ANN. Мо. Вот. Garb., Vor. 14, 1927 PLATE 97

NEAL—COTTON WILT

[Vor. 14, 1927]
422 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 38
Fig. 1. Cotton fertilizer experiments at South Mississippi Experiment Station,

Poplarville. At left, plot 12, fertilized with 400 pounds acid phosphate, 300 pounds
ammonium sulphate, 250 pounds potassium sulphate. At right, check. Season
1926.

А view of a cotton field near Aberdeen, Mississippi, showing the effect
of а complete fertilizer on wilt control. Тһе rows at left were not fertilized. Тһе
rows at right received 700 pounds per acre of an 84-4 fertilizer. Season 1926!

ANN. Мо. Вот. Garb., Vor. 14, 1927 PLATE 38

NEAL—COTTON WILT

[Уоь. 14, 1927]
424 ` ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION ОЕ PLATE

PLATE 39

Fig. 1. Cotton fertilizer experiments, South Mississippi Experiment Station,
Poplarville. Plot 1, fertilized with 400 pounds acid phosphate and 150 pounds am-
monium sulphate (referred to in plate 37, but photographed one month later when
cotton was mature). Note missing hills and dead plants caused by the wilt fungus.
Season 1920.
ig. 2. Cotton fertilizer experiments, South Mississippi Experiment Station,
Poplarville. Plot 4, referred to in plate 37 but photographed one month later when
cotton was mature. Fertilized with 400 pounds acid phosphate, 150 pounds am-
monium sulphate, and 100 pounds potassium sulphate. Note healthy appearance.
Season 1926.

ANN. Mo. Вот. Ganp., Vor. 14, 1927 PLATE 39

NEAL—COTTON WILT

SPECIES ОЕ CERCOSPORA ОМ SMILAX IN THE
UNITED STATES
L. O. OVERHOLTS
Professor of Botany, Pennsylvania State College
Formerly M ycologist to the Missouri Botanical Garden and Visiting Professor in the
Henry Shaw School of Botany of Washington University

In the identification of a Missouri Cercospora on Smilax leaves
it was found desirable to examine rather critically authentic
material of the different species reported on that host. These
species are as follows: C. Smilacis de Thuem; C. Smilacina Sace.;
C. Petersii (Berk. & Curt.) Atk.; C. mississippiensis Tracy &
Earle; C. subsanguinea Ell. & Ev.; and C. nubilosa Ell. & Ev.
Of these, C. nubilosa can be left out of the subsequent discussion,
since an examination of the type collection in the herbarium of
the Missouri Botanical Garden shows that it is on leaves of
Dioscorea villosa L. and not on тах as originally reported. On
the original label the host is recorded as “Smilax ?," but the
doubt indicated by the question mark was not incorporated in
the record when the species was described. However, compari-
sons of the leaves with those of Dioscorea villosa L. shows con-
clusively that they are from plants of the latter species.

C. subsanguinea has been available only from one collection, on
Maianthemum canadense, distributed in ‘Fungi Columbiana,’
No. 4110. On this host it forms irregular necrotic areas 1 cm.
or more broad and long, not bordered by a distinct margin.
This is in sharp contrast to the type of spot produced by the
other species on Smilax. The fasciculate conidiophores are
abundant on the lower side of the leaf over all the dead area. They
measure 40-240 x 6 u. The spores also are distinctive, being
practically cylindric, 10-celled or more when mature, and measure
45-120 x 5-6 и. If these are the characters of that species
when on Smilax as well, it need not be confused with the species
discussed below.

The other collections examined, numbering in all about thirty,
readily fall into two groups on the basis of size and shape of the
spores. Before describing these in detail it may be well to state
that an examination of this series of collections has demonstrated

ANN. Мо. Вот. Garb., Vor. 14, 1927 (425)

[Vor 14
426 ANNALS OF THE MISSOURI BOTANICAL GARDEN

conclusively that one can easily be misled by failing to recognize
the variability in the shape and coloration of the conidiophores in
young and old specimens. Probably in all species of Cercospora
the conidiophores are at first straight and pale-colored (pl. 41,
fig. 4), i.e., pale fuscous or pale cinnamon. Тһе conidia originate
apically, and one may find an entire collection in which the conid-
iophores and the conidia are in this stage of development. If
the fungus continues growth, however, the apical conidium is
pushed aside as the conidiophore grows distally from a point
near the point of origin of the first conidium. This manner of
growth forms an offset in the conidiophore hypha at that point.
Another conidium is produced on the apex of the new growth, and
it in turn may be pushed aside and another offset results. Asa
result of this method of growth an old conidiophore may become
very irregular at its apex. At times the direction of growth
assumed by the conidiophore after the production of one coni-
dium may be almost at right angles to that of its previous
growth, in consequence of which the mature conidiophore may
present a sharp elbow near its distal end (pl. 41, fig. 7). Like-
wise the color of the conidiophore becomes darker with age, so
that while ір а young condition it may appear rather pale, іп ап
older stage it assumes а darker color, in the species discussed
here becoming dark reddish-brown or chestnut on the lower
part, the apex remaining somewhat paler.

Length of the conidiophores is also a factor on which little reli-
ance can be placed in the genus if the conditions in the species on
Smilax can be taken as a criterion. For example, in a collection
distributed by Nash in ‘Plants of Florida,’ No. 1872, the conidi-
ophores measure 40-75 y long, and essentially the same measure-
ments are obtained from de Thuemen, ‘Мус. Univ.' No. 1670.
However, in Nash, ‘Plants of Florida,’ No. 1893, they measure
45-135 џ and in a collection by Peck at Manor, І. I. (Herb. N. Y.
State Mus.) they measure 72-225 u. Yet in other characters
these three collections are so similar that there can be no doubt
they should be referred to the same species. Likewise a col-
lection by Peck at Wading River, N. Y., gives conidiophores
50-140 y, while one at Arcadia, Mo. (Overholts Herb. 10426)
has them 75-210 u; in ‘Plants of the Gulf States,’ No. 7802, they

1927]
OVERHOLTS—SPECIES ОЕ CERCOSPORA ON SMILAX 427

are 150-180 џ long; and in Nash, ‘Plants of Florida,’ No. 2125,
they аге 105-165 и. Moreover, I have found that mounts from
different spots on the same leaf give about as much variation as
may be exhibited by different collections, so that it has become
quite evident that, in the species оп Smilax at least, little reliance
can be placed on the length of the conidiophores.

After examining all the available collections it is apparent,
as stated above, that they readily fall into two categories on the
shape and size of the spores, so that of the four described species
yet to be considered, two only can remain as valid. Опе of these
is C. Smilacis de Thuemen, іп which the conidia are subcylindric,
tapering very gradually to a narrowed apex, and hence in reality
slightly obelavate when seen on the conidiophores. Frequently
they are considerably curved. They measure for the most part
60-135 x 4-5 u, but some as short as 40-50 џ are to be found,
and perhaps longer ones also could be located. The septations
are most frequently only two or three in number but may vary
to ten or eleven, or perhaps more (see pl. 40).

Тһе other collections must be referred to C. Smilacina. Amer-
ican specimens were first so identified by Saccardo, the material
being the same collection that de Thuemen had apparently re-
ferred for Peck to his own C. Smilacis. Since Peck reported it as
this species (Ann. Rept. N. Y. State Mus. 33: 29. 1880) it is
sometimes cited in synonymy as C. Smilacis Peck. Тһе species
differs from C. Smilacis in the conidia that are usually rather
abruptly narrowed at one end and have a tendency to be snow-
shoe-shaped or the shape of a tailed pumpkin seed. Most of the
conidia measure 40-65 x 5-6 u, but a few are as short as 30 в
and 4-5 y in diameter, while occasionally a spore as much as 75 ш
long has been seen. They may be as much as 8-celled, but usu-
ally show less than that number. In general, therefore, they
are much shorter than those of C. Smilacis and are more sharply
contracted at one end (see pl. 41).

How often it occurs that Cercospora conidia are clavate and not
obclavate on the conidiophores, remains for further study to
determine. Аз originally founded by Fresenius (‘Beitrage zur
Mykologie,’ p. 91, 1863) the genus is based on C. Ари, which he
illustrates as with obclavate conidia, and the usual conception

[Vor. 14
428 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of the genus involves that characteristic. While I have not seen
a large number of clavate spores оп conidiophores in C. Smilacina,
yet they have been observed several times, though always ob-
Aiously immature, and at a later stage of maturity they may
possibly take а more obclavate form.

I find nothing in the character of the spots produced by these
two species to aid in their macroscopic separation. They vary
to а considerable extent on the different hosts, with variation in
thickness and texture of the leaves. Always the spots produced
by both species are subcircular in form and at first dark in color,
the center usually becoming lighter in color with age, leaving a
narrow, dark red or purplish, often raised margin. Yet in some
collections the margin is not well marked. Тһе conidiophores are
usually hypophyllous in both species, but at times are amphigen-
ous, though this tendency to occur on both surfaces seems more
marked in C. Smilacis than in C. Smilacina.

А few words may be said regarding C. Petersii. The species
was originally described as a Helminthosporium (Grevillea 3:
102. 1875), but was transferred to Cercospora by Atkinson
(Jour. Elisha Mitchell Scientif. Soc. 8: 25. 1892). I have ex-
amined mounts from the type collection of this species but failed
to find the conidia and found only a few conidiophores, and was
not able to arrive at any decision as to the exact status of the
species. In the sense of Atkinson (l.c.) and as determined іп
several collections by Ellis it most certainly is C. Smilacina, and
I am of the opinion that it must be regarded as a synonym of
that name. It is recorded in Saccardo’s host index as on the
stems of Smilax, evidently an error, since the original description
specifically locates it on the lower surface of the leaves of Smilax
and of Laurus.

The following diagnoses of the two species most concerned in
this treatise are appended:

Cercospora Smilacis de Thuemen, Myc. Univ. No. 1670. 1880.

РІ. 40.

С. mississippiensis Tracy & Earle, Bull. Torr. Bot. Club 22:
179. 1885.

Spots subcircular, usually limited by the veins, 1-7 mm. in

diameter, at first dark purplish red, then becoming paler at the

1927]
OVERHOLTS—SPECIES OF CERCOSPORA ON SMILAX 429

center while retaining a dark conspicuous margin; conidiophores
fasciculate, hypophyllous or amphigenous, septate, pale cinnamon
to fuscous or chestnut, at first straight and paler at the tips, in
age becoming chestnut color and the tips nodulose or wavy,
45-225 y. long, 3-5 u broad; conidia subcylindric, gradually nar-
rowed at the apex, hence somewhat obclavate, 3-4-celled or
becoming 10- to 14-celled, pale fuscous, 60-135 X 4-5 u.

On living leaves of various species of Smilax.

The species differs from C. Smilacina Sacc. in the longer spores
that are sub-cylindric and not sharply narrowed.

Specimens examined: de Thuemen, Myc. Univ. Nos. 1670
(type) and 1768; Starkville, Miss., 1891, Tracy (type of C. mis-
sissippiensis Tracy & Earle) (Mo. Bot. Gard. Herb. and Herb.
N. Y. State Mus.); Manor, L.I., Peck (Herb. №. Y. State Mus.);
Columbus, Miss., 1895, Tracy (Mo. Bot. Gard. Herb.); Nash,
Plants of Florida, Nos. 1893, 1872, and 2073 (Mo. Bot. Gard.
Herb.); Ellis, N. Am. Fungi, No. 1251 (Mo. Bot. Gard. Herb.);
Bairds, Miss., Tracy (Mo. Bot. Gard. Herb.).

Cercospora Smilacina басс. Michelia 2: 364. 1881. FL AL

? C. Petersii (Berk. & Curt.) Atk. Jour. Elisha Mitchell Sci-
entif. Soc. 8: 25. 1892.

? Helminthosporium Petersii Berk. & Curt. Grevillea 3: 102.
1875.

Spots subcircular, usually limited by the veins, 1.5-5 mm. in
diameter, at first uniform dark red or purplish black, becoming
paler at the center as the spot expands, retaining (usually) a
dark, somewhat raised border; conidiophores hypophyllous or
rarely amphigenous, fasciculate, septate, pale cinnamon to
chestnut below, usually paler above and there quite wavy or
nodulose in extreme age, sometimes bent almost at right angles
near the upper end, 50-215 џ long; conidia terminal and lateral,
rather abruptly narrowed toward one end, pale fuscous to brown-
ish, usually 2- to 4-celled, sometimes 6- to 8-celled, 40-65(-75)
х би.

On living leaves of various species of Smilaz.

'The species is easily separated from C. Smilacis by the conidia
that are rather abruptly narrowed near the middle, and less
than 75 y long.

[Vor. 14, 1927]
430 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined: Wading River, N. Y., 1879, Peck (Herb.
N. Y. State Mus.); Shelter Island, N. Y., Clinton (Herb. N. Y.
State Mus.); Plainville, Conn., 1883, A. B. Seymour (Mo. Bot.
Gard. Herb.); Ravenel, Fungi Am. Nos. 166 and 616 (Mo. Bot.
Gard. Herb.); Hume, Florida Fungi, No. 62 (Mo. Bot. Gard.
Herb. and Herb. N. Y. State Mus.); Tracy, Plants of the Gulf
States, No. 7802, Auburn, Ala., 1897 (Mo. Bot. Gard. Herb.);
Auburn, Ala., 1896, Underwood (Mo. Bot. Gard. Herb.); Nash,
Plants of Florida, No. 2125 (Mo. Bot. Gard. Herb.); Seymour &
Earle, Economie Fungi, No. 199 (Mo. Bot. Gard. Herb.);
Bartholomew, Fungi Col. No. 2808 (Mo. Bot. Gard. Herb.);
Arcadia, Mo., 1926 (Overholts Herb., No. 10426, and Mo. Bot.
Gard. Herb.).

EXPLANATION оғ PLATE
PLATE 40
Cercospora Smilacis

Fig. 1. Conidia from de Thuemen, Myc. Univ. 1768. 400.

Fig.2. Fascicle of conidiophores emerging through a stoma. Ellis, М. Am.
Fungi, 1251. х 160

Fig. 3. Conidiophores, one bearing an obclavate conidium. From de Thuemen,
Myc. Univ. 1768. X 400.

Fig. 4. Conidiophores, one bearing a conidium. From deThuemen, Myc.
Univ. 1670. x 400.

Fig. 5. Conidia. From Ellis, М. Am. Fungi, No. 1251. x 400.

Fig. 6. Conidia. From the type collection of C. mississippiensis. X 400.

Fig. 7. Conidia. From collection at Starkville, Miss., by Tracy, 1892. х 400.

ГУ оз. 14, 1927]
432 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 41
Cercospora Smilacina

Fig. 1. Conidia. From Tracy, Plants of the Gulf States, No. 7802. X 420.

Fig. 2. Conidia. From collection at Arcadia, Mo., 1926, Overholts Herb. 10426.
x 420. |

Fig. 3. Conidia. From Seymour & Earle, Economic Fungi, No. 199. X 420.

Fig. 4. Conidiophores in young condition. From collection at Auburn, Ala.,
1896. х 420.

Fig. 5. Conidia. Same as fig. 4.

Fig. 6. Conidia and two conidiophores. From Bartholomew, Fungi Col. 2808.
x 420.

Fig
Overbólte Herb. 10426.

Fig. 8. Old conidiophore. Same as fig. 7. X 420.

Fig. 9. Old conidiophore. From Seymour & Earle, Economic Fungi, No. 199.
x 420.

Fig. 10. Conidiophore. From Tracy, Plants of the Gulf States, No. 7802.
x 420

Fascicle of conidiophores. From collection at Arcadia, Mo., 1926,
65.

Fig. 11. Two conidiophores and two conidia. From collection at Wading River,
N .Y., Peck, 1879. x 420

GENERAL INDEX TO VOLUME XIV

New scientific e of plants and 0 сп им н pa new combinations -

printed in bold face type; synonyms

ving reference to figur

and plates, in italics; and previously ا‎ радо scene Peu and all other imn

in ordinary ty

A

А. & М. College, ra
wilt experiments а at, 360, 398
Agaricus a 139: aureus, Б
aurive 152; 224 nalis, 134:
mm 125; dactyliotus, 174; aug
161; : flamman 8, 148; hor
i Johnsonianus, 130; luci E А
eons te m ; mutabili is, 138;
, 106; pud us, pA ismod dad
115; togularis, "114, соот, 184
Aluminium compounds, a of cot-
wilt to, 383, oe 395
жй аррив “forifer, 1 , уат. В, 21
Aster Wrightii, 32; oe 28

cotton

B
Bystropogon axillare, 82; minutus, 83;
uniflorus, 82
C

Calcium, sig of, on development of
cotton w
буз вресїев of, оп Smilax іп the

nited States,
Cercospora, 425; Apii, 427; mississip-
piensis, 428; Жаз T Pet d
429 Жайык». 429, 432; Smilacis

8, 430;
Cotton, wilt-resis
Cotton’ wilt, a pathological and physio-

logieal investigatio n, 359; description
of, 364; factors involved in producing,

Cryptantha, section Oreocarya, 237;

affinis, 307; aperta, 295, 204; Bakori,
331, 358; ' Bradburiana, 307, 3

246, var. disticha, 248, "var. laxa, 246,
ANN. Мо. Вот. Garp., Vor. 14, 1927

var. multicaulis, 244, 348, var.
typica, 248; Jon nesiana, 323, 356;
leucophaea, 202, 348; longiflora,
326, 358; Macounii, 303,

mensana, 333, bi modesta, 277,
352; nana, r. commixta, 312,
var. ovina, 314, guns Shantzii, 313,

1 Б пп
gena, 265, 320; 'oblat ta, 254, 348;
Osterhoutii, 829, 328; Palmeri, 253;

30 5
354; stricta, 264, 350; suffruti-
Ж) 4 242; tenuis, 327, 358; thyrsi-
flora, 2 = 352; tumulosa, 216, 8 852;
utahen nsis, 270; virgata, 270, 350;
virginensis, 274, 352; Wetherillii,

324, 358
Cynoglossum glomeratum, 307

D
Dioscorea villosa, 425
E
Epling, Carl C. Studies on South Ameri-
n Labiatae, Ш, 47
Fees florifer,
Eritrichium section Ps eudomyosotis, 237
Eritrichium ссы, 319; glomera-
de 303, 307, var. fu vocaneseens, 278,

ar. hispidissimum, 2 h umile,

var. one n, 270; Jam mesi,
m, 262; Sault coude, 244;

setosissimum, 268; vir "gatum, 70

F

Fertilizers, commercial, effect of, on
development of cotton wilt, 39

Flammula ЕЕ folia, 148; unicolor,
136; M эң 148

Fungicides а in cotton wilt ехрегі-
ments,

(433)

434

Fusarium vasinfectum, 360; mor ры

logical and cultural characters of, 3
pathogenicity of, 382; et on r і]
А of, 366

с

Gardoquia, 47; acutifolia, а 4
ns by viflora - chilen 56; dis
, 02; ell iptic 54; ne а. 72;
foliolosa, бі; Gillis, 56; glabra ta, 66;
incana, 64; Jan sonit, 891 crophylla,
70; multi, ‘flora "53: obov don revoluta,
74; rugosa, 76; мк 84; sericea,
73; stri rata, 68; "ta axifolia, 67; thymoides,
61: ‚ tomentosa, '60

Hartwell & Pember's nutrient solution,

eloma hortense, 105, 107
иот Petersii, 429
Hesperothymus of Satureia, 50
Hydrogen-ion concentration, effect of,

on germination of fungi,
Hypodendrum oregonense ‚ 140

I

Iron compounds, relation of, to cotton
wilt, 383, 395

Krynitzkia «ids oe оен? и
fe ar.

ulvocanescen idahoensis,
an glomerata, 307, ae acuta, 32 4°
var. virginens 274; amesti, 248;
wii қ 269, ta, 256;
mensana, 338; и" var. abor-
a, 250, var. setosa, 244; oblata, 254;

ust data, 307; sericea,
cated, 268; virgata,

237,

316;

Krynitzkia, section Plerygium,
section Pseudokrynitzkia, 237

L

E, Studies on South American,
ПІ,

и. "Ты her Г. А revision of the
ge nus Коли. 1

М

Maianthemum canadense, 425
gig б, әгі boliviana, 80; bonarien-
8, 84; Darwinii, 78; eugen nioides, 83;
Gilliesti, 83; АШЫ) ena, 76; past illa, 79;
pulchella,

[Vor. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Micromeria section Hesperothymus, 50
be" сад conditions, effect of, on cotton

мын 6 the genus Pholiota in the
United Sta

Monogra P of pS section Oreocarya of
Cryptan

Myosotis pene 307 ; leucophaea, 262;
suffruticosa, 242, 248

N

Neal, David С. Cotton wilt: a patho-
logical and ина investiga-
tion, 359

Neocosmospora vasinfecta, 360

Nitrogen, effect of, = _ development of
cotton wilt, 393,

Nutrition, effect of, oe the develop-
ment of cotton wilt, 389

O

erroe of сурет а monograph
ines sectio

ya, жы; орот, аа ; affinis, 307,
dn nis, 288; alata, 256; aperta,
295; argentea, 287; Baker i, 331; brevi-
flora ; caespitos а, 316;
ойгоо, 299, 301; Kos Vue 298;
cinerea, 246; com ommixta, 312; confe rti-
a, 256; cri cristata, маг depressa, 271;
disticha, 248; dolosa, 313; dura, 283;
E" oodae ‚ 334; cohinoides 278, 321;
285; 'eulophus, 1; flava ‚ 259;

Fro ud 334, var. eire
fulvocanescens 8,

307;

milis, 278; in зона
273. ие 296; дерде Жең 323;
Lemmoni, 246; leucophaea, 262,
йн, 256; Ыт 326;

acbri йй,

815; mitida, 81 ubigena, 265;
oblata, 255; Osterhoutii, 329; Palmeri,
253; paradoxa, 330; Paysonti, 255;
perennis, 307; procera, 287;
propria, 3 pustulosa, 252; sa,
295; salmonensi 263; sericea, 287,
301; setosissima, 268; Shanizii, 313;
Sheldonii, 301; Shockleyi, 334; spicata
270; spiculifera, 298; stricta, 264;
suffruticosa, 248, var. abortiva, 250,

inerea, 246, multicaulis, 244;

urticacea, ; virgata, 270
ma spicata, 270; virginensis, 274;
оппа арі 324

1927]

INDEX

yes L. O. A monograph of the
s Pholiota a in the United States,
87; Speci cies of Cercospora on Smilax in

the United States, 425

F

Payson E. monograph of the

section Oreocarya of кай», 211

ve a eae of the genus, in
he United Sta 8,

Pholiota, 99 гоча 108, 182; adi-
posa, 154, 202, 204; reeling 139;
Aegerita var. strobiloidea, 140 ; aerugi-

nosa, 147,

174; albivelata, ; albocrenulata,
150, 200; angustipes, 166, 208; anoma-
la, 127; append a, 174; aurea, 121,
186; aurivella, Я 0; &urivel-
loides, 151; autumnalis, 134, 174;
blattaria, 113; caperata, 123, 188;
cerasina, K omosa, 169, 74; con-
agosa, 160, 208; curvipes, 164;
dactyliota, 174: destra s, 168; eter-

sibilis, 161, 174; olo
dura, 105, 174, уаг. zanthophylla, 105
aba уму 129; 'erebia, 118, 184; e

ari 6; flam ааа; "148,
o-squam 67, ; fur-
‚ 136; het оода, 169, i74; hormo-

ucifera lutea,
АТР 145; nina. 175; margi-
marginelin, -

cula, 142; rugosa 2

; Schraderi, 159; speciosa, 175;
spectabilis, 144, 96; sphalero-
morpha, 109, 175; squarrosa, 157,
204; squarrosoides, , 206; sub-
nigra, 120; subsquarrosa, 151, 156, 176;
temnophylla, 112; ; 1201
terrigena, 127, 176; togularis, 114, 192;
togularis var. filaris, 11 зрога,
125; tuberculosa, 164, 190; unicolor,
34; ventricosa, , 176; vermiflua,

> А 78; ^ e 170; washingtonensis,

19,

of, on development of‏ ا
cotton wilt, 392, 3 395‏

Poplarville Experiment Station, Missis-
t experiments at,

of, on development of
cotton wilt, 397

Potassium, effect of, on development of
cotton wilt, 392, 395

435

R

Revision of the genus Townsendia, 1
izoa, 47; ovatifolia, 5
Rochelia glomerata, 307

5

Satureia, synopsis of the genus, 47
Satureia NDS 63; ei

; eugenioides, 83; Ў
foliolosa, 61; Gilliesii, 56; Gilliesii, 83;
6; guamaniensis, 58;

striata, bi ta К 67; thymoides,
P tomentosa, 60; vàna, 79; Webe
au

Sm Be ony А4 сез Ж еке on, іп the
United States, 425

Stachys speciosa, 54

Ste ken er, 19, var. В, 2

Stropharia Howeana, 110; Schr PR 159

Stu ie on South American Labiata ae,
ІП, 47

T

Temperature, influence of, on cotton
wilt, 366

revision y the ии 1

Townsendia, 8; alpigena, 17; alpina, 22;
ambigua, 21; arizonica, 27; zr a
X incana, 27; Bakeri, 16; а ta
22; dejecta, 17; eximia, 1

р

, ; 0
formosa, 11, 3
glabella, 16; grandiflora, 13; incana,
25; incana var. ambigua,
proliza, 26; enn 20; lepotes, "o8;
leptotes ‚ 27; mensana, 30; mexicana,
ies. montana, 17; go rryi, 9; Parry

r. alpina, 2; pinetorum, 11; Roth-

[Vor. 14, 1927]

436 ANNALS OF THE MISSOURI BOTANICAL GARDEN
rockii, 16; морем 20; scapigera Toxic substances in cotton wilt fungus,
var. ambigua, 21, caulescens, 21; 380
sericea, 30, 44, 46; serian - „45 46,
var. leptotes, 27, В 29; Ж.

gos E 40;
strigosa, 18, 19; texensis, 15, 38; Хепорота, 47, 76; bolivianum, 80, var.
Vreelandii atsoni, 18; Wil- tarijense, 81; eugenioides, 88
coxiana, 95; Wrightii, 32

JUN © 1977

Volume XIV

| Лобин

of the |
Missouri Botanical

Garden

‚ Contents
February, 1927
А Revision of Ps Genus Townsendia

еле? ае 92- ө ¢ 9 9. 9.949

: Esther L. Lorsen
Studies on South American Labiatae. aH. d ud Carl C. Epling

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